Proceedings of the International Workshop on

Red Coral Science, Management, and Trade: 

Lessons from the Mediterranean

September 23 - 26, 2009 

Naples, Italy

Edited by

Ezio Bussoletti, David Cottingham, Andrew Bruckner, 

Glynnis Roberts, and Roberto Sandulli

U.S. Department of Commerce

National Oceanic and Atmospheric Administration 

National Marine Fisheries Service 

NOAA Technical Memorandum CRCP-13

February 2010
This publication should be cited as: 

Bussoletti, E., D. Cottingham, A. Bruckner, G. Roberts, and R. Sandulli (editors). 2010. Proceedings 

of the International Workshop on Red Coral Science, Management, and Trade: Lessons from the 

Mediterranean. NOAA Technical Memorandum CRCP-13, Silver Spring, MD 233 pp. 

Significant support for the development of this document was provided by NOAA Fisheries, Office 

of Habitat Conservation, and NOAA’s Coral Reef Conservation Program. The views expressed in 

this document are those of the authors and the participants of the workshop, and do not necessarily 

reflect the official views or policies of the U.S. Government, NOAA or DOC.

Additional copies of this publication may be requested from: 

Nerie Canasa

National Oceanic and Atmospheric Administration

NOAA Fisheries 

Office of Habitat Conservation 

1315 East West Highway 

Silver Spring, MD 20910-3282 

Nerie.Canasa@noaa.gov
Proceedings of the International 

Workshop on Red Coral Science, 

Management, and Trade: Lessons from 

the Mediterranean

Edited by Ezio Bussoletti, David Cottingham, Andrew Bruckner, 

Glynnis Roberts, and Roberto Sandulli 

National Oceanic and Atmospheric Administration 

Jane Lubchenco, Ph.D

Under Secretary for Oceans and Atmosphere 

National Marine Fisheries Service 

Jim Balsiger, Ph.D

Assistant Administrator for Fisheries (acting)

Office of Habitat Conservation

Habitat Protection

NOAA Fisheries

Coral Reef Conservation Program

and I.M. Systems Group, Inc. at NOAA 

1315 East-West Highway 

Silver Spring, MD 20910-3282 

U.S. Department of Commerce 

Gary Locke, Secretary 

NOAA Technical Memorandum CRCP-13
Workshop Coordinators:

Ezio Bussoletti, Italy 

David Cottingham, U.S.A.

Proceedings of the International 

Workshop on Red Coral Science, 

Management, and Trade: Lessons from 

the Mediterranean

Sponsors:

Italian Ministry of Foreign Affairs 

(MAE)

Italian Ministry of Environment, Land, and Sea 

(MATTM)

University of Naples “Parthenope”

U.S. Department of Commerce

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Coral Reef Conservation Program

National Oceanic and Atmospheric Administration

The Ocean Foundation

Università degli Studi di Napoli

“Parthenope”
Acknowledgements

The Workshop Coordinators wish to thank:

•	 The Italian Ministry of Environment, Land and Sea (MATTM) for the initial incentive to organize 

this Workshop and for its economic, professional and technical support; 

•	 The Italian Ministry of Foreign Affairs (MAE) for its institutional support;

•	 Prof. Gennaro Ferrara, Rector of the University of Naples “Parthenope,” who has provided the 

access to the University structures, and for his introductory speech;

•	 Dr. Aldo Cosentino, Director General for Nature Protection at MATTM, for his strategic 

coordination, his continuous stimulus to operate, and for his introductory speech;

•	 Dr. Raffaele Celentano, on behalf of the Italian Under-Secretary of State at MAE Hon. Vincenzo 

Scotti, for the introductory speech;

•	 Prof. Raffaele Santamaria, Dean of the Faculty of Science and Technology - University of Naples 

“Parthenope,” for having made the University facilities available during the Workshop, and for his 

introductory speech;

•	 The Local Organizing Committee, for its continuous effort in the organization of the event which 

made it a success;

•	 The complete logistic structure of the Centro Direzionale building of University of Naples 

“Parthenope,” for the technical support;

•	 U.S. Fish and Wildlife Service for its institutional support and subject matter expertise;

•	 U.S. National Oceanic and Atmospheric Administration’s Coral Reef Conservation Program for 

providing financial support for supporting many of the international travelers.

Particular thanks go to Dr. Antonio Maturani, Head of Division in the Nature Protection Directorate 

at MATTM, who, with enthusiasm and competence, supported the preparatory work of the whole 

event.

Very appreciative thanks are due to ASSOCORAL, the National Association of Coral and Cameo 

producers, for hosting an informative visit to Torre del Greco - Naples, and to its President, Dr. 

Ciro Condito, to its past President, Dr. Mauro Ascione, and to Dr. Antonino De Simone, President 

of the Banca di Credito Popolare of Torre del Greco. Workshop participants observed red coral 

processing and had a very pleasant evening party.

Last but not least, a particular thanks is due to all the scientific contributors and managers who 

assured the Workshop full success in the respect of competing different positions. 

The Workshop Coordinators

David Cottingham Ezio Bussoletti

v
Preface

vi
STATEMENT OF ALDO COSENTINO

General Director, Protezione della Natura

MATTM

Following the commitment taken in occasion of the last COP, the Italian Government has organized 

this International Workshop on the Mediterranean Red Coral (Corallium rubrum) jointly with 

U.S. Government. Aim of this event, which followed that held in Hong Kong in March 16th-20th 

2009, was to define a management strategy for the Mediterranean Red Coral analyzing the various 

problems of conservation and the most suitable methods of sustainable harvest.

Despite different positions on the subject, we believe that everybody shares the wish to implement 

a process able to improve the relationships between “science” and “political decisions” in order to 

use them for a better support, in a qualified and not a prioristic approach.

This is one of the major rendez-vous that the international community concerned in biodiversity 

protection will face in 2010; in this sense UNEP is operating helping States to better define the 

mechanisms allowing the scientific results to be promptly available and fully understood by 

“political stakeholders” so that they could be able to identify strategies and initiatives responding 

to the actual needs for a correct survival and development of our planet. 

When we talk of “conservation” and “sustainable use” of biodiversity, it is important to be aware 

that we operate in a peculiar situation in which science, integrated with international legislation, can 

make the difference between an efficient and a not efficient choice. On purpose I state “efficient” 

and not right or wrong.

In fact, when we talk about the genus Corallium, as well as other marine organisms, once the 

biologists have correctly identified the species distribution, they have to be able to understand all 

the parameters necessary to guarantee their sustainable utilization. The question arising at this 

stage is then: which is the best international instrument capable to regulate and control that any use 

is performed following the due rules?

In the case of Corallium rubrum, as well as other marine species, international trade represents an 

important element of its management; however, we should evaluate how reasonable and justified is 

to consider the control of trade as the best choice rather than the control of more critical parameters. 

We believe that we need to understand the difference between the various available instruments 

in order to use those which appear the most suitable, case by case, to get the maximum efficiency.

Instruments of “knowledge” are the IUCN “red lists” which, however, pertain to a global level. If 

a species is globally threatened, this does not mean necessarily that it is so in every area where it 

is present. Then, red lists are an important instrument which however needs to be re-applied and 

tested in the field to get the best and more realistic results.

The Washington Convention is another fundamental instrument of safeguard; we all know, 

however, that it became really operational and useful only when countries felt committed to it, 

deciding its application in the due form.

vii
This Workshop represents in my view an important step ahead in the knowledge of the Mediterranean 

Red Coral as it has been possible to compare the experience of different and very qualified scientists 

representing the leading front on the subject. We have also compared this knowledge with that of 

experts operating in areas other than the Mediterranean Sea. This has allowed filling knowledge 

gaps of scientists who did not worked in this region and, then, did not share the day by day history 

of care and harvest of the Mediterranean precious coral in the last 150 years as it occurred in Italy.

We had hoped, taking advantage of the Workshop, to arrive to a shared, agreed position on a 

strategy concerning the care, safeguard and harvesting of this species on the ground of the positive 

Italian experience on the subject as a result of an activity existing for one and a half century and 

which sustains the economy of a sector of our country where about 15,000 persons are involved.

Concerning the scientific results and the further needs of knowledge, the Workshop was a clear 

success as all the attendants arrived to a shared position agreed unanimously and is reported in the 

Appendix of the present publication. Important statements are done on future actions devoted to 

deepen the research filling the present gaps. Indications on sustainable management measures and 

enforcement in the Mediterranean are also given.

Unfortunately, attendants were not able to arrive to a similar shared position on trade related 

issues. While some participants stressed the importance of an inclusion in CITES Appendix-II for 

Mediterranean Red Coral according to the U.S. official position, others, according to the Italian 

official position, disagreed, mainly because of serious economic and bureaucratic implications of 

a listing, as the source appears not in danger and is managed in a sustainable way. This document 

reports also the report of the trade group.

Despite a full consensus was not reached, we consider that the Workshop has been a success as 

put together experts and experiences coming from different parts of the world allowing people to 

discuss, compete, agree or friendly disagree and always in a very open minded atmosphere. 

In conclusion, this event, as well as the Hong Kong workshop, represents an important step ahead, 

providing a useful contribution to the general problem: the journey has started and we all hope to 

reach some useful results enabling us to provide significant solutions to policy-makers in the near 

future.

 Aldo Cosentino

 

viii
STATEMENT OF JAMES BALSIGER

Acting Director, National Marine Fisheries Service

National Oceanic and Atmospheric Administration

The National Oceanic and Atmospheric Administration’s Coral Reef Conservation Program was 

pleased to co-sponsor the International Workshop on Mediterranean Red Coral (Corallium rubrum) 

in Naples in September 2009. The workshop brought together leading scientists and managers 

from the Mediterranean and around the world to discuss the status of Corallium rubrum and its 

management. The presentations highlighted the most current information available on the status of 

Mediterranean populations of Corallium rubrum as well as the importance the trade plays in local 

economies and traditions. This workshop and a similar workshop on red and pink coral trade held 

in Hong Kong in March 2009 enabled scientists, resource managers, and diplomats to learn about 

these precious corals. Improving our understanding of coral biology and how to manage it is a key 

to its long-term sustainable use.

The spirited discussions during the workshop lead to common understandings and broad acceptance 

of the current state of Mediterranean red coral resources and the knowledge gaps that the scientific 

community needs to address. When the research and management plans called for by working 

group participants are completed, they should lead to greatly improve management throughout the 

range of the species. Representatives of governments from many Mediterranean countries spoke 

of their commitment to preparing the plans, conducting the research, and managing the stocks 

better.

Some of the most contentious discussions at the meeting focused on the appropriateness of 

additional measures to monitor and regulate red coral trade. Some participants expressed their 

view that each country should adopt its own management regulations without involvement of 

international organizations. Because of a lack of meaningful coral management in many countries, 

the United States representatives and others felt that international oversight of national programs 

could provide an impetus for better national management. From that perspective, a listing on 

Appendix II of the Convention on International Trade in Fauna and Flora (CITES) would build on 

national management by ensuring that all exporting countries implemented sustainable management 

strategies. While it is regrettable that participants at the workshop did not agree on the role of red 

coral trade and management, the information presented at the Workshop laid the basis for future 

productive discussions on the science and management of Corallium rubrum. 

The United States would welcome opportunities to participate in those discussions on research and 

management in hopes that all nations develop, adopt, and implement effective measures to better 

understand and manage these precious resources.

ix
x
Table of Contents

Acknowledgments .......................................................................................................................... v

Preface .......................................................................................................................................... vi

Executive Summary ....................................................................................................................... 1

White Papers and/or Abstracts ....................................................................................................... 4

Section I 

Mediterranean Red Coral (Corallium rubrum): Historical Assessment ............................ 5

Section II

Harvest, Manufacturing, and Trade ............................................................................... 100

Section III

Science: Biology and Ecology of Red Coral ................................................................. 139

Section IV

Management .................................................................................................................. 183

Working Group Reports ............................................................................................................ 208

Science/Management .................................................................................................... 209

Trade ............................................................................................................................. 211

Appendix .................................................................................................................................... 214

Participants and Affiliations ........................................................................................... 215

Workshop Agenda .......................................................................................................... 225

Workshop Coordinators: Ezio Bussoletti and David Cottingham ................................ 232

Steering Committee ...................................................................................................... 233

xi
Executive Summary

The Italian Ministry of Agriculture and Ministry of Environment and the U.S. National Oceanic 

and Atmospheric Administration convened the International Workshop on Red Coral Science, 

Management, and Trade: Lessons from the Mediterranean in Naples Italy in September 2009. 

Scientists, managers, representatives of the coral fishery and manufacturing industries, policy 

makers, and environmental organizations from Europe, Africa, Asia, and North America attended 

the workshop. The workshop provided an opportunity to discuss the best available science on the 

natural history of Mediterranean red coral (Corallium rubrum L.) and how it is managed throughout 

the region and utilized around the world.

The workshop was the result of deliberations on a 2007 proposal by the United States to list red 

and pink corals in Appendix II of the Convention on International Trade in Endangered Species 

of Wild Fauna and Flora (CITES). CITES member nations did not adopt the U.S. proposal in 

2007. While the international community recognized the conservation needs of these species, 

questions were raised about species distribution and biology, domestic management practices, 

product identification, and other issues related to implementation of a CITES Appendix-II listing. 

The United States held a workshop in Hong Kong in March 2009 to address these issues, with 

an emphasis on Pacific species. The Naples workshop focused on the Mediterranean red coral, 

Corallium rubrum. 

Corallium rubrum is widely distributed throughout the Mediterranean Sea and into the neighboring 

Atlantic Ocean from Portugal to Cabo Verde. It is slow-growing and inhabits a wide depth range 

from 15 to 600 meters, and possibly even deeper. In shallow waters (between about 15 and 70 

meters), C. rubrum colonies grow in caves, crevices, overhangs and other protected interstices 

in dense patches with as many as 127-1300 colonies per square meter, the majority of which are 

less than 5 cm tall, with dense patches confined to small areas with large expanses of uncolonized 

habitat. In deeper waters, between 70 and 130 meters, coral colonies tend to be larger and more 

dispersed, and typically settle on open surfaces. It is rare to find colonies that reach their historical 

maximum size of 30-50 cm, except in areas where fisheries have never operated. Below 130 

meters, colonies tend to be even larger and less dense. 

C. rubrum has been harvested for more than two thousand years. As early as the 1800s, fisherman 

from Italy, Spain, and France sailed to North Africa in search of new stocks as local areas were 

depleted. Traditional harvest techniques included free divers and non-selective coral dredges such 

as the famed St. Andrew’s cross and the “barra italiana,” both of which harvesters dragged across 

the sea floor to ensnare broken colonies of coral. The invention of SCUBA in the 1950s allowed 

fishermen to harvest red coral more selectively and efficiently. It also lead to over-exploitation of 

coral beds within 15-50 meter depth, especially in areas that were inaccessible to coral dredges. 

As SCUBA technology improved, divers began to harvest down to depths of 100-130 meters. The 

high value of red coral and the boom and bust exploitation patterns made for unstable yields, with 

various peaks and drops; the progressive overexploitation of red coral in shallow water forced 

divers to descend to greater depths in search of new resources. The last major peak recorded lists 

100 t in 1978, but since then landings have remained below 30 t, on average.

1
Although effective management of red corals is lacking in many Mediterranean nations, others 

have taken steps to promote sustainable harvesting. Since 1994, European Union countries have 

banned the use of the St. Andrew’s cross because it unselectively destroys coral populations and 

habitat. Most but not all nations in the region ban harvest of colonies smaller than 7 mm basal 

diameter, while some countries have no minimum size of harvest, relying instead on a quota or 

limits on license numbers. Some governments, like Sardinia and Morocco, have developed coral 

management plans and regulations that establish harvest quotas and restrict access to coral beds. 

In some countries, poaching has become a growing problem and illegal use of coral dredges are 

still occurring. 

Artisans along Mediterranean coastal communities have worked raw coral into precious jewelry 

and handicrafts for centuries. Red coral jewelry and carved pieces became a staple in trade between 

Europe and Asia and the Americas. It is highly prized in many cultures around the globe. Since 

1805, the center of the red coral processing and manufacturing industry has been Torre del Greco, 

Italy. In Torre del Greco, many artisans working in small family businesses convert raw coral into 

precious jewelry and artistic crafts, which are shipped and appreciated throughout the world.

After listening to a series of presentations on the biology and ecology of red coral and its importance 

in local cultures, and trade, workshop participants divided into 2 working groups on 1) science and 

management, and 2) trade. Abridged conclusions of the working groups are presented below. The 

complete statements of the working groups can be found beginning on page 208 of this report.

Abridged Conclusion of the Science Working Group

The working group reached consensus on the following conclusions and presented them to the 

plenary of the Workshop, which approved them: 

1. A comprehensive Mediterranean approach to red coral management – for example through 

the General Fisheries Commission for the Mediterranean (GFCM) – based on a common 

assessment of current management, monitoring, and research findings across the region is 

needed.

a. Local management plans under this Mediterranean ‘umbrella’ are needed to 

improve management.

b. A working group needs to develop these plans and to seek funding to develop and 

implement them.

2. Management should be based on key scientific parameters such as population structure, 

density genetics, colony and population growth rates (reproduction and recruitment), and 

connectivity. 

3. Management should be adaptive based on feedback from fishery dependent and independent 

data, and input from relevant stakeholders.

4. Populations above 50 meters depth are well studied, at least in Spain, France, and Italy, 

but deeper water populations and populations in other countries are poorly known. 

Extrapolating from shallow water studies (in selected locations) may not be appropriate. 

5. The majority of shallow water populations are, or have been, over-exploited, and the 

overall extent of red coral populations and of their decline is not known at a Mediterranean 

scale. Therefore, there is a clear need to determine the extent of populations and their 

2
decline (or recovery) trends. 

6. Shallow water (less than 70 meters) populations need to be fully protected from harvesting. 

7. Deep water populations (from 70-150 m) are a priority for research and fishery management 

because a) these are thought to contain abundant resources and fisheries are rapidly 

expanding into these areas because shallower areas cannot support commercial fisheries, 

and b) demographic parameters of deep-dwelling populations are poorly known. 

a. These deep water populations should be managed by selective harvesting using 

appropriate size limits based on suitable demographic models. 

b. The deepest populations (> 150 m) are not currently harvested and should not be 

harvested in the future, as they could provide refugia for the species.

8. The decision to ban non-selective gears was a major step forward in the regulation of the 

fishery. Management measures and enforcement in the Mediterranean overall should be 

improved. 

a. Such management measures should be population-specific and include, but not be 

limited to, minimum basal diameter1 and height of colony and branching pattern, 

quotas, regulation of collection methods, harvest reporting and monitoring, rotating 

harvest and an increase in the number and extent of no-take zones/protected areas. 

b. Management measures should be adaptive and be informed by proper and 

comprehensive scientific research on, inter alia, stock status and management 

models. 

c. Absence of scientific information is not a sufficient excuse not to manage 

populations on a precautionary basis, but it is an important incentive to study deepdwelling populations.

9. Management regimes that are not adequately enforced fail to yield the anticipated 

conservation benefits. 

a. There is a strong need for existing and future management measures to be properly 

enforced with inter alia sufficient resources and penalties – including confiscation 

– that are commensurate with the offence and represent a meaningful deterrent. 

10. The working group emphasised the need for further research to underpin the management 

and conservation of the species. 

a. Research should be collaborative and the results shared between Mediterranean 

countries and internationally. 

b. Regular stock assessments and national reporting, in the context of regional 

management, is desirable.

Conclusions of the Management and Trade Working group

It was not possible for the workshop, when considering the report of the trade working group, 

to reach any appropriate consensus on trade related issues because some participants objected 

to any reference to any CITES listings, while others felt that discussion of the implications of 

a hypothetical CITES Appendix-II listing were essential. The final report of the working group, 

pages 211-213 of this report, represents general agreement of the working group but consensus 

was not reached on all of the text.

1 For instance, the working group felt that the minimum basal diameters used in some areas (typically 7 mm) 

should be increased to enable sustainable fisheries.

3
White Papers and/or Abstracts
Section I

Mediterranean Red Coral (Corallium rubrum): 

Historical Assessment

5
THE PRECIOUS RED CORAL OF THE MEDITERRANEAN:

AN HISTORICAL, CULTURAL, AND ECONOMIC HERITAGE

Caterina Ascione

Since the most ancient times, coral has had an extraordinary power of attraction for all peoples. Its 

history is lost in the mists of time, and is intertwined with the ongoing development of Mediterranean 

civilization from East to West. 

It is the history of gods and men, of fears to be overcome, of magic and rituals, of ornaments and 

talismans. Both an amulet and a jewel, then.

The unique biological parabola that is the distinctive characteristic of coral, the complex 

mythological content that surrounds it, tying it to the blood of the head of the Medusa that Perseus 

decapitated, blood that would color and petrify the sea algae, the extraordinary ambiguity of its 

nature, in a sense part of all three kingdoms – animal, vegetable, mineral – brought ancient peoples 

to attribute protective and curative qualities to the “red tree.”

Diverse cultures have indicated it is an antidote for poisons, a remedy against the bite of scorpions, 

able, if mixed with grains of corn during planting, to prevent drought, to keep away the plague, 

calamities, and parasites.

Coral is the exaltation of the symbol of the life force, of the balance between life and death, 

of the possibility of rebirth, subordinate to the complete transformation of one’s essence. In 

this sense, associated to the figure of the Saviour, it indicates, in early Christian symbolism and 

sacred iconography, the blood pact and redemption, the Cross, the palingenesis of Resurrection, 

wonderfully expressed in that marine element that, according to a millenary belief, did not die 

when brought to the surface but, on the contrary, transmuted into a mineral on contact with air, 

becoming perfect and incorruptible, fons vitae perennis.

In addition to being a talisman, a protection for good health and good luck, able to keep away 

negative influences and neutralize the evil eye, coral has always been a widely used ornament, too, 

both plebeian and not.

The art of working coral has criss-crossed the Mediterranean: from Marseilles to Genoa, from 

Trapani to Leghorn to Naples, finally coming to Torre del Greco, a coastal city briefly mentioned 

in various documents and diaries as a continuous font of men dedicated to harvesting the bounty 

of the sea, both fish and coral.

The tie between Torre del Greco and coral has been often noted. A very ancient tie that has had 

two phases: harvesting, testified to from remote times, and the working of the raw material, which 

began early in the 19th century. A tie, beyond the economic and productive aspects, that has brought 

about a true culture of coral that has affected both the life and customs of this place.

Coral harvesting, concentrated in the difficult and dangerous waters of the Mediterranean, is a 

particularly fascinating and dramatic chapter in the history of the city. The earliest documents, 

6
indeed the first to mention Torre del Greco as an autonomous civitas, go back to the end of the 15th 

century. It deals with a request to abolish the taxes imposed on coral fishermen. The tax itself is 

indicative of a large-scale and financially conspicuous activity, extant for some time.

Leaving aside anecdotes, detailed and vivacious though in large part invented, among the 

historically significant milestones that should be mentioned are: the founding of the Pio Monte 

dei Marinai in 1639; a charity created thanks to the initiative of 58 ship owners, with its seat in 

the church of Santa Maria di Costantinopoli; the promulgation of the Codice Corallino in 1789, a 

document of great legal importance that aimed at creating definitive regulations in the turbulent 

world of coral fishing. At the beginning of the 19th century, a new and decisive chapter begins, 

regarding the relationship between Torre del Greco and coral.

On the 27th of March, 1805, Paul Bartelémy Martin was given a ten-year concession for the 

working of “red gold.” This was the decisive step towards the creation of a complete economic 

cycle – harvesting, working, sales – that many had supported and requested. Only a few decades 

later, in 1837, with the end of the concession, there were already numerous shops and eight large 

factories with more than fifty workers. The entire population seemed, therefore, to be involved in 

activities tied to coral, not only in harvesting and decoration, but also in related concerns, from 

supply to shipbuilding.

From the very beginning of coral working in Torre del Greco, the manufactures were quite 

diversified: not only necklaces of innumerable types – pallini, barilotti, cannettine, to mention 

only a few – but also the creation of jewelry and other luxury items was a distinctive feature, using 

complex and refined sculpting techniques.

The rigorous lines of Neoclassic style marked the early works. From the middle of the 19th century, 

instead, the rise of archeological and neo-Renaissance styles enriched the various methods of 

decoration: mythological figures, putti, cherubs and later baccanti, unicorns, amphorae, combined 

and mounted in imposing and wondrous jewels. Virtually in the same years, the consecration of the 

naturalistic style offered the artisans the possibility of creating a vast production of “flowers, leaves, 

and fruit” compositions: splendid bracelets and necklaces celebrate the high level of technique of 

the master sculptors, surprisingly almost all anonymous.

For almost all of the 19th century, only Mediterranean culture was harvested and it was this coral, 

with its shapes and dimensions, that set the outer and inner limits of production, that naturally 

was also conditioned by the political and economic events, in particular on Europe and northern 

Africa. Great interest and surprise were the result of the French government’s decision in 1875 to 

prohibit harvesting with metal tools, which were damaging the coral banks and the seabed, and to 

allow coral harvesting on the Algerian coast for a limited number of boats, restricted to only three 

areas with a ten-year rotation, in order to allow the abundant re-growth of the banks. It was the first 

time that the quantity of coral brought to market was subject to limitations, to protect both the raw 

material, and prevent excessive exploitation.

In those very years an unexpected event shook the world of coral: the discovery of the Sciacca banks, 

a finding that moved the major interest to the coast off Sicily, towards the island of Pantelleria. The 

report by Giovanni and Riccardo Canestrini, on the behalf of the Ministry of Agriculture, Industry 

and Commerce, studying the case, makes a significant point: “The coral harvested (in the sea off 

7
Sciacca) is dead, that is, lacking the shell or sarcosma, in which the polyps live and reproduce (...

This) was announced in the report to the Ministry of Agriculture, Industry and Commerce on the 

results of the search among the coral banks off Sciacca (...) We certainly brought up thousands 

of samples from these banks and many showed the recent fractures caused by our tools, but no 

samples were found with a sarcosma” (Giovanni and Riccardo Canestrini, Il Corallo in the Annali 

dell’Industria e Commercio, 1882).

Wide and rich banks, but of dead coral, with no trace of sarcosma or polyps. These very characteristics 

made reproduction of the banks impossible. The events that the followed the singular harvest at 

Sciacca are well known: an excessive abundance of coral gave rise to a crisis, caused by market 

saturation and depreciation of a large part of the production. In just ten years, the excess of supply 

made its dramatic effects felt and many manufacturers were forced to close. The grave crisis 

and the recognized inadequacy of the Sciacca coral, by now devalued and unsuitable for detailed 

and elaborate workings that the market wanted, caused some companies, in the second half of 

eighties, to put aside the supplies already acquired and to try other sources. A new type of coral 

was discovered in the markets at Madras and Calcutta: it came from Japan.

Coral which was highly-prized and morphologically different from that previously used. Above 

all, imported coral. This broke the continuous economic cycle (harvest-manufacture-sales) that 

had marked Torre del Greco for the entire 19th century. The need to invest large amounts of capital 

for the purchase of the raw material caused a severe selection process among the companies of the 

sector. Its value heavily influenced production, which abandoned lower levels and moved up to 

luxury jewelry, the prerogative of the upper middle class and an aristocratic elite. Modernist and 

Art Decò influences inspired original geometric creations, with strong chromatic contrasts (red 

and white matched with black onyx or green malachite) and heterogeneous pairings of materials: 

iridescent pearl with nuances of coral, with inserts in tortoise shell, painted wood and plastic. 

The predominant use of Oriental coral in the most expensive and exclusive production did not mean 

that Mediterranean coral was no longer used, which on the contrary was kept alive to safeguard 

the harvesting of raw material, fundamental for the economy of Torre del Greco. This was used 

primarily for low-cost items and necklaces in traditional styles, both for domestic and foreign 

markets. In the 1950s, after World War II, there was a increase in sales, slow at first but slowly 

growing. With the reopening of the North African markets, in Nigeria, in the United States, and 

in the Far East, demand grew for both raw and worked coral, which in the meantime had been 

rediscovered by important jewelers such as Cartier, Boucheron, and Van Cleef.

This represented a new phase for Torre del Greco, which continues today. A story which is the history 

of the very city and its people: a unique culture, a universal treasure, an Immaterial Patrimony, 

as cited by Article 2 of the Convention for the Safeguard of Immaterial Cultural Patrimony of 

UNESCO, it is “transmitted from generation to generation, constantly recreated by the community 

and those related to it, in conformity with its environment, by their interaction with Nature and 

their history, and gives them a sense of identity and continuity, by so promoting the respect of 

the cultural and creative diversity.” A patrimony, therefore, to be protected and safeguarded from 

destruction.

Today a renewed sensibility imposes the need to take a different view of the sea and its treasures, 

coral among these. And it points a way, perhaps the only one which can actually be undertaken. 

8
The eco-sustainable exploitation of the resources that the Earth offers. Intelligence and rigorous 

regulation. In the case of coral, the adoption of a system under strict control regarding times, areas 

and methods of harvesting would be not only a guarantee of the on-going defense of coral species, 

but also the certainty, historically documented and proven, of the safeguard of employment levels 

and quality of production, and therefore the maintaining over time of a profitable relationship 

between supply and demand, with undoubtable advantages for the continuing stability of the 

market and prices. A wise choice, and above all with excellent returns, both in terms of ecology 

and economics.

9
LACAZE-DUTHIERS’ LEGACY: ADVANCES IN SCIENTIFIC 

KNOWLEDGE OF CORALLIUM RUBRUM FROM FRENCH WORKS AFTER 1864

Jean-Georges Harmelin

Centre d’Océanologie de Marseille

Station Marine d’Endoume, 13007 Marseille, France

Introduction

The French Mediterranean coast has been for centuries the center of active harvesting, manufacturing, 

trading and scientific study of precious red coral. Antique vestiges found at Marseilles, the oldest 

port in France, attest that red coral was already collected there 26 centuries ago. This activity was 

permitted by the abundance and easy accessibility of red coral stocks along the Provence coasts. 

This opportunity was also seized by naturalists, such as L.F. de Marsigli and J.A.Peyssonnel, the 

last actors with opposite opinions of the old controversy about the mineral, vegetal, or animal 

nature of the red coral, which was solved at Marseilles in the early 18th century (McConnell 1990).

The scientific knowledge of Corallium rubrum benefited greatly from the work by Lacaze-Duthiers 

(1864) during his stay at El Kala (Algeria), where he was commissioned by the French government. 

This major step was followed by a long gap of nearly one century in French research devoted to the 

precious coral, except for two papers. The renewal of interest for C. rubrum started at the end of the 

1950s when SCUBA diving became a common tool for studying hard bottom communities. This 

method was particularly workable in the NW Mediterranean as red coral commonly thrives there at 

shallow depths, especially along the French coasts. The number of references increased noticeably 

from the 1970s and reaches now 60 (including the works realised at the Centre Scientifique de 

Monaco). One of the conclusions of the first FAO meeting on red coral held at Mallorca in 1983 

was that all questions addressed by Lacaze-Duthiers were still unsolved. Uncertainties concerned 

particularly the growth rate of colonies and thus the capacity of red coral banks to replenish after 

exploitation. Since then, significant advances were provided by works realized in France and 

Monaco, which are resumed below, while the entire list of references is given in bibliography.

Habitat Distribution of Red Coral

The ecological attributes of C. rubrum and features of its distribution among hard-bottom habitats 

were described in a series of papers (Bianconi et al. 1988; Laborel and Vacelet 1961; Stiller and 

Rivoire 1984; Weinberg 1975, 1978, 1979a, 1979b, 1979c). The most outstanding peculiarity 

of the Mediterranean red coral compared to all other Corallium or Paracorallium species is its 

capacity to occupy near-surface habitats as well as much deeper biotopes. On the upper side of this 

range, patchy populations bordered by photophilic algal assemblages can be found in cavities and 

overhangs at few meters below the surface (3-6 m). On the opposite, deeper side of this distribution, 

C. rubrum can coexist with the bathyal cold-water coral Madrepora oculata at several hundred 

meters depth and short distance from the coast. This was observed, for example, on the abrupt 

walls of Cassidaigne submarine canyon, near Marseille, at 200-325 m (Bourcier and Zibrowius 

1973; Laborel et al. 1961; and unpublished ROV observations during Hermes and AAMP surveys, 

2009). Although these nearshore and deep-water habitats may appear in stark contrast, they share 

10
some common environmental characters, such as dim light, steep slope (i.e. limited sediment 

deposits), and circulating water.

Thermal Sensitivity of Red Coral 

Field evidences indicate that the upper depth at which C. rubrum occurs presents a great geographical 

variability around the Mediterranean, hypothetically related to local surface temperatures and 

deepening of the thermocline in summer (Laborel and Vacelet 1961; Stiller and Rivoire 1984). 

Moreover, red coral is subjected to mass mortality in the shallower sites when positive thermal 

anomalies happen during summer periods (Garrabou et al. 2001, 2009; Harmelin 1984). The 

sensitivity of red coral to high temperatures had thus to be tested experimentally.

The thermo-tolerance patterns of red coral were studied by O. Torrents by experiments in thermoregulated tanks with four types of thermal regime applied during two consecutive years to 

specimens coming from shallow and deeper sites. The responses (coenenchyme necrosis, polyp 

activity, calcification rate) of red coral colonies in experimental conditions were similar to those 

observed during mortality events and brought precise information on the thermo-tolerance limits 

of this species. Long exposure of colonies to 24° C and shorter exposure to 25° C are critical to 

both shallow and deep specimens; necrosis occurs in less than five days if temperature is higher 

than 25° C and deep populations are more sensitive than shallower ones (Torrents et al. 2008).

These experiments confirm the particular thermal sensitivity of C. rubrum, and therefore that (i) 

the shallower populations of the NW Mediterranean are under the threat of global warming, and 

(ii) high summer temperatures are a key factor of its exclusion from shallow depths in the warmer 

regions, such as the eastern Mediterranean (Zibrowius 1979). 

Diet of Red Coral 

As other azooxanthellate passive suspension-feeders, red coral depends entirely on food supplied 

by the water movements. This constraint limits, for example, its penetration inside caves when 

confinement predominates. Red coral is known to consume particulate organic matter, both live 

and detrital. However, feeding experiments by Picciano and Ferrier-Pagès (2007) showed that 

red coral is also able to prey effectively on bacterioplankton. Although pico- and nanoplankton 

represent only 4.5% of the total carbon ingested, this food appears to be essential in terms of 

nitrogen and phosphorus. 

Resilience of Harvested Populations 

It is evident from field observations that, at least in some regions such as Provence (France), red 

coral persists in shallow sites despite a long history of harvesting pressure and easy accessibility. 

Thus, how these shallow populations can persist and replenish?

The study of reproduction of C. rubrum (Torrents et al. 2005) showed that the size of the smallest 

fertile colonies was surprisingly small, only 2.4 cm in average (height), the smallest being 1.5 cm 

tall. Thus, these smallest fertile individuals are far below the threshold size that can trigger a selective 

harvesting. This feature together with the poor dispersal of larvae, which favour self-seeding in 

populations aggregated in cavities (Weinberg 1979b), allow local populations to persist with 

11
small colony size in highly frequented sites. However, age analysis showed that these small fertile 

colonies were not juvenile, but 7-10 years old (Torrents et al. 2005). It was also clear from this study 

that the larval productivity increases considerably with the colony size. Therefore, conservation of 

large-sized colonies, most efficient in producing larvae and icons of the Mediterranean landscape 

heritage, is imperative. This is particularly acute when local populations are formed of scattered 

colonies, more subjected to the hazards of recruitment than patchy populations.

Recruitment Patterns and Colonization 

Underwater observations show that assemblages including red coral in dimly-lit habitats are 

dominated by long-lived, highly-competitive components (sponges, tunicates, bryozoans, serpulids, 

scleractinian corals, etc.) (Laborel and Vacelet 1961; Bianconi et al. 1988; Weinberg 1978). How 

does C. rubrum participate to the establishment and functioning of this type of communities? 

Limestone panels deployed in 1969 within a Corallium-rich community were photographically 

monitored yearly since 1979, i.e. two years after a first pulse settlement of red coral colonies. Only 

another recruitment event was observed during the 22 years-long monitoring, but survivorship of 

colonies was high (60%). In situ measurements of colony branches showed that linear growth was 

in average only 5.6 mm year-1 (Harmelin 1984), while measurements of the basal diameter of 22-

year old colonies showed that the radial thickening of branches was very slow, only 0.24 mm year1

 (Garrabou and Harmelin 2002). This long-term colonisation experiment brought indisputable 

indications on the red coral growth rate, but could hardly be repeated for testing the variability of 

growth in diverse conditions.

Skeleton Features and Growth Rate Determination

The tree-like growth-form of red coral suggested the possible use of dendrochronology to date 

colonies. The first attempt was made by Dantan (1928), who made horizontal cross sections in 

the axial calcified skeleton and found a few concentric zones, e.g. only five rings in a 1 cm thick 

branch. To validate this technique, he proposed another pioneer idea: monitoring of transplanted 

colonies by divers. However, this petrographic method leads to a considerable over-estimation of 

growth-rate because apparent rings are pluriannual, as proved by tests on colonies from panels, i.e. 

of known age (Marschal et al. 2004).

Another approach was to look for patterns in the organic part of the calcareous skeleton. Fundamental 

advances in the biochemical characterization of the organic matrix of C. rubrum and its role in 

biomineralization processes were done by D. Allemand and his team at Monaco (Allemand 1993; 

Allemand and Grillo 1990, 1992; Allemand et al. 1994; Grillo et al. 1993). Biochemical evidences 

show that spicules and axial skeleton have a different origin although the organic matrix is present 

in both skeletal elements with a water-insoluble fraction (76-80%) and a soluble matrix, which is 

richer in proteins. 

The finding by C. Marschal (Marschal 2004; Marschal et al. 2004) that the organic matrix reflected 

an annual periodicity in its concentration resulting in concentric dark and clear rings was a major 

step. The annual periodicity of these rings was validated first with colonies of known age settled 

on panels (Garrabou and Harmelin 2002) and also by using repeated calceine staining on colonies 

transplanted during fixed times. The organic matrix method, now used in routine, allowed for 

12
example to show evident differences in growth rate at small spatial scale according to the exposure 

of colonies to water circulation.

A quite different approach, based on features of the calcareous skeleton mineralogy, was tested 

by D. Vielzeuf. Mineralogical analyses of skeleton cross-sections showed that the magnesium/

calcium ratio presents oscillations, which are annual with more magnesium deposited during the 

warm season. Interestingly, the comparison of the two methods, organic matrix and magnesium 

cartography, on the same skeleton sections, proved that they are perfectly congruent (Vielzeuf et 

al. 2008).

Biocalcification and the Skeleton Construction 

It was admitted, following Lacaze-Duthiers (1864), that the construction of the axial skeleton 

resulted from the incorporation and cementation of spicules. It was argued that spiny protuberances 

occurring at the surface of the axial skeleton were spicules in the process of being incorporated. 

This hypothesis was tested with different modern tools. Biochemical analyses and other methods 

such as radioisotope incorporation were used by D. Allemand and his team to trace the uptake of 

calcium and describe the calcification of the axis (Allemand 1993; Allemand and Grillo 1990, 

1992; Allemand and Bénazet 1992; Allemand and Bénazet-Tambutté 1996; Grillo and Allemand 

1990). It is now proved that the skeleton growth follows two different patterns, one at the tip of 

branches, where spicules are incorporated, and another in the rest of the colony with the activity of 

a peripheral skeletogenic epithelium which is responsible of the axis thickening. 

The mineral microstructure of the axial skeleton has been recently studied by Vielzeuf and his team. 

The outstanding hierarchic structure of the biomineral skeleton was revealed from the nanoscale 

to the macroscale with nanograins, fibers, blades, and microprotuberances (Vielzeuf et al. 2008, 

in press). This microstructure approach also allowed to confirm that microprotuberances are not 

partly incorporated sclerites, as believed by Lacaze-Duthiers.

Red Coral Color 

The color of the skeleton of C. rubrum is a fundamental attribute for jewelry, which may vary among 

regional populations from dark red to pink, but also within local populations with occasional albino 

colonies (Harmelin 2000). The red color was first attributed to iron oxide (Ranson and Durivault 

1937). However, analyses by resonance Raman spectroscopy (Merlin and Delé 1983; Merlin and 

Delé-Dubois 1985) indicated that carotenoids were responsible of the skeleton red color. The use of 

high-performance liquid chromatography (HPLC/DAD and HPLC/MS) by Cvejic et al. (2007) on 

both soft and hard tissues confirmed the carotenoid origin of the pigmentation and more precisely 

showed the presence of canthaxanthin, a dominant carotenoid in aquatic animals. Canthaxanthin 

is contained in the organic matrix and its concentration is notably greater in spicules than in the 

skeleton. Carotenoids are provided by a wide variety of food items and their role in the skeleton is 

still unknown, as well as the origin of the wide variety of tones.

Size Structure in Harvested vs. Non-Harvested Populations 

It is manifest from underwater observations in the 0-60 m zone that large colonies are now 

extremely rare while they were accounted to be not infrequent during the 50s or early 60s even 

13
at shallow depth. On the other hand, the difference in colony size structure between populations 

located in harvested zones and those in areas protected for long is also visually obvious. These 

size differences were evidenced by direct measurements (basal diameter, height, width) of the 

largest colonies found in multiples spots within a region including harvested and protected sites 

(Garrabou and Harmelin 2002; Harmelin et al. 1999). The “harvesting erosion” affecting the 

size of colonies is impressive: the mean basal diameter of the largest colonies was 6-7 mm in 

unprotected sites versus 12 mm in a marine reserve. Precise in situ measurements of colonies 

are time-consuming and entail long decompression stages. Now this method is advantageously 

replaced by the underwater photogrammetry of colonies (Graille 2007), which can provide rapid 

and reliable size data. Photogrammetric surveys of harvested populations compared to those 

sheltered in three marine protected areas have confirmed the obvious effect of harvesting on the 

population size structure (Bianchimani 2005) and the existence of size thresholds that determine 

collect attractiveness.

Conclusions: Lessons to Management

The Mediterranean red coral presents an outstanding capacity to cope with different local conditions 

leading to a broad ecological distribution. However, a particular sensitivity to warm temperatures 

and thus to global change can be a grave threat to the shallowest populations. It is a remarkable 

model for biomineralisation studies with peculiar patterns of skeleton structure and growth. It is now 

proved that its growth is very slow, even in the most productive sites. However, despite harvesting, 

it is not a threatened species thanks to its capacity to reproduce at very small size. The large, very 

old colonies (equivalent to the biggest trees in forests) are no longer present in most sites and will 

soon totally disappear. Permanent sanctuaries of significant size must be established all around the 

Mediterranean in order to allow some local populations to reach a natural size structure including 

very old colonies in the next centuries. Harvesting must be strictly controlled everywhere with 

size limits, education and precise registration of professionals, and with eradication of poaching. 

Regulations should be adapted to local conditions.

References 

Compilation of papers on Corallium issued from French and Monaco marine laboratories since 

Lacaze-Duthiers’ book: Lacaze-Duthiers H (1864) Histoire naturelle du corail. J.B. Baillière (Ed.) 

Paris, 371 p.

Allemand, D. 1992. Le corail rouge, cet inconnu. In: Association Monégasque pour la Protection de 

la Nature (Ed.) Réserve sous-marine à Corail de Monaco. Expériences de coralliculture en 

milieu naturel. AMPN, Monaco, 1-7.

Allemand, D. 1993. The biology and skeletogenesis of the Mediterranean red coral. A review. 

Precious Corals and Octocoral Res 2: 19-39.

Allemand, D., Grillo, M.C. 1990. La biocalcification chez le corail rouge, Corallium rubrum. 2. 

Approches biochimique et physiologique. Rapp Comm Int Mer Médit 32, 1: 148.

Allemand, D., Bénazet, S. 1992. Dynamique de la calcification chez le corail rouge, Corallium 

rubrum. Rapp Comm Int Mer Médit 33: 27.

14
Allemand, D., Cuif, J.P., Watanabe, N., Oishi, M., Kawaguchi, T. 1994. The organic matrix of 

skeletal structures of the Mediterranean red coral, Corallium rubrum. In: Allemand, D., 

Cuif, J.P. (Eds.) 7th Int. Symp. Biomineralization, Monaco, 17-20 November 1993. Bull Inst 

Océanogr Monaco, no spécial 14, 1: 129-139.

Allemand, D., Debernardi, E., Seaman, Jr. W. 2000. Artificial reefs in the Principality of Monaco: 

Protection and enhancement of coastal zones. In: Jensen, A., Collins, K., Lockwood, A. 

(Eds.) Artificial reefs in European Seas. Kluwer Academic Publishers, Dordrecht. 151-166.

Allemand, D., Grillo, M.C. 1992. Biocalcification mechanism in Gorgonians: 45Ca uptake and 

deposition by the Mediterranean red coral Corallium rubrum. J Exp Zool 262: 237-246.

Allemand, D., Bénazet-Tambutté, S. 1996. Dynamics of calcification in the Mediterranean red coral, 

Corallium rubrum (Linnaeus) (Cnidaria, Octocorallia). J Exp Zool 276: 270-278.

Arnoux, A., Harmelin, J.G., Monod, J.L., Romaña, L.A., Zibrowius, H. 1992. Altérations des 

peuplements benthiques de roches profondes en Méditerranée nord-occidentale: quelques 

aspects biologiques et molysmologiques. CR Acad Sci Paris 314 (III): 219-225.

Association Monégasque pour la Protection de la Nature. 1992. Réserve sous-marine à Corail de 

Monaco. Expériences de coralliculture en milieu naturel. AMPN publ., Monaco, 1-17.

Bianchimani, O. 2005. Evaluation des effets des aires marines protégées sur les populations de corail 

rouge (Corallium rubrum) : le cas des réserves marines françaises. Rapport DU. Université 

de la Méditerranée.

Bianconi, C.H., Rivoire, G., Stiller, A., Boudouresque, C.F. 1988. Le corail rouge Corallium rubrum 

dans la réserve naturelle de Scandola (Corse). Trav Scient Parc Nat Région Réserves Nat 

Corse 16: 1-80.

Bourcier, M., Zibrowius, H. 1973. Les «boues rouges» déversées dans le Canyon de la Cassidaigne 

(région de Marseille). Observations en soucoupe plongeante SP 350 (juin 1971) et résultats 

de dragages. Téthys 4 (1972): 811-842.

Calderón, I., Garrabou, J., Aurelle, D. 2006. Evaluation of the utility of COI and ITS markers as tools 

for genetic studies of temperate gorgonians. J Exp Mar Biol Ecol 336: 184-197.

Carpine, C., Grasshoff, M. 1976. Les gorgonaires de la Méditerranée. Bull Inst Océanogr Monaco 

1430: 1-140.

Cvejic, J., Tambutté, S., Lotto, S., Mikov, M., Scalanin, I., Allemand, D. 2007. Determination of 

canthaxanthin in the red coral (Corallium rubrum) from Marseille by HPLC combined with 

UV and MS detection. Mar Biol 152: 855-862.

Cuif JP, Lafuste J, Lawniczak A (1985) Evolution microstructurale au cours de l’ontogenèse chez 

un Alcyonaire actuel: passage fibres/lamelles dans le squelette axial de Corallium johnsoni

Gray (Cnidaria, Octocorallia). CR Acad Sc Paris 300 sér. II, 5: 181-184.

Dantan, J.L. 1928. Recherches sur la croissance du corail rouge (Corallium rubrum Lamarck). Bull 

Soc Zool France 53: 42-46.

Dauphin, Y. 2006. Minelarizing matrices in the skeletal axes of two Corallium species (Alcyonacea). 

Comparative Biochimistry and Physiology part A, 145: 54-64.

15
Garrabou, J., Pérez, T., Sartoretto, S., Harmelin, J.G. 2001. Mass mortality event in red coral 

(Corallium rubrum, Cnidaria, Anthozoa, Octocorallia) populations in the Provence region 

(France, NW Mediterranean). Mar Ecol Prog Ser 217: 263-272.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study of life history traits of a harvested long-lived 

temperate coral in NW Mediterranean: insights into conservation and management needs. J 

Anim Ecol 71: 966-978.

Garrabou, J., Bianchimani, O., Fourt, M., Graille, R., Ledoux, J.B., Marschal, C., Zuberer, F., 

Dominici, J.M. 2006. Etat actuel de conservation et caractérisation des populations de corail 

rouge (Corallium rubrum) de la réserve naturelle de Scandola, Parc Régional de Corse. 

Rapport COM-DIMAR and Parc régional de Corse, 18 p.

Garrabou, J., Coma, R., Bensoussan, N., Bally, M., Chevaldonne, P., Cigliano, M., Diaz, D., Harmelin, 

J.G., Gambi, M.C., Kersting, D.K., Ledoux, J.B., Lejeusne, C., Linares, C., Marschal, C., 

Pérez, T., Ribes, M., Romano, J.C., Serrano, E., Teixido, N., Torrents, O., Zabala, M., 

Zuberer, F., Cerrano, C. 2009. Mass mortality in Northwestern Mediterranean rocky benthic 

communities: effects of the 2003 heat wave. Global Change Biology doi: 10.1111/j.1365- 

86.2008.01823.x

Graille, R. 2007. Mise au point et utilisation des techniques photogrammétriques pour la mesure 

d’organismes marins benthiques dans leur milieu. Mémoire diplôme Ecole Pratique des 

Hautes Etudes. Sciences de la Vie et de la Terre. Laboratoire EPHE Ecosystèmes aquatiques 

tropicaux et méditerranéens, Perpignan and Centre d’Océanologie de Marseille, 73 p.

Grillo, M.C., Allemand, D. 1990. La biocalcification chez le corail rouge, Corallium rubrum. Rapp 

Comm Int Mer Médit 32, 1: 21.

Grillo, M.C., Chessa, L.A. 1992. Développement larvaire de Corallium rubrum. Rapp Comm Int 

Mer Médit 33: 40.

Grillo, M.C., Goldberg, W.M., Allemand, D. 1993. Skeleton and sclerite formation in Mediterranean 

red coral. Mar Biol 117: 119-128.

Harmelin, J.G. 1980. Etablissement des communautés de substrats durs en milieu obscur. Résultats 

préliminaires d’une expérience à long terme en Méditerranée. Mem Biol Marina e Oceanogr 

Suppl.X: 29-52.

Harmelin, J.G. 1984. Biologie du Corail Rouge. Paramètres de populations, croissance et mortalité 

naturelle. Etat des connaissances en France. In: Charbonnier, D.Y., Garcia, S. (Eds.) 

Rapport de la Consultation technique du CGPM sur les ressources de Corail Rouge de 

la Méditerranée occidentale et leur exploitation rationnelle: 99-103. Palma de Mallorca, 

Espagne, Déc. 1983.

Harmelin, J.G. 2000. Le corail rouge de Méditerranée: quelques aspects de sa biologie et de son 

écologie. In: Morel, J.P., Rondi-Costanzo, C., Ugolini, D. (Eds.) Corallo di Ieri, Corallo 

di Oggi, Atti del Convegno, Ravello, Villa Rufolo, 13-15 Dicembre 1996: 11-20. Centro 

Universario Europeo per i Beni Culturali, Ravello. Travaux Centre Camille Jullian, 

Université de Provence, 25. Edipuglia, Bari.

Harmelin, J.G., Zibrowius, H., Arnoux, A., Romaña, L.A. 1991. Evaluation de l’état des peuplements 

benthiques du haut-fond St Julien (région de St Tropez). Campagne Cyana de mars et juin 

1990. IFREMER Rapport n° DRO/EM 91-01.

16
Harmelin, J.G., Sartoretto, S., Francour, P. 1999. Mise en place d’une stratégie de suivi de 

l’ichtyofaune et des peuplements de gorgonaires de l’archipel de Riou. Rapport Ville de 

Marseille, Direction de l’Environnement et des Déchets, 110 p.

Laborel, J., Pérès, J.M., Picard, J., Vacelet, J. 1961. Etude directe des fonds des parages de Marseille 

de 30 à 300 m avec la soucoupe Cousteau. Bull Inst océanogr Monaco 1206: 1-16.

Laborel, J., Vacelet, J. 1961. Répartition bionomique du Corallium rubrum Lamarck dans les grottes 

et les falaises sous-marines. Rapp Comm Int Mer Médit 16, 2: 465-469.

Lawniczak, A. 1987. Les modalités de croissance de l’axe calcaire chez Corallium johnsoni. 

Senckenberg Marit 19: 149-161.

McConnell, A. 1990. The flowers of corals. Some unpublished conflicts from Montpellier and Paris 

during the early 18th century. History and Philosophy of the Life Science 12: 51-66.

Marin, J., Reynal de Saint-Michel, L. 1981. Premières données biométriques sur le corail rouge, 

Corallium rubrum LMCK, de Corse. Rapp Comm int Mer Médit 27, 2: 171-172.

Marschal, C. 2004. Mise au point d’une nouvelle technique de détermination de l’âge de Corallium 

rubrum basée sur la mise en évidence de cernes d’accroissement dans la matrice organique. 

Mémoire diplôme Ecole Pratique des Hautes Etudes. Sciences de la Vie et de la Terre. 

Laboratoire EPHE Biologie Marine et Malacologie, Perpignan and Centre d’Océanologie 

de Marseille, 91 p. 

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in the precious red coral Corallium rubrum (L.). Coral Reef 23: 423-432.

Merlin, J.C., Delé, M.L. 1983. Etude par spectroscopie Raman de résonance de la pigmentation des 

squelettes calcaires de certains coraux. Bull Soc Zool France 108, 2: 289-301.

Merlin, J.C., Delé-Dubois, M.L. 1985. Resonance Raman characterization of polyacetylenic pigments 

in the calcareous skeleton. Comp Biochem Physiol 84B: 97-103.

Mokhtar-Jamaï, K., Ledoux, J.B., Garrabou, J., Féral, J.P., Aurelle, D. 2009. Interest and application 

of genetic markers for the study and conservation of Mediterranean sessile invertebrates. 

In: Pergent-Martini, C., Brichet, M. (Eds.) Actes du 1er Symposium méditerranéen sur la 

conservation du Coralligène et autres bio-concrétions calcaires (Tabarka, 15-16 Janvier 

2009): 112-116 CAR/ASP publ, Tunis.

Picciano, M., Ferrier-Pagès, C. 2007. Ingestion of pico- and nanoplankton by the Mediterranean red 

coral Corallium rubrum. Mar Biol 150: 773-782.

Ranson, G., Durivault, A. 1937. Le pigment d’Heliopora coerulea et de quelques autres alcyoniaires. 

CR Soc Biol Paris 126: 1149-1151.

Ranson, G. 1987. Existence de champs de corail rouge et de gorgones morts en profondeur entre 

Cassis et Nice. Rapport Direction Affaires Maritimes de Méditerranée, 16 p. 

Ranson, G. 1991. Mortalité de corail et de gorgones en profondeur au large des côtes provençales. 

In: Boudouresque, C.F., Avon, M., Gravez, V. (Eds.) Les espèces marines à protéger en 

Méditerranée: 53-59. GIS Posidonies, France. 

Stiller, A., Rivoire, G. 1984. Biologie et écologie du corail rouge en Méditerranée française. In 

Charbonnier, D.Y., Garcia, S. (Eds.) Rapport de la Consultation technique du CGPM sur les 

ressources de Corail Rouge de la Méditerranée occidentale et leur exploitation rationnelle: 

89-93. Palma de Mallorca, Espagne, Déc. 1983.

17
Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in the 

commercially exploited red coral Corallium rubrum (L.) (Octocorallia, Gorgonacea) in the 

Marseilles area (France, NW Mediterranean). Biological Conservation 121 (3): 391-397.

Torrents, O. 2007. Biologie des populations du corail rouge Corallium rubrum (L. 1758) de 

Méditerranée nord-occidentale. PhD thesis Université d’Aix-Marseille II, 221 p.

Torrents, O., Tambutté, E., Caminiti, N., Garrabou, J. 2008. Upper thermal thresholds of shallow vs. 

Deep populations of the precious Mediterranean red coral Corallium rubrum (L.): Assessing 

the potential effects of warming in the NW Mediterranean. J Exp Mar Biol Ecol 357: 7-19.

True, M.A. 1970. Etude quantitative de quatre peuplements sciaphiles sur substrat rocheux dans la 

région marseillaise. Bull Inst Océanogr Monaco 69: 1-48.

Vielzeuf, D., Garrabou, J., Baronnet, A., Grauby, O., Marschal, C. 2008. Nano to macroscale 

biomineral architecture of red coral (Corallium rubrum). American Mineralogist 93, 11-12: 

1799-1815.

Vielzeuf, D., Floquet, N., Chatain, D., Bonneté, F., Ferry, D., Garrabou, J., Stolper, E.M. (in press) 

Multilevel modular mesocrystalline organization in red coral. American Mineralogist.

Weinberg, S. 1975. Ecologie des octocoralliaires communs du substrat dur dans la région de Banyulssur-Mer. Essai d’une méthode. Bijdragen tot de Dierkunde 45, 1: 50-70.

Weinberg, S. 1976. Revision of the common Octocorallia of the Mediterranean circalittoral. I. 

Gorgonacea. Beaufortia 313: 63-103.

Weinberg, S. 1978. Mediterranean octocorallian communities and the abiotic environment. Mar Biol 

49: 41-57.

Weinberg, S. 1979a. Autoecology of shallow-water Octocorallia from Mediterranean rocky substrata. 

I. The Banyuls area. Bijdragen tot de Dierkunde 49, 1: 1-15.

Weinberg, S. 1979b. The light dependent behavior of planula larvae of Eunicella singularis and 

Corallium rubrum and its implication for octocorallian ecology. Bijdragen tot de Dierkunde 

49, 1: 16-30.

Weinberg, S. 1979c. Transplantation experiments with Mediterranean gorgonians. Bijdragen tot de 

Dierkunde 49, 1: 31-41.

Zibrowius, H. 1979. A propos du corail rouge en Méditerranée orientale. Rapp Comm Int Mer Médit 

25/26, 4: 121-122.

Zibrowius, H., Monteiro, M., Grasshoff, M. 1984. La répartition de Corallium rubrum dans 

l’Atlantique. Thétys 11: 163-170.

18
THE HISTORY OF RESEARCH AND HARVESTING OF THE 

MEDITERRANEAN RED CORAL: POTENTIAL AND LIMITS

Giovanni Santangelo1

, Lorenzo Bramanti1,2, Ilaria Vielmini1, Mimmo Iannelli3

1

 Zoology and Animal Ecology. CSIC University of Pisa, Italy.

2

 Institut de Ciencias del Mar CSIC Barcelona, Spain

3

 Dep. Mathematics, University of Trento Italy

Summary

The history of the research on the Mediterranean red coral (Corallium rubrum L 1758) is strictly 

intertwined with that of harvesting. For more than three centuries all the information on red coral 

habitat and all the material examined by naturalists was gathered by fishermen; moreover, the 

oscillating succession of red coral fishing vicissitudes deeply affected the development of red coral 

research. Up to the XVII century, red coral was considered to be a plant, and the first naturalist who 

stated the animal nature of red coral was the Neapolitan alchemist Farinella, and only subsequently 

Peyssonel in 1723 confirmed this statement. A following milestone in red coral research was the 

beautiful book of the French scientist Lacaze-Duthiers, who, in 1864 published: History naturelle 

du corail. Only in the second half of the XX century the research on red coral started again and 

in situ researches were carried out by Laborel and Vacelet (1958, 1961) and Weinberg (1978, 

1979a, 1979b), who described the sensitivity of red coral to light, sedimentation, hydrodynamics, 

and the red coral habitat selectivity. In the same period, a team of Italian researchers (Barletta, 

Marchetti, Vighi 1968) described, on the basis of data collected directly by SCUBA diving, several 

red coral populations living in shallow-waters (between 20 and 50 meter depth) along the Italian 

coasts. Moreover, they described for the fist time the reproductive cycle of this species. In 1986, 

Spanish researchers (Rodriquez and Massò 1986) described red coral population size structure 

and allometric relationships on the basis of Catalan fishing crops; moreover, they try to determine 

the age of red coral colonies. In 1991 and 1994, after two meetings organised by FAO in Torre 

del Greco and Alger, and the submission of a Spanish proposal to include red coral in CITES 

Appendix-II, new impulse was given to red coral research by the Italian Government, who funded 

two preliminary research projects. These researches led to two basic books on red coral biology 

and ecology (Red coral in the Mediterranean Sea, Art history and Science 1993; Red Coral and 

other Mediterranean Octocorals: Biology and Protection 1999). It will be worth to remember that 

nearly all the papers already published deal with shallow-water populations which have a limited 

economic value. More recently red coral research was addressed towards the study of population 

structure and dynamics by means of demographic models (Abbiati et al. 1992; Santangelo et al. 

2004, 2007; Bramanti et al. 2009). This approach, widely applied in Conservation Ecology, will 

supply the tools to match harvesting to population growth rate fostering both conservation and 

rational management of the populations of this precious species, if sound data on deep-dwelling 

populations will be at least gathered and enforcement will be really applied at local population 

level.

A Long History

The history of the research on the Mediterranean red coral (Corallium rubrum L 1758) is strictly 

intertwined with that of harvesting as oscillating succession of red coral fishing vicissitudes deeply 

19
affected the development of red coral research. Moreover, along three centuries all the material 

examined by naturalists was gathered by fishermen. This first phase continued up to the second half 

of 1950s, when SCUBA-Diving techniques allowed to researchers to observe red coral populations 

directly in the field.

Three different phases in red coral research can thus be sketched:

1. XVII-XIX Naturalists gathered all the samples they examined and all information by fishermen.

2. Since 1950s to nowadays SCUBA-Diving scientists acquired the possibility to collect 

their data directly in situ; these studies were limited to the shallower portion of red 

coral bathymetric distribution (20–50 m) At the same time electron microscopy 

allowed to gain insights into the small-scale structure of the red coral skeleton. 

3. More recently, the use of ROV and submarines has extended research to deeper populations 

and new, reliable methods allow to determine colony age and to apply demographic models, 

structured in age classes, which will foster a rational management of this precious Mediterranean 

resource.

In the course of the long history of red coral exploitation (two thousand years, at least) and the 

even longer history of red coral use as precious raw material for carving talismans, art objects, and 

jewellery, several legends and fancy hypotheses raised on its origin and real nature. Up to the XVII 

century, red coral was considered firstly to be a mineral and then a plant. The first naturalist who 

stated the animal nature of this species was the Neapolitan alchemist Finella in 1649; only later 

the Italian naturalist Cestoni (1718), and finally the French physician Peyssonnel (1725) confirmed 

the animal nature of red coral. 

Fig. 1. Red coral colony with expanded polyps in a crevice at 35 m depth.

Fig. 2. Red coral, flowered plant or animal? The truth revealed in the XVIII 

century.

20
The fist naturalist who tried to estimate the growth rate of red coral colonies was the Italian botanist 

Marsili (1725), who observed at about 5 meters depth red coral colonies living in a shallow-water, 

shaded cave. Red coral still dwells nowadays at such depth in some long, dark, cold caves like 

those at Capo Caccia (Sardinia) and near Marseille. 

In 1785, Cavolini provided a microscopic description of the skeleton of red coral. Linnaeus in his 

“Sistema naturae” (1758) named the species Madrepora rubra and finally Lamark (1816) gave to 

the species its present name: Corallium rubrum.

The British naturalist Ellis, in 1753, discovered the similarities between gorgonians and Corallium 

rubrum, which was thus included in the Family Keratopytae = Gorgonacea.

A following milestone in red coral research was the beautiful book of the French scientist LacazeDuthiers, who, in 1864 published: History naturelle du corail. This eminent naturalist gave an 

exhaustive description of geographic distribution, ecology, and colony anatomy (Weinberg 1993). 

His skill in observing and describing by drawing magnificent color plates (Fig. 3) of the main 

morphological features of red coral at a macroscopic and microscopic scale was incredible, if 

we consider the limited microscope supply researchers have at that time. It is worth to remember 

that, also in this case, top-quality research was produced in the framework of a fishing campaign 

promoted in Algeria by the France Government. 

Lacaze-Duthiers’ high quality observations and insights were confirmed only at the beginning of 

XX century, when more efficient microscopic tools were developed by two German researchers 

(Koch 1991; Müller 1910), working at Stazione Zoologica di Napoli. 

In the mid XX century, the research on red coral received new impulse due to SCUBA diving 

gears development. In situ research was carried out by Laborel and Vacelet (1958, 1962) and then 

Weinberg (1978, 1979a, 1979b), who described the sensitivity of red coral to light, sedimentation, 

hydrodynamics, and red coral habitat selectivity. Carpine and Grashoff described the features of red 

coral sclerites and the geographic distribution of the species (Carpine and Grashoff 1975). At the 

Fig. 3. From Lacaze-Duthiers 

(1964). At left: the first description 

of red coral recruits; red coral 

female polyp with oocytes in 

two different phases of their 

reproductive cycle. 

21
same time, a team of Italian researchers (Barletta, Marchetti, Vighi, 1968) described, on the basis 

of data collected directly by SCUBA diving, several red coral populations living in shallow-waters 

(between 20 and 50 meters depth) along the Italian Western coasts. Moreover, Vighi described for 

the fist time by means of histological tools, the reproductive cycle of red coral (Vighi M.; Etude 

sur la reproduction du Corallium rubrum Vie et Milieu 1972).

Overfishing at Alboran and the Following Research Impulse in Spain

In 1983, the discovery of rich red coral banks at Alboran (Santangelo et al. 1993b) led to their 

uncontrolled harvesting and to a consequent rapid depletion by fishermen from Sicily; all this led 

to the first submission of a proposal of red coral inclusion in CITES Appendix-II, by Spain, the 

country to which Alboran Islet and surrounding sea belong, with the aim to control overexploitation 

by controlling trading. Under this new situation the Spain Government fostered and funded some 

research on red coral. This is a further example of the way in which the oscillating succession of 

red coral fishing vicissitudes deeply affected the development of red coral research. 

The results of these researches were collected in a monographic volume of the Boletin Espanol 

de Oceangrafia (1986) dedicated to different topics of red coral natural history (Figs. 4, 5, 6). 

There the Spanish researchers Garcia-Rodriquez and Massò (Garcia-Rodriquez and Massò 1986) 

described for the first time the frequency distribution of size and the allometric relationships of 

red coral colonies in Spanish fishing crops. Moreover, they tried to determine the age of colonies 

to convert the size structure they found in an age structure but, unfortunately, their petrographic

method revealed to be erroneous (Garrabou and Harmelin 2002). Only 18 years later a new, reliable 

method, based on organic matrix staining of thin-sections of colony skeleton revealed the real 

annual growth rings of red coral (Marschall et al. 2004).

Fig. 4. Survivor curve (Number of colonies 

of different size/age) drawn by GarciaRodriguez and Massò on the basis of Spanish 

fishing crops (Cala Sardina Gerona) (GarciaRodriguez and Massò 1986).

Fig. 5. Summary of the papers on red coral 

in the Spanish Boletin del’Istituto Espanol de 

Oceanografia dedicated to red coral in 1986.

22
Between 1988 and 1994, after two meetings organised by FAO at Torre del Greco (1988) and Alger 

(1989), a new impulse was given to red coral research by the Italian Government, who funded 

two preliminary research projects. These researches led to two basic books on red coral biology 

and ecology (Red coral in the Mediterranean Sea, Art history and Science 1993; and Red Coral 

and other Mediterranean Octocorals: Biology and Protection 1999; Fig. 7). It will be worth to 

remember that nearly all the papers already published at that time (and nowadays too) deal with 

shallow-water populations which have a limited economic value. 

A real, positive contribution to rational management and conservation of red coral derived from 

these researches: in 1994 highly unselective, destructive fishing gears (Ingegno and St. Andrew 

Croix), which were trawled on red coral banks heavily damaging both red coral populations and 

the surrounding benthic community, were banned by the European Countries and, within a few 

years, from the whole Mediterranean Sea. Nowadays fishing is allowed by SCUBA diving and 

pick only (a quite selective fishing method!) in the whole Mediterranean.

Fig. 6. Garcia-Rodriquez and Massò 

applied the Beverton and Holt yield 

per recruit model to find the age of 

maximum yield. Their result was 80 

years, not far from the values found 

by Tsounis et al. (2007); this model 

will be discussed in paragraph 9 and 

following).

Fig. 7. Red Coral and other Mediterranean Octocorals: Biology and 

Protection, 1999.

23
Shallow and Deep-Dwelling Populations

This schematic subdivision has not any genetic implication (also if some genetic difference between 

shallow and deepest-dwelling populations has been recently found: Costantini et al. 2009), but it is 

operative, based only on some different gross features characterising red coral populations living 

at different depths. 

According to of the current literature, two main conclusions can be drawn on shallow-water 

populations:

1. The majority of papers deal with such populations living within the bathymetric range between 

20 and 50 meters (5 meters in caves);

2. These populations show some common dominant features: small colony size, small age-size at 

firs maturity (Santangelo et al. 2003; Torrens et al. 2005; Tsounis et al. 2006), high densities, 

high incidence of boring sponges (more than 50% of colonies are affected by boring sponges 

Corriero et al. 1997). It leads, together with small colony size, to a limited economic value 

of these populations. Shallow-water populations resulted to be genetically structured at a 

small spatial scale (Abbiati et al. 1993; Costantini et al. 2007). These populations show high 

reproductive output (Cattaneo-Vietti et al. 1993; Santangelo et al. 2003) and high recruitment 

rates (Bramanti et al. 2003, 2005, 2007). 

These shallow-water populations thrive at a depth that can easily be reached by scuba divers, thus 

they have been or are still now deeply harvested (Santangelo et al., Tsounis et al. in press); their 

peculiar size/age structure in which small/young colonies are dominant is the consequence of the 

overharvesting they suffered in the past, and the highest density (up to 3.000 col. m2

 ) some of 

them show could be the consequence of overharvesting (Bramanti et al. 2009). 

In recent years (late summer 1999), some shallow-water red coral populations have been affected 

by mass mortality associated to anomalous temperature increase in the Eastern Ligurian Sea 

(Calafuria-Italy; Bramanti et al. 2005), as well as Western Ligurian Sea (Calafuria), and in the 

Gulf of Lion (Marseille; Garrabou et al. 2001). Mass mortality events affecting shallow-water 

populations occurred also in Corsica and North Sardinia. Such mass mortality events (putatively 

linked to global climate change: Santangelo and Fronzoni 2008) affected the populations of several 

benthic suspension-feeders in the Ligurian Sea (Cupido et al. 2008).

Being composed by small-sized colonies, shallow-water populations have a limited economic 

value, but they are important ensuring a pool of breeders for the species. Moreover, due to their 

beauty, red coral colonies are an important attraction for touristic diving. MPAs which harbor red 

coral populations have a unique value for both conservation and recreational diving. 

The Deep-Dwelling Populations

The populations composed of larger colonies having a higher economic value (up to 20 fold higher 

than the smaller shallow ones) generally live at a deeper depth (60-150 meters); in these populations, 

colonies are more sparse and less affected by boring sponges than in the shallower populations. 

Also, in some preliminary studies recently carried out (Angelillo et al. 1999; Bavestrello et al. 

2009; Pedoni et. al. 2009), nothing is known about colony growth, population age, sexual structure, 

24
age at first maturity, reproductive output, recruitment, and natural mortality of these populations. 

It is worth to recall that Rivoire (Rivoire 1991) reported a wide mortality of unknown origin in the 

red coral populations living between 80 and 200 meter depth in the Western Ligurian Sea and in 

the Gulf of Lion (Rivoire 1991). 

One of the few papers dealing with deep-dwelling populations was published in 2008 by the 

Spanish research team working on red coral at Barcelona ICM by Rossi and German and Italian 

co-workers (Rossi et al. 2008). This study presents the first quantitative data gathered by a ROV 

survey on the spatial distribution and structure of a population extending between 50 (common 

SCUBA limits) and 230 m depth, and compared it with the shallow populations previously studied 

in the same area. The findings suggest a lower exploitation level in deep-dwelling populations at 

Cap de Creus.

Increased Knowledge on the Bathymetric Distribution Range of Red Coral and its Implications for 

Conservation and Management

Recently (Costantini et al. 2009; Taviani et al. in this volume), the bathymetric limits of red coral 

distribution have been greatly expanded: from 350 to 600-800 meter depth. As no commercial 

exploitation never occurred at this depths, there is a large area in which red coral lives still 

unexploited (Taviani et. al. in this volume).

Three different depths and exploitation levels can be thus distinguished: a shallow one (20–50 m.), 

in which red coral has been (over)harvested in the past, and in some area (like Cap de Creus, Spain) 

still nowadays, a second, deeper one (between 60–150 m) in which the main part of commercial 

harvesting occurs and a third one (from 150 to 600 m and more) in which deepest populations have 

never been harvested (until now). Such deepest populations too, if undisturbed, could represent 

a pool of sure breeders for the species. Both shallow and deepest populations must be protected.

Tools for a Sustainable Management of Red Coral Harvesting

In order to simulate population growth and exploitation, some population models have been 

applied to red coral; in particular, two different kind of models have been applied: Beverton and 

Holt model (or “yield per recruit model;” Beverton and Holt 1957; Garcia-Rodriquez and Massò 

1986; Grigg 1984, 2001; Tsounis et al. 2007; Goffredo and Lasker 2008), and life-history table 

and Leslie-Lewis matrix model (a discrete, age-structured demographic model; Abbiati et al. 

1992; Santangelo et al. 1993a; Ebert 1999; Caswell 2001; Santangelo et al. 2007; Bramanti et al. 

2009). Moreover a “rotating harvesting strategy,” on which basis exploitation in different countries 

should be regulated according to colony growth rate (biomass), was proposed by Caddy (Caddy 

1989, 1993). The objective of Caddy’s management proposal was to define harvesting strategies 

which take into account the time intervals since the resource was last harvested and the time the 

resource needs to reach the optimum yield. Caddy calculated, on the basis of the Beverton and Holt 

model, the best yield in biomass (the best colony age for harvesting), integrating it in a “rotating 

harvesting plan” at a Mediterranean spatial scale. The limits of rotating harvesting are due to the 

difficulty to apply such a model at a Mediterranean level, coordinating fishing enforcement for 

many years (10–15) in several different countries and to the characteristics of the Beverton and 

Holt models, as will be explained in the following paragraph. A rotating harvesting strategy could 

25
be successfully applied to red coral management, but it needs to be applied at a local level to single 

populations (Abbiati et al. in this volume).

Virtues and Limits of the Beverton and Holt Models

• Beverton and Holt “yield per recruit” models are based on: population biomass growth

(weight) and mortality. They aim at finding a trade-off between biomass growth of an initial 

cohort (first age class) and the loss of biomass over time due to mortality: there will be a time 

at which the difference between yield growth and mortality loss (in biomass) will reach its 

maximal value. 

• One of the main advantages is that data can be collected only once (in a single harvesting/

sampling season).

• Such “simple” models could be useful to initiate a harvest regime, but the effects of reproduction 

and density-dependence of biomass growth, which play a major role on the growth of a real 

population, are not included in the model; moreover the original model assumes that all the 

colonies above a determined (age) size will be completely removed (Goffredo and Lasker 

2008).

• Beverton and Holt models have been recently applied to the Cataluña red coral population by 

Tsounis (Tsounis et al. 2007). Tsounis applied to the model different growth rates and different 

mortality rates gathered from the literature obtaining the times of maximum yield reported 

above (Fig. 8).

New Tools for Red Coral Management and Conservation: The Demographic Approach

More recently, red coral research was addressed towards the study of population structure and 

dynamics by means of demographic models (Santangelo et al. 2004, 2007; Bramanti et al. 2009). 

This last approach, widely applied in Conservation Ecology (Dobson 1998; Ebert 1999; Fujiwara 

and Caswell 2001), will supply the tools to match harvesting to population growth rate fostering 

Fig. 8. Yield per recruit curve for the 

Cataluña C. rubrum population (Modified 

from Tsounis et al. 2007, 2009). Different 

ages of maximum yield have been found 

according to different growth and mortality 

rates of an initial cohort. A maximum yield 

after 96 years was reached when a colony 

growth rate overestimating growth, (GarciaRodriquez and Massò 1986; Marschall et al. 

2004) was applied.

26
both conservation and rational management of the populations of this precious species, if sound 

data on deep-dwelling populations will be at least gathered and enforcement will be really applied 

at local population level. The compilation of life-history tables greatly helped to foster the 

progression of species and populations towards survival; they supply the drastic data on population 

structure and demography (density, number of colonies in each size/age class, reproductive output, 

mortality). These data are then included in an algebraic matrix (Leslie-Lewis transition Matrix; 

Caswell 2001), which allows to project population trends overtime (Fig. 9).

The main advantages of these models are: 

1. Contrary to Beverton and Holt models, the contribution of each cohort to population growth 

rate by reproduction (and survival) is now included into the model. 

2. The model allows to simulate every possible variation in harvesting intensity and a selective 

harvesting affecting different age classes in a different way.

3. Demographic models too (as Beverton and Holt models) can be applied to a data set collected 

only once (in a single harvesting/sampling season, under the assumption of population steadystate).

To apply demographic models, based on Leslie-Lewis transition matrixes to deep-dwelling 

populations (those having the major economic value), it needs to be known:

• Colony growth rate 

• Population size-age structure and density

• Reproductive output and mortality rates (mortality can be gathered by the population sizeage structure, under the assumption of population steady-state)

Fig. 9. Sketch of the steps for demographic modelling. In the simulation different 

colors represent different age classes. 

27
This last approach, widely applied in Conservation Ecology, will supply the tools to match 

harvesting to population growth rate fostering both conservation and rational management of the 

populations of this precious species.

Some Conclusive Remarks

From this condensed revision of the scientific research on the Mediterranean red coral Corallium 

rubrum, some focal points can be stressed. There is no real risk of global extinction for the species, 

due to its early sexual maturity, wide bathymetric and geographic distribution, comprehensive of 

several, connected populations (Monte-Luna et al. 2007). Nevertheless, an increased frequency 

of anomalous mortality events, superimposed on overharvested shallow-water populations, could 

lead some of them to local extinction. Moreover, economic extinctions of some populations (i.e. 

overharvested populations that cannot be further harvested for a long while) have happened and 

will happen in the future.

Corallium fishing in the Mediterranean and in the Pacific are completely different. Contrary to 

Pacific species, which are harvested mainly by trawling, the Mediterranean Corallium rubrum

is harvested only by SCUBA diving and by pick. As pick-harvesting could be highly selective, 

culling of larger-older colonies within the populations can be planned. Recent development of 

demographic models for age-structured populations supply the tolls to plan sustainable harvesting 

matching exploitation to population growth rates (Grigg 2001; and in this volume).

Rotating harvesting strategies, suggested in the past by Caddy (Caddy in FAO 1998) could be 

successfully applied at regional or national scale.

Shallow-water populations have been and are overharvested. The Cap de Creus shallow population 

has been and is nowadays heavily harvested; moreover, harvesting rates recently increased after 

the proposal of inclusion of red coral in CITES Appendix-II was known by poachers. These 

populations, of limited economic value, should be protected. Similarly, the deepest red coral 

populations, recently discovered, should remain unharvested.

Deep-dwelling populations (those living between 60 and 150 meter depth), having a higher 

economic value, should be harvested following selective exploitation plans based on sound 

demographic data. As there is no knowledge on the demographic features of these populations 

(i.e. colony growth rate, age structure, reproduction and reproductive output, genetic structure), 

research efforts must be addressed towards such populations and common research projects must 

be fostered and funded at an European and Mediterranean level.

A demographic approach, based on life tables and transition matrices, could supply the tools to 

match harvesting to population growth rate fostering both conservation and rational management 

of this precious species, if sound data on deep-dwelling populations will be at least gathered and 

enforcement will be really applied at local population level.

28
References

Abbiati, M., Buffoni, B., Caforio, G., DiCola, G., Santangelo, G. 1992. Harvesting, predation and 

competition effects on a red coral population. Netherlands Journal of Sea Research 30: 219-

228. 

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Mar. Ecol. Prog. Ser. 95: 

245-250.

Angiolillo, M., Canese, S., Salvati, E., Giusti, M., Cardinali, A., Bo, M., Greco, S. 2009. Presence of 

Corallium rubrum assemblages below 50 m along the Calabrian coasts. In: UNEP-MAPRAC/SPA 2009. Proceedings of the 1st symposium on conservation of the coralligenous 

bio-concretions (Tabarka 16-19 January 2009). Pergent-Martini, C., Brichet (Eds.) RAC/

SPA publ. Tunis.

Bavestrello, G., Cerrano, C., Cattaneo-Vietti, R. 2009. Biological interactions affecting the 

growth rates of red coral (Corallium rubrum) colonies. In: UNEP-MAP-RAC/SPA 2009. 

Proceedings of the 1st symposium on conservation of the coralligenous bio-concretions 

(Tabarka 16-19 January 2009). Pergent-Martini, C., Brichet (Eds.) RAC/SPA publ. Tunis.

Barletta, G., Marchetti, R., Vighi, M. 1968. Ricerche sul corallo rosso IV. Ulteriori osservazioni sulla 

distribuzione del corallo rosso nel Tirreno. Rend. Sci. Ist. Lomb. 102: 119-144.

Beverton, R.J.H., Holt, S.J. 1957. On the dynamics of exploited fish populations. Fisheries 

investigation (II) London 19: 1-533.

 Bramanti, L., Magagnini, G., Santangelo, G. 2003. Settlement and recruitment: the first stages in the 

life cycle of two epibenthic suspension feeders (Corallium rubrum and Anomia ephippium). 

It. J. Zool. 70: 175-178

Bramanti, L., Magagnini, G., DeMaio, L., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean Red Coral Corallium rubrum (L 1758), a four-year study. J. 

Exp.l Mar. Biol. Ecol. 314: 69-78.

Bramanti, L., Rossi, S., Tsounis, G., Gili, J.M., Santangelo, G. 2007. Settlement and early survival 

of red coral on settlement plates: some clues for demography and restoration. Hydrobiol.

580: 219-224.

Bramanti, L., Iannelli, L., Santangelo, G. 2009. Mathematical modelling for conservation and 

management of gorgonian corals: young and olds, could they coexist? Ecological modelling 

220: 2851-2856. 

Caddy, J.F. 1989 Background concepts of for rotating harvesting strategy for the Mediterranean red 

coral resource. In: Rep. Second GFCM Tech. Consult. Red Coral in the Mediterranean. 

FAO Fish Rep. 413: 87-89.

Caddy, J.F. 1993 Background concepts of for rotating harvesting strategy with particular reference 

to the Mediterranean red coral, Corallium rubrum. In: Cicogna, F., Cattaneo, R. (Eds.) Red 

coral in the Mediterranean sea: Art, History and Science. Min. Ris. Al. Agr. For. Roma.

Caswell, H. 2001. Matrix Population Models: Construction, Analysis and Interpretation. 2nd Ed. 

Sinauer Associates, Sunderland, Massachusetts. 

29
Cattaneo-Vietti, R., Bavestrello, G., Senes, L. 1993. Red coral from the Portofino promontory. 

In: Cicogna, F., Cattaneo, R. (Eds.) Red coral in the Mediterranean sea: Art, History and 

Science. Min. Ris. Al. Agr. For. Roma.

Corriero, G., Abbiati, M., Santangelo, G. 1997. The sponge complex inhabiting a Mediterranean red 

coral population. P.S.Z.N. Marine Ecology 18(2): 147-155. 

Costantini F, Fauvelot C, Abbiati M. 2007. Fine-scale genetic structuring in Corallium rubrum: 

evidence of inbreeding and limited effects of larval dispersal. Mar. Ecol. Prog. Ser. 340: 

100-119.

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembrini, P.J., Abbiati, M. Deep-Water Corallium 

rubrum L. (1758) from the Mediterranean Sea: preliminary genetic characterization. Marine 

Ecology (in press).

Cupido, R., Cocito, S., Sgorbini, S., Bordone, A., Santangelo, G. 2008. Responses of a gorgonian 

(Paramuricea clavata) population to mortality events: recovery or loss? Aquat. Cons. Mar. 

Freshw. Ecosyst. 18: 984-992. 

Dobson, A.P. 1998. Conservation and Biodiversity. Scientific American Library. New York.

Ebert, T.A. 1999. Plant and Animal Populations. Methods in demography. Academic Press, S. Diego 

CA.

FAO. 1988. GFCM technical consultation on red coral of the Mediterranean. FAO Fisheries Report 

413, 1-159.

FAO. 2007. Capture Production 2006. Yearbook Fishery Statistics 101/1. 

Fujiwara, M., Caswell, H. 2001. Demography of endangered North Atlantic right whale. Nature 414, 

537-541.

Garcia-Rodriguez, M., Massò, C. 1986. Estudio biometrico de poblaciones de coral rojo del litoral de 

Gerona. Bol. Inst. Esp. Oceanog. 3: 65-74. 

Garrabou, J., Perez, T., Sartoretto, S., Harmelin, J.G. 2001. Mass mortality event in red coral 

(Corallium rubrum) populations in the Provence region (France, NW Mediterranean). Mar. 

Ecol. Prog. Ser. 217: 263-272.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in NW Mediterranean: insights into conservation and management needs. 

J. An. Ecol. 71: 966-978.

Goffredo, S., Lasker, H.R. 2008. An adaptive management approach to an octocoral fishery based on 

the Beverton-Holt model. Coral Reef 27: 751-761.

Grigg, R.W. 2001. Black coral: History of a sustainable fishery in Hawai’i. Pacific Sciences 53: 291-

299. 

Kock, G. Die Alcyonacea des Golfes von Neapel. Mitt. Zool. Stat. Neapel 9(4): 652-676.

Laborel, J., Vacelet, J. 1958. Etude des peuplements d’une grotte sous-marine du Golfe de Marseille. 

Bull. Inst. Océan. Monaco 1120: 11-20.

 Laborel, J., Vacelet, J. 1961. Répartition bionomique de Corallium rubrum dans les grottes et falesies 

submarines. Rapp. Réun. Comm. Int. Explor. Sci. Mer Médit. 16: 465-469.

30
Lacaze-Duthiers, H. 1864. Histoire naturelle du corail. Bailière, J.B. et fils, Paris. 137 p.

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in the precious red coral Corallium rubrum (L). Coral Reefs 23: 43-432. 

Muller, R. 1910. Ueber die Bildung des des Achsenskelets von Corallium. Mitt. Zool. Stat. Neapel, 

20: 652-676.

Pedoni, C., Follesa, M.C., Cannas, R., Matta, G., Pesci, P., Cau, A. 2009. Preliminary data on 

red coral (Corallium rubrum) population of Sardinian Sea (Western Mediterranean). In: 

UNEP-MAP-RAC/SPA 2009. Proceedings of the 1st symposium on conservation of the 

coralligenous bio-concretions (Tabarka 16-19 January 2009). Pergent-Martini, C., Brichet 

(Eds.) RAC/SPA publ. Tunis.

Rivoire, G. 1991. Mortalitè du corail et des gorgones en profondeur au large des cotes Provencales. 

In : Les Espècies Marines à protegér en Méditerrané. Bouduresque, C.F., Avon, M., Graves, 

V. (Eds.) GIS Posidonie publ. Fr. 53-59. 

Roark, E.B., Guilderson, T.P., Dumbar, R.B., Ingram, B. L. 2006. Radiocarbon-Based ages and 

growth rates of Hawaiian deep-sea corals. Mar. Ecol. Prog. Ser. 327: 1-14.

Rossi, S., Tsounis, G., Orejas, C., Padrón, T., Gili, J.M., Bramanti, L., Teixidó, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Mar. Biol. 152: 429-439.

Santangelo, G., Abbiati, M., Caforio, G. 1993a. Age structure and population dynamics in Corallium 

rubrum. In: Cicogna, F., Cattaneo, R. (Eds.) Red coral in the Mediterranean sea: Art, History 

and Science. Min. Ris. Al. Agr. For. Roma.

Santangelo, G., Abbiati, M., Giannini, G., Cicogna, F. 1993b. Red coral fishing trends in the Western 

Mediterranean Sea. Scientia Marina 57: 139-143. 

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an overexploited 

Mediterranean species. Aquat. Cons. Mar. Freshw. Ecosys. 11: 253-259.

Santangelo, G., Carletti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Mar. Ecol. Prog. 

Ser. 248: 99-108.

Santangelo, G., Maggi, E., Bramanti, L., Bongiorni, L. 2004. Conservation biology and population 

demography of a Mediterranean overexploited species. Sci. Mar. 67(6): 199-204.

Santangelo, G., Bramanti, L., Iannelli, M. 2007. Population dynamics and conservation biology of 

the over-exploited Mediterranean Red coral. J. Theor. Biol. 244: 416-423. 

Santangelo, G., Fronzoni, L. 2008. Global Climate Change and the ecology of the next decade. ETS 

Pisa. 

Santangelo, G., Bramanti, L., Vielmini, I., Iannelli, M. 2009. What we have learned about re coral 

and what we need to learn for its rational management. In: Bruckner, A.W., Roberts, G.G. 

(Eds.) Proceedings of the First International Workshop on Corallium Science, Management 

and Trade. NOAA Technical Memorandum NMFS-OPR-43 and CRCP-8, Silver Spring 

MD.

31
Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean). Biol. Cons. 121: 391-397.

Torrents, O. 2007. Biologie des population du corail rouge Corallium rubrum (L 1758) de Mèditerranée 

Nord-occidentale. PhD Thesis. Université de la Méditerranée de Marseille. 

Tsounis, G., Rossi, S., Aranguren, M., Gili, J.M., Arntz, W. 2006. Effects of spatial variability and 

colony size on the reproductive output and gonadal development cycle of the Mediterranean 

red coral (Corallium rubrum L.). Mar. Biol. 148: 513-527.

Tsounis G, Rossi S, Gili J.M., Arntz W.E. 2007. Red coral fishery in the Costa Brava (NW 

Mediterranean): case study of the overharvested precious coral. Ecosystems 10: 975-986.

Tsounis, G., Rossi, S., Grigg, R., Santangelo, G., Bramanti, L., Gili, J.M. The Exploitation and 

Conservation of Precious Corals. Oceanography and Marine Biology: An Annual Review, 

Volume 48 (in press).

Vighi M. (1972) Etude sur la reproduction du Corallium rubrum. Vie et Milieu 23: 21-32.

Weinberg S. 1978. Mediterranean octocorallian communities and the abiotic environment. Marine 

Biology 49: 41-57.

Weinberg, S. 1979a. Autoecology of shallow-water Octocorallia from Mediterranean rocky substrates. 

I. The Banyul area. Bijdr. Dierk. 49: 1-15.

Weinberg, S. 1979b. The light-dependent behaviour of planulae larvae of Eunicella cavolinii and 

Corallium rubrum. Bijdr. Dierk. 49: 16-30.

Weinberg, S. 1993. Red coral research through the centuries. In: Red coral in the Mediterranean Sea: 

History, Art and Science. Cicogna, F., Cattaneo-Vietti (Eds.) Min. Ris. Agr. Al. For. Roma.

32
IDENTIFYING POPULATION DECLINE IN CORALLIUM RUBRUM

BY USING HISTORICAL INFORMATION

Georgios Tsounis1, Sergio Rossi2, Josep-Maria Gili1

1

Departamento de Biología Marina, Institut de Ciències del Mar (CSIC)

Passeig Marítim de la Barceloneta, 37-49, 08003 Barcelona, Spain

E-mails: georgios@icm.csic.es, gili@icm.csic.es 

2

Institut de Ciència i Tecnologia Ambientals (Universitat Autònoma de Barcelona), 

Edifici Cn Campus UAB, Cerdanyola del Vallés (Barcelona) 08193, Spain

E-mail: sergio.rossi@uab.cat

Introduction

Corallium rubrum has been fished for 5000 years, and since the 1800s its fishery reached an 

industrial scale with several hundred boats dredging the Mediterranean for this valuable resource 

of the jewelry industry. Dramatic yield declines occurred in the 1980s, and the second FAO ad hoc 

Corallium rubrum consultation in 1988 concluded that the species was already overharvested (FAO 

1988; Cognetti 1989). Subsequent research confirmed not only that the yields had dropped, but 

throughout the Mediterranean the known red coral populations in diving range consisted entirely 

of much smaller colonies (ca. 3 cm on average) than the maximum size of the species (more than 

50 cm) (Garcia Rodriguez and Massó 1986; Santangelo et al. 1993; Santangelo and Abbiati 2001; 

Garrabou and Harmelin 2002; Santangelo et al. 2007; Tsounis et al. 2007).

Despite of these studies, recent efforts to improve its management and conservation were slowed 

down by discussions over the extent of the decline, as management or conservation organs like 

FAO and CITES need to quantify population decline precisely. However, in the case of precious 

corals, a precise estimate will probably never be possible, since ecologists have to deal with a 

lack of a natural baseline. In other words, we cannot know the population structure of undisturbed 

populations, because the first studies on its population status have been conducted only in recent 

decades, while fishing began five millennia ago (Tescione 1973). This makes it difficult, but not 

impossible, to identify decline and propose more effective management measures.

In order to provide the best possible estimate of population trends, we briefly summarize the most 

important historical information that is scattered among grey literature and has therefore been 

unavailable to ecosystem managers looking at scientific texts. It is the only information available 

that allows an evaluation of the early state of Corallium rubrum populations in the Mediterranean 

Sea. The purpose of this summary is therefore not to review recent scientific studies and their 

application in precious coral management. For a detailed review of precious coral fisheries please 

refer to Tsounis et al. (2010). This summary attempts to briefly present a historical frame that may 

serve to understand the extent of fishing impact on C. rubrum populations, and put the modern 

studies into context.

The Two Different Fractions of Corals Harvested

a) Dredging

33
Information from the period when dredging was still allowed is scarce, but data compiled in 

Tescione (1973) show fluctuating yields of up to several metric tons per year throughout centuries. 

Fortunately, the FAO started compiling continuous yield statistics in the 1970s, which was early 

enough to document the depletion of the coral fraction accessible to dredges: The data show a 

dramatic decline beginning in about 1980 which reached its low point before Spain prohibited 

dredging in its waters in 1986 (Cognetti 1989). Sardinia followed this example soon afterwards 

(Cognetti 1989), and in 1994 the EU banned dredging. It is important to understand that this 

decline occurred before dredging was banned or fishing effort decreased, and is therefore a strong 

indication that dredging had removed all coral that grew in exposed positions. In fact, most fisheries 

experienced a similar decline due to the industrialization after the 2nd world war, and show similar 

decline curves (Pauly 2002).

Furthermore, instead of neglecting the structure of coral habitat and its relation to the fishery, we 

can identify two different coral fractions within the populations. These fractions are determined 

by their position in their habitat, which affects their accessibility to different harvesting methods: 

Coral may grow in an exposed position on top of the substrate, or rather cryptic in crevices, cave 

entrances and overhangs. Dredges however only access the exposed part of the population. We 

postulate therefore that dredging with the “Saint Andrews Cross” and the “Barra Italiana” removed 

the vast majority of exposed coral on the rocks, down to the limit of 100 – 200 m, as evidenced 

by the above mentioned collapse. The recovery of the populations began only in the late 80s/early 

90s, so if they immediately re-colonized the free habitat, today the colonies may have reached 25% 

of the maximum size (extrapolating linear growth in relation to C. rubrum maximum lifespan of 

ca. 100 years). However, there are indications that other deep coral species grow much faster than 

red coral (Orejas et al., submitted), and may have occupied habitat freed up by dredging (Rossi et 

al. 2008).

We furthermore argue that intensive dredging over centuries may thus well be the reason that C. 

rubrum appears to grow a cryptic, sciafillic species. It is found in shady places only, but light levels 

and other abiotic factors have not been found to affect larvae behavior or survival in both, larvae 

and adults (Weinberg 1987, 1979). Historical information reports that red coral was present at only 

few meters depth (Tescione 1973), which has been confirmed in recent surveys, where (small) 

colonies were found at 7.5 m depth at Cap de Creus (Tsounis et al. 2006).

b) Scuba Diving

A second phase of the industrial fishery began in the 1950s with the invention of the Aqualung. 

SCUBA divers were able to access the second fraction of colonies, hidden in caves, overhangs 

and crevices that dredges cannot reach. Historical information from this period of coral fishing 

reveals that even a surprisingly small fishing effort allowed extremely high yield initially: The 

first SCUBA harvester was the sport diver Leonardo Fusco, who became a professional coral diver 

in 1954 after discovering red coral while free diving in Palinuro near Naples (Liverino 1983). In 

his first season he harvested 250 kg. Quickly the small circle of SCUBA pioneers in Italy (Guido 

Garibaldi, Alberto Novelli, and Ennio Falco) became coral fishermen and discovered further banks 

in Sardinia, Elba, and Corsica (Roghi 1966). The fishery evolved similarly at the Costa Brava 

(Spain): At Cap de Creus, the diver George Kontos began to fish coral in the 1950s, but also other 

red coral fishermen (e.g. the Boix brothers or Narcís Flaqué) collected huge amounts of red coral 

at the Montgrí Coast and the Medas Islands. It has been reported that in some cases, they collected 

34
all large colonies from a cave of almost 100 meters length in less than one week (Plujà, pers. com.). 

These caves contain only very few tiny C. rubrum branches today (pers. obs.).

One of the most spectacular and valuable source of C. rubrum was discovered at that time in the 

legendary Capo Caccia Cavern in Sardinia at a depth of 37 m (Liverino 1983). Liverino (1983) 

reports that in 1956 divers worked at 30 – 35 m, but in 1958 at 40 – 45 m. By 1964 an ever growing 

group of divers was working at depths of 72 m, and inevitably a long list of accidents were the 

result of the spreading ‘coral fever’ among the young divers (Liverino 1983). Leonardo Fusco 

reported that in 1955 he harvested at 60 m in the Gulf of Naples but in 1964 he had to descend 

to 90 m. Similarly, the pioneer Fausto Zoboli is reported to have said that he worked as one of 

the first at 60 m in 1964 (near Rome), while in 1971 he was forced to work at 100 m in Alghero, 

Sardinia (Liverino 1983). Others similarly documented that by the late 1950s divers in France and 

Italy already had to descend to depths of 80 m, and at times to even more than 100 m, to find coral 

(Galasso 2000). In 1974 helium based mixed gas diving techniques developed by the French ocean 

engineering company COMEX started to spread among coral divers, permitting them to work at 

120 m for 20 min without the dangers of nitrogen narcosis (Liverino 1983).

The 14 mile long Scherchi Channel from Sicily to Tunisia was regarded as a coral ‘el dorado,’ 

with 80 divers from Italy, France, and Spain harvesting 70 – 120 t in 1978, starting at 60 m and 

gradually working their way down to 130 m. This submarine canyon was described as an oasis for 

marine fauna and flora, uncontaminated and exceptionally rich in rare species. In 1979 there were 

366 boats at work (283 of them were registered in Italy) and 150 divers (Liverino 1983). It thus 

appears that the operating depth of the divers reached a limit of SCUBA in the late 1970s/early 

1980s, at about the time when the yields collapsed. We therefore conclude that the second fraction 

of corals (especially large colonies) was largely depleted down to 130 m in the early 1980s.

Furthermore, the management measures (or their degree of efficiency) that led to this situation 

are more or less still in place today, since they have not been significantly revised during the last 

decades. Thus the trend of depletion of 35 – 40 m habitats in the 1950s, then intermediate depths 

at 60 – 70 m, and today’s harvesting at 90 – 150 m depth, will continue deeper and deeper if 

management is not fundamentally revised.

The fact that enormous amounts of coral were harvested initially by a small number of divers 

implies that large colonies existed in shallow water. The extremely large annual yield of several 

hundred tons until the 1980s implies another significant conclusion in the case of such a slow 

growing species. If we extrapolate from today’s young populations with today’s relatively low 

yields, to determine what kind of populations afforded several hundred tons of annual yield for 

decades, it becomes unquestionable that vast C. rubrum “red forests” disappeared before they 

could be documented. These tons of C. rubrum were creating a habitat complexity that must have 

been comparable to coral reefs. The extremely slow growth rate of C. rubrum (Marschal et al. 

2004) lets us conclude that the populations cannot have recovered yet from such an intensive 

fishery.

Trends of Colony Size through Time 

The above conclusions are furthermore confirmed by comparisons of the size of the corals in time. 

The size of the largest coral colonies that can be found in a population is a good descriptor for 

35
the degree of the fishing pressure they are exposed to, as the fishery is size specific. Furthermore, 

historical reports on the size of corals in their habitat exist, and enable us to make comparisons: 

In the 1960s corals as large as a man’s palm could still be found in 35 m depth in Cap de Creus 

in Spain (J.G. Harmelin and A. Plujà, pers. com.), whereas today this size of colony can only be 

found in marine protected areas, and only beyond safe SCUBA limits (pers. obs.). 

In other precious coral populations, such as black coral in Hawaii and New Zealand, we know that 

a considerable percentage of old, large individuals exist in undisturbed populations (Grange 1997; 

Grigg 2001). The fraction of old colonies in a population is a function of the natural mortality (Grigg 

1976). Like most precious coral species, C. rubrum is characterized by low mortality (Garrabou 

and Harmelin 2002), and we can therefore expect large colonies in undisturbed populations. In 

fact, video footage from coral divers in Sardinia (90 m depth) confirms that large corals of more 

than 45 cm are frequent in undisturbed or moderately harvested populations (M. Scarpati, pers. 

com.). 

In contrast to these few fairly well preserved populations, surveys in Spain show that the average 

size of shallow water populations is ca. 3 cm of height, while deep populations reach 10 – 16 cm 

(Rossi et al. 2008). Most data in Italy and France originate from the Marine Protected Area in 

Calafuria, while unprotected populations are hardly studied (the difficulty of finding unprotected 

coral in diving range is the reason).

Deep populations are slightly larger, as they are recovering since 1986, but still only ca. 25% of the 

potential maximum size. Finally, analyzing the catch of poachers working in shallow water (less 

than 70 m) confirms that they are not able to find any large colonies either (Tsounis et al. 2007). 

Shallow water populations are therefore overexploited. It is alarming that their young populations 

do not appear to have the resilience to survive a combination of anthropogenic impact and natural 

catastrophes (Santangelo et al. 2007; Bruckner 2009).

Remarkably, we can deduce a considerable decline without the existence of abundance data, which 

are less useful in these species (Tsounis et al. 2010). Size structure immediately reflects overfishing, 

since coral fishing is always size specific, targeting large individuals first. Considering the biomass 

removed before the 1980s, the gradually increasing operating depths and the small colonies that 

remain in overharvested stocks, we have no doubts that this species meets any criteria for decline. 

The data imply that the biomass present today is only a minute fraction of the natural state some 

centuries ago, while the trend is continuing decline.

Implications for Management

This summary demonstrates that a century ago the Mediterranean Sea was still inhabited by red 

coral populations that can be described as underwater forests. Corals in general are regarded as 

ecosystem engineers that increase biomass and biodiversity of their ecosystem. In the case of 

deep corals, they are assumed to be fish nurseries, and management plans in the US do not need 

to prove this in order to conserve deep coral habitat (Magnuson-Stevens Fishery Conservation 

and Management Act). However, during the recent consultation-workshops the focus has been 

on single species extinction and local extinction rather than using a holistic habitat conservation 

approach. It should be stressed that the habitat function of large C. rubrum colonies needs to be 

taken into consideration. Marine protected areas in the Mediterranean are rather small and not all 

36
of them contain red coral populations. Also, the protection started only very recently, just a few 

decades ago, so that their populations are still young and recovering. 

The implication of the above for sound management starts with increased minimum size limits 

and decreased yield quotas. Above all, poaching must be stopped and yields of legal divers better 

controlled. Furthermore, rotation harvest schemes should be abandoned as, even at small scale, 

populations and patches should not be harvested in a way that completely depletes the largest size 

class (Tsounis et al. 2010). Populations consisting of only young colonies recover slower, and 

do not provide their ecological services as fish nurseries. It is known for example, that another 

Mediterranean precious coral (Gerardia sp.) is crucial to several shark species as they lay their 

eggs in its branches (Öztürk, pers. com.). Care must also be taken to not interrupt continuous 

populations, as this can interrupt the gene flow (Costantini et al. 2007). 

The above arguments lead to the conclusion that populations with only small, unbranched colonies 

should be allowed to recover, instead of continuing to be harvested. Unfortunately this is the case 

in all known shallow water populations, so that a ban of coral fishing in shallow water must be 

recommended. Industry representatives characterized today’s shallow water populations as having 

little commercial value (Ciro Condito, pers. com.).

Finding a way to implement these conclusions and install new management plans is a challenge 

however, as twenty years ago a FAO workshop already reached the same conclusions that we 

reached today: C. rubrum is overharvested, and a cross-national management plan should be 

created under the responsibility of the General Fishery Commission of the Mediterranean (GFCM) 

(Cognetti 1988). 

References

Bruckner, A.W. 2009. Rate and extent of decline in Corallium (pink and red coral) populations: are 

existing data adequate to justify a CITES Appendix II listing? Marine Ecology Progress 

Series (in press). doi: 10.3354/meps08110.

Cognetti, G. 1989. FAO Congress on Corallium rubrum. Marine Pollution Bulletin 20, 95 only

Costantini, F., Fauvelot, C., Abbiati, M. 2007. Genetic structuring of the temperate gorgonian coral 

(Corallium rubrum) across the western Mediterranean Sea revealed by microsatellites and 

nuclear sequences. Molecular Ecology 16: 5168-5182.

FAO. 1988. GFCM technical consultation on red coral of the Mediterranean. FAO Fisheries Report 

413: 1-159.

Galasso, M. 1998. Unterwasserfunde in West-Sardinien. Zeitschrift für Unterwasserarchaeologie 1: 

18-31.

García-Rodríguez, M., Massó, C. 1986a. Estudio biométrico de poblaciones de coral rojo (Corallium 

rubrum L.) del litoral de Gerona (NE de España). Boletin Instituto Español de Oceanografia 

3: 61-64.

37
Garrabou, J.M., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: Insights into conservation and management 

needs. Journal of Animal Ecology 71: 966-978.

Grange, K.R. 1997. Demography of black coral populations in Doubtful Sound, New Zealand: 

Results from a 7-year experiment.. In: Proceedings of the 6th International Conference on 

Coelenterate Biology. Leiden: National Museum of Natural History, 185-193.

Grigg, R.W. 1976. Fisheries management of precious and stony corals in Hawaii. UNIHI-SEAGRANT, 

TR-77-03. Honolulu.

Grigg, R.W. 2001. Black coral: History of a sustainable fishery in Hawaii. Pacific Science 55: 291-

299.

Liverino, B. 1983. Il Corallo – Esperienze e ricordi di un corallaro. Banca di credito populare Torre 

del Greco. Li Causi Editore. Torre del Greco, Italy.

Marschal, C., Garrabou, J. and Harmelin, J.G. 2004. A new method for measuring growth and age in 

the precious red coral Corallium rubrum (L.). Coral Reefs 23: 423-432.

Orejas, C., Ferrier-Pages, C., Reynaud, S., Tsounis, G., Alemand, D., Gili, J.M. (submitted). The 

deep-sea coral Madrepora oculata from the Mediterranean, grows at similar rates than the 

tropical coral Galaxea fascicularis. Marine Ecology Progress Series.

Pauly, D., Christensen, V., Guénette, S., Pitcher, T.J., Sumaila, U.R., Walters, C.J., Watson, R., Zeller, 

D. 2002. Towards sustainability in world fisheries. Nature 418: 689-695.

Roghi, G. 1966. La febre rossa. La strada del corallo: una storia italiana. Mondo Sommerso, 6. 

Online. Available HTTP: http://www.gianniroghi.it/ (accessed: 7 October 2009).

Rossi, S., Tsounis, G., Padrón, T., Orejas, C., Gili, J.M., Bramanti, L., Teixidor, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Marine Biology 154: 533-545. Santangelo, G., Abbiati, M., Giannini, F., 

Cicogna, F. 1993b. Red coral fishing trends in the western Mediterranean Sea during the 

period 1981-1991. Sciencia Marina 57: 139-143.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an over-exploited 

Mediterranean species. Aquatic Conservation Marine Freshwater Ecosystems, 11, 253-259.

Santangelo, G., Bramanti, L., Iannelli, M. 2007. Population dynamics and conservation biology of 

the over-exploited Mediterranean red coral. Journal of Theoretical Biology 244: 416-423.

Tescione, G. 1973. The Italians and Their Coral Fishing. Naples: Fausto Fiorentino.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W.E. 2006b. Population structure of an exploited benthic 

cnidarian: The case study of red coral (Corallium rubrum L.). Marine Biology 149: 1059-

1070.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W.E. 2007. Red coral fishery at the Costa Brava (NW 

Mediterranean): Case study of an overharvested precious coral. Ecosystems 10: 975-986.

Tsounis, G., Rossi, S., Grigg, R., Santangelo, G., Bramanti, L., Gili, J.M. 2010. The Exploitation and 

Conservation of Precious Corals. Oceanography and Marine Biology: An Annual Review, 

Volume 48 (in press).

38
Weinberg, S. 1978. Mediterranean octocoral communities and the abiotic environment. Marine 

Biology 49: 41-57.

Weinberg, S. 1979. The light dependent behaviour of planula larvae of Eunicella singularis and 

Corallium rubrum and its implication for octocorallian ecology. Bijdragen tot de Dierkunde 

49: 145-151.

39
MULTIDISCIPLINARY DATA ON THE STATUS OF 

RED CORAL (CORALLIUM RUBRUM) RESOURCE IN 

SARDINIAN SEAS (CENTRAL WESTERN MEDITERRANEAN).

Cannas R., Caocci F., Follesa M.C., Grazioli E., Pedoni C., Pesci P., Sacco F., Cau A.

Department of Animal Biology and Ecology – 

Centre of Competence in Marine Biodiversity – University of Cagliari

Viale Poetto, 1 – 09126 Cagliari - ITALY

E-mail: cau@unica.it

Abstract

In this study, data gathered from several fields are presented in order to give an historical and 

actual picture of the status of the red coral resource in Sardinian seas. A review of historical data is 

provided and the first picture of spatial distribution and genetic structure of red coral populations in 

deep waters of Sardinian seas (below 80 meters) are illustrated. The historical analysis shows that 

no variation occurred in mean yields/boats in the last four centuries, suggesting that the amount 

of red coral in Sardinian seas is not decreased with time, differently from what observed in other 

parts of the Mediterranean. Analyses carried out from 2007 to 2008, through ROV surveys and 

samplings of entire colonies, highlight that the populations show dimensions (> 80% with maximum 

height > 6 cm) able to ensure the reproductive potential needed for their survival. Finally, strong 

genetic heterogeneity was measured between Sardinian samples and corals from other western 

Mediterranean localities (e.g. Corsica and Sicily). High level of genetic differentiation was found 

even among Sardinian samples, over different spatial scales from hundreds to less than 1 km. 

In conclusion, the strong genetic structuring found, confirms the need for management plans 

and conservation programs to be locally defined. Moreover, all historical, population structure 

and genetic data coherently indicate that the harvesting effort should not be increased, and the 

actual Sardinian management plan, enacted by the regional administration, seems to be effective 

in protecting the resource from overexploitation. However, the strict respect of this regulation is 

binding to ensure an optimal balance between harvesting and reproductive rate and hence allow 

the long term persistence of the species.

Introduction

Corallium rubrum (L. 1758) is one of the components of the Mediterranean coralligenous 

community (Sarà 1973). Because of their high economic value, the red coral has long been heavily 

exploited in the Mediterranean Sea since ancient times (Morel et al. 2000; Santangelo et al. 1993; 

Santangelo and Abbiati 2001). Harvesting on an industrial scale began as early as in the Neolithic 

age (Grigg 1977). During the industrialization period the stocks declined and the catches dropped 

dramatically in 1976–1984 (FAO 1984, 1988). Nonetheless, this resource is still present in great 

quantity in the Northwestern Mediterranean basin (Torrents et al. 2005), where it is still successfully 

harvested today (FAO 1988). 

Sardinia (Central Western Mediterranean) has always been very rich in red coral banks and the 

main of these are – and were – located mainly in the Western part of the island, almost with no 

interruption from North to South. A documentation of these banks, most of which are still exploited 

40
at present, can be seen in the maps of an old paper of Parona (1883). Unofficial information 

let us know that Sardinian red corals were already exploited in the 13th century by Provençal 

fishermen (Doneddu and Fiori 2003). The first official document on red coral fishing dates back to 

the early 14th century (Zanetti 1946). It is reported that the Maritime Republic of Pisa exploited 

the Southern part of Sardinia, while Maritime Republic of Genoa the Northern one. In this times, 

Alghero, in the Northern part, is the leading town, due to its very rich and high quality coral banks. 

In 1354, Alghero was conquered by Catalans who exploited its coral banks. Later on, in the 16th 

century, Sardinian banks started to be exploited by people form Campania (Torre del Greco) who 

became later on the main harvester. 

A lot has changed with time in harvesting red coral. First of all the catch methods: while in previous 

centuries the gear used was the “ingegno,” a gear generally made by a wooden cross with some 

weight or rake and some net, at present the harvesting of the coral is made by professional coral 

fishermen who dive to find the coral banks and pick it, generally helped by a ROV. These are two 

very different way of catch coral which determine a very different effort on the resource. Only 

the latter can select the colonies to be harvested thus leaving smaller ones alive. The change of 

the catch method has been driven by management advice: since 1979, in Sardinia the harvest is 

regulated by a detailed regional law, redacted with the strict collaboration of technician, local 

administrators and fisherman (Regional law n. 59, 5 July 1979). Today, the harvesting, practiced 

only manually, is regulated annually by the imposition of a period of harvesting (generally from 

May to September/October), by the issue of a limited number of licenses and by the definition of 

size and weight limits (basal diameter < 10 mm with a tolerance of 20%; maximum harvesting 

of 2.5 kg per dive/day) (Regional law N.16/V, 22 May 2006). Since 2008, the fishery above 80 m 

depth is also forbidden (Regional law N.1409/Dec A/49, 11 June 2008). 

Generally, it is stressed that strategies to foster conservation and management need to be based 

on a sound knowledge of population demographic features and life history traits (Bramanti et 

al. 2007). Some studies on demography (Santangelo and Abbiati 2001; Garrabou and Harmelin 

2002), reproduction patterns (Vighi 1972; Santangelo et al. 2003; Torrents et al. 2005, Tsounis et 

al. 2006a), recruitment, settlement and mortality (Garrabou and Harmelin 2002; Bramanti et al 

2003, 2005, 2007), growth (Marschal et al. 2004; Bramanti et al. 2005), distribution and space 

competition (Giannini et al. 2003), feeding ecology (Tsounis et al. 2006c; Picciano and FerrierPages 2007) have contributed to know the ecology of the species essentially situated in shallow 

waters. Little is known on its distribution and population structure below 50 m depth (Tsounis et 

al. 2007), because of the logistical difficulty of working at greater depths (Genin et al. 1992). The 

understanding and knowledge of the distribution and population structure along its complete depth 

range, the magnitude and direction of the exchange of individuals among geographically separated 

populations are fundamental to understand population dynamics, to forecast how corals are likely 

to recover from disturbance events both of natural and anthropogenic origin and hence to plan a 

realistic management for a species such as Corallium rubrum (Rossi et al. 2008). 

An indirect measure of reproductive population connectivity can be obtained following a genetic 

approach. The results of early genetic studies on red coral populations, based on allozymes, indicate 

that Mediterranean C. rubrum populations are genetically heterogeneous, and provide evidence of 

some breeding isolation and population sub-structuring among samples (Abbiati et al. 1993). More 

recently, microsatellite data indicate that effective larval dispersal in red coral in the Northwestern 

Mediterranean Sea is highly restricted (Costantini et al. 2007a, 2007b).

41
The totality of the over-mentioned genetic studies examined shallow waters colonies (mainly up 

to 30 m of depth). Only a very recent paper provide the genetic characterization of 12 fragments 

from live red coral colonies collected at the exceptional depths of ca. 600–800 m: these were found 

to be slightly different, suggesting the possible occurrence of genetic isolation among shallow and 

deep-water populations (Costantini et al. 2009). However, given the exiguity of colonies analyzed 

further samplings of red coral deep colonies are needed to confirm these results. 

In this paper we reported data gathered from several fields in order to give an historical and actual 

picture of the status of the red coral resource in a part of Central western Mediterranean, Sardinian 

seas. A review of historical data (from early 1700 until now) is provided and the first picture of 

spatial distribution and genetic structure of red coral populations in deep waters of Sardinian seas 

(below 80 meters) are illustrated. 

Materials and Methods

Historical Data

To assess the evolution of the red coral resource in Sardinia from 1978 to 2005, yearly landings 

data (kg) from FAO fisheries statistics and CPUE (Catches per Unit of Effort, kg) were analyzed. 

The CPUE here was obtained as the kg of the harvested coral divided by the number of boats 

(referred to the years in which the “ingegno” was towed) or the number of divers (referred to the 

years after 1989 when only divers could harvest red coral in Sardinia). CPUE then represents the 

yearly mean of the red coral fished for unit of effort. 

Statistical trends were evaluated with the Spearman rank test (Zar 1999).

Extensive Survey

The Extensive ROV survey was carried out between 2007 and 2008 at depths below 80 meters by 

professional divers to identify locations of Corallium rubrum colonies in the Sardinian Sea and to 

identify the size structure of colonies. The main objective of this survey was in fact to compile an 

extensive set of data without causing any destruction whatsoever to the colonies themselves. 

This part of the study consisted of an exploration made by a ‘Velociraptor’ type ROV (Remotely 

Operated Vehicles). This is a scuba vehicle run by cave, equipped with an autonomous propulsion 

system and a Sony HDV colour camcorder, which lets vehicle machinist to stay outside while the 

ROV works suspended in mid-water. The vehicle can develop a maximum of 3-4 knot speed and is 

composed of an exterior control unit and an umbilical cave 250 m long made in ‘Kevlar’ fibre that 

carries current and control signals.

During the monitoring, images, gotten by ROV, were obtained in analogical format, were passed 

on VHS support by a video recorder, and then converted into digital format by the auxiliary of a 

DVD recorder. Video transect positioning and distance covered by ROV were relieved by a Global 

Positioning System (GPS). An echo-sounder, put on the boat, provided bathymetric data. 

42
A total of 33 transects were performed and 19307 colonies were examined (Table 1). In each colony 

the height was measured, using a rod of 2 cm as the reference size, from a point two centimeters 

from the base of the stem to the furthest tips of the longest branches (Tsounis et al. 2006a), with 

the image analysis program, tpsdDig 2 (Rohlf 2005). Finally all colonies were grouped in six 

classes according to height with five centimeters of difference between one group and the next (I, 

0-5 cm; II, 5-10 cm; III, 10-15 cm; IV, 15-20 cm; V, 20-25 cm; VI, > 25 cm). The data have been 

analyzed according to the coasts from near to which samples were taken (Fig. 1). These coasts are: 

the Northern coast in the areas of Santa Teresa and Castelsardo, where samples come from depths 

of between 84 and 93 meters; the Northwestern coast in the areas of Bosa and Alghero, where 

samples come from depths of from 80 to a 122 meters; the Central-Western coast in the area of 

Oristano, samples here come from one 113 to one 115 meters down; and the Southwestern coast in 

the area of Carloforte, harvested colonies from this site were brought up from depths of between 

85 and 107 meters. The data relative to each zone are compared by the G method (Zar 1999). 

Genetic Analyses

Samples were gathered from the three main islands of the Western Mediterranean basin: Sicily, 

Corse, and Sardinia. Most of the samples (n=7) were collected off Sardinian coasts in fall 2008 and 

summer 2009 from deep waters (from 84 up to 121 m of depth). Sardinian samples were harvested 

in three main areas: the Northern coast (Santa Teresa di Gallura and Castelsardo), the Central 

Western coast (Bosa and Oristano) and the Southwestern coast (Carloforte) (Fig. 1). Two further 

samples were collected: one off Corse and the other one in Sicilian waters. At each location, 

branch fragments from a min of 20 to a max of 40 individual colonies were picked by professional 

scuba divers; corals were then preserved in 90% ethanol. Total genomic DNA was extracted 

from polyps using the NucleoSpin® Tissue (Macherey-Nagel). Five microsatellite loci (COR9, 

COR15, COR46, COR48, and COR58m) were amplified following the PCR conditions described 

in Costantini and Abbiati (2006). Allele sizing was scored using the software Genemarker v1.75 

(Softgenetics).

Allelic diversity was measured through estimates of the Number of alleles (Na), the Allelic richness 

(Ra), the mean observed heterozygosity (Ho), and the unbiased expected heterozygosity (UHe) 

using the software GenAlEx v.6.2 (Smouse et al. 2008), Genepop v.4.0 (Rousset 2007) and Fstat 

2.9.3. Genepop was also used to test genotypic deviations from Hardy–Weinberg Equilibrium 

(HWE) measured as heterozygosity deficiency, both for each locus and globally (Markov chain 

1000 dememorization steps, 100 batches and 1000 iterations per batch). Significance levels 

were adjusted with sequential Bonferroni corrections for multiple tests using the program SGoF 

(Carvajal-Rodriguez et al. 2009). To determine whether null alleles were present, the FreeNa 

package (Chapuis and Estoup 2007) was used: a maximum-likelihood estimate was calculated 

for the frequency of null alleles according to Dempster et al. (1977). Differentiation of allele 

frequencies within and among samples was estimated by an AMOVA analysis and the calculation 

of a Fixation index (FST) with Arlequin version 3.1 (Excoffier et al. 2005). The significance of 

FST values were tested using a nonparametric Monte Carlo approach (100000 permutations). 

Significance levels for pairwise FST were adjusted with sequential Bonferroni corrections. The 

software Genetix (Belkir et al. 1996) was used to construct a factorial component analysis (FCA) 

to visualize the genetic distance between all genotypes. 

43
To estimate the effects of isolation-by-distance, we calculated the correlation between pairwise 

genetic differentiation (FST) and geographic distance, measured as the direct sea path between 

sampling locations, using a Mantel-test implemented in the software Isolation by Distance Web 

Service (IBDWS) http://ibdws.sdsu.edu/~ibdws/aboutibdws.html (Jensen et al. 2005). 

Results

Historical Data

The analysis of FAO data show that from 1978 a marked decline in red coral landings can be 

observed (Fig. 2). This reduction resulted to be statistically significant (Spearman rank test: 

P-value < 0.01). However it doesn’t necessarily reflect a decrease in the production: it can be often 

referable to the decrease in the number of boats and, as well, to the market demand.

The CPUE was then analyzed (Fig. 3). The greater values of CPUE are recorded in the period 

1989–1994 (with a peak in 1992: 335.7 kg by diver). On the whole, these data are more regular 

and show the clear absence of a decreasing trend (Fig. 3). The absence of a statistical trend is 

highlighted by the Spearman test (P-value = 0.784).

Extensive Survey

The population structures of Corallium rubrum appeared statistically different (G = 449 d.f = 15 P 

< 0.005) among all four investigated areas.

On the Northern coast, from the 10 ROV transects carried out in Santa Teresa at depths of between 

84 and 93 meters, populations that in the past were exposed to considerable human impact were 

identified. In this area there is in fact a near absolute dominance of small colonies, with 90% of 

the colonies falling into either one or the other of the smallest of the two height classes (I and II) 

(Tables 2-3). In particular, 49% of the colonies are in the height class of up to 5 cm, and 41% in 

the height class of from 5 to 10 cm. Our ROV survey has not revealed any colonies within this 

sector having heights in the classes of over 15 cm. In contrast, Corallium rubrum resources on the 

Northwestern coast are not just in a different state but in a better and healthier one. From the 18 

transects carried out in the Bosa area in 2007, the highest values are observed for the medium-size 

colonies comprised in the two classes of from 10 to 15 cm in height and 15 to 20 cm in height, 

with percentages of over 30%. Only 5% of registered colonies pertain to smaller-height categories 

(Class I) (Tables 2-3). On the Central Western coast the structure of C. rubrum populations is 

seen to be similar to those on the Northwestern coast. Finally, an intermediate structure between 

that of the Northern and the Northwestern coasts has been found in the Carloforte area on the 

Southwestern coast. The 11 ROV transects carried out in 2007 and 2008 at depths of between 85 

and 107 meters show that the colonies prevalently found here do not exceed 10 cm in height. Only 

17% of all colonies in this sector belong to the height class of between 10 and 15 cm. 

Genetic Analyses

A total of 185 individuals were successfully genotyped, and the 5 microsatellite loci were 

polymorphic across all samples. Estimators of genetic diversity varied through loci and populations, 

with variable levels of polymorphism in number of alleles (Na), allelic richness (Ra), observed 

44
heterozygosity (Ho), and unbiased expected heterozygosity (UHe) (Table 4). In all samples, the 

inbreeding coefficient FIS resulted to be significantly > 0, indicating that breeding among close 

relatives seems to occur, but with different levels of consanguinity in the different samples (Table 

4). Significant heterozygosity deficiency was detected for almost all populations in the HWE tests 

(Table 4). Different frequencies of null alleles were computed in the five loci (Table 4). 

Analysis of molecular variance (AMOVA) revealed that populations were genetically differentiated 

(Table 5). The AMOVA assigned 8.28% (FST = 0.083, P-value = 0) of the genetic variation to 

among-site variation, suggesting that recruitment is highly localized. This is supported by the 

indices of pairwise FST, ranging from 0.015 to 0.16, values statistically significantly > 0 (data 

not shown), and providing evidence of a genetic subdivision between almost all populations even 

when the allele frequencies corrected with FreeNa for technical artefacts (null alleles) were used. 

When all Sardinian samples were gathered in one group and compared with the other samples, 

Corse and Sicily, placed in two other distinct groups, these 3 groups were significantly differentiated 

(Table 5). When Sardinian samples were split in three groups (north, central, and south groups) 

according to their geographical origin, the 5 groups (3 Sardinian groups + Sicily + Corse) were still 

significantly different (Table 5).

The FCA plot (Fig. 4) show three well defined and separated clouds: Sardinian corals, Corsican 

and Sicilian samples. Moreover, focusing on the Sardinian samples, three clusters, corresponding 

to colonies from the Northern, Central Western, and Southwestern coast are distinguishable. The 

genetic structuring found on geographical basis with the previous analysis is further confirmed 

by the Mantel test (r = 0.6087, p = 0.0060, Fig. 5) showing a significant correlation for the deep 

colonies between genetic and geographical distance. 

Discussion and Conclusions

This study employs a multidisciplinary approach to define the status of red coral populations off 

the Sardinian coasts, an area that harbors the majority of the commercial banks still existent in the 

Mediterranean Sea. In fact, according to the FAO data, for the last 20 years Sardinia results to be 

the main producer of red coral in Mediterranean Sea; the annual yield jumped from 35% in 1983 

to more than 95% of the total Italian yield in 1991. Since 2000, the Sardinian production reached 

about the 99% of the overall Italian yield of C. rubrum.

Historical landings data, population structure analysis, and genetic surveys were performed in 

order to gather useful information for a sustainable management of this precious resource.

As concerns the historical data, the analysis of both landings data and CPUE (Catches per Unit of 

Effort, kg) was performed. In fact, the use of landings data alone is neither adequate nor correct to 

assess a resource like red coral, strongly dependent on market demand. Variation in landings, in 

fact, cannot be referred only to a decrease in the population amount but also to a number of events 

not directly related to the increase in fishing effort. This can be the case when a crash in the market 

determines a reduction in the red coral demand. 

This is an important statement, since landings data variation does not always reflect a decrease in 

the population amount. The CPUE is a better approach to study the red coral resource. It in fact 

45
gives an average of the amount of the red coral caught by single boats or diver, independently from 

how many boats were used to fish it and so also independently from how intense is the marked 

demand. 

In particular, in Sardinia no statistical decrease of CPUE was observed in the analyzed years and 

the values obtained from 1978 to present are comparable. This should indicate that the resource 

has not been depleted and that, the regional laws, enacted from 1978, seem to work very well for 

a sustainable exploitation of the red coral. 

Concerning the population structure analysis, the results indicate that, in general, C. rubrum 

resources present a “good ecological condition” in the Sardinian Sea. Only the North coast resources 

seem to have been more affected by human impact. It is for this very reason that the area has been 

closed to harvesting since 2007 (Regional law N. 16/V, 22 May 2006). In all the other areas we 

have investigated, the colonies show a mean height higher than 6 cm, a size defined by Tsounis 

et al. (2006a) as the measure at which the colonies reached the 100% of fertility and therefore 

able to ensure the reproductive potential needed for the survival of the populations. Even in the 

Northwestern coast the considerable density of adult colonies does not allow the settlement of 

new larvae so much that a higher harvesting pressure of commercial colonies could be permitted. 

Whereas size structure of C. rubrum resources is the key to the detecting of the species’ response 

to environmental and anthropogenic factors and forms the basis for management decisions, from 

the gathered data we can definitely confirm that harvesting pressure has not to be increased in the 

Sardinian Sea. The actual regional law, which is based on three main measures —time of harvesting 

number of licenses and a harvest per dive/day — could in fact ensure sustainable management. 

Although, its strict respect should be binding to ensure an optimal balance between capacity of 

recovery and the harvesting.

Finally, this study utilized high resolution genetic markers, microsatellites loci, to indirectly 

measure reproductive population connectivity among populations of Mediterranean red coral 

Corallium rubrum. Extensive data on the genetic variability of the Mediterranean red coral deep 

colonies are presented here for the first time, contributing to fill a crucial knowledge gap.

The first finding of this study is that heterozygote deficits recorded in all populations; this situation 

was already reported for C. rubrum populations in other Mediterranean areas (Costantini et al. 

2007a, 2007b) and it is commonly found in several other species of brooding and broadcasting 

hard corals and in marine species in general (Underwood et al. 2007, and reference therein). 

Non-random mating within sites, a product of restricted dispersal of larvae, could be invoked as 

an explanation and may have contributed to the heterozygote deficiencies observed here. In fact, 

positive and significant FIS values were measured, indicating that breeding among close relatives 

seems to occur in red corals, but at different levels in the different locations, that is samples are 

characterized by different levels of consanguinity. 

Furthermore, a limited mixing among all populations is evident as pairwise Fst values were 

significant, even for sites separated by less than a kilometer (e.g. Oristano113 and Oristano115 

are separated by < 100 meters). Consequently, the interpretation is that dispersal over the scale of 

kilometers to tens of kilometers is sparse among deep coral populations. 

46
AMOVA analyses confirmed a high genetic heterogeneity among red corals across the Western 

Mediterranean basin. When Sardinian samples are split in three groups (Northern, Central Western 

and Southwestern groups), these are genetically differentiated and all are different from Corse and 

Sicily samples (Table 5); the Mantel test confirms that the more distant are the samples the more 

divergent they are (Fig. 5). 

All of these results together permit to gather useful information for management and conservation 

of this important resource. This study highlight the substantial genetic differentiation of red coral 

populations on the basis of their geographical, with limited connectivity and gene flow among the 

different areas and it gives strong support for locally defined management units and conservation 

plans strategies. 

47
Table 1. Number of transects and examined colonies for each investigated area.

Investigated Area n° transects n° examined colonies

Nothern coast 9 11795

Northwestern coast 10 1423

Central Western coast 3 1496

Southwestern coast 11 4593

Total 33 19307

Table 2. Initial and final coordinates and relative main depth of ROV transect conducted in each zone.

Zone Initial position Final position Depth (m)

Northern coast of the transect of the transect

001COR07 41°17.182’ 009°01.448’ 41°17.253’ 009°01.465’ 90

002COR07 41°17.059’ 009°02.143’ 41°17.170’ 009°02.130’ 84

003COR07 41°14.225’ 009°05.105’ 41°14.100’ 009°03.182’ 90

004COR07 41°14.206’ 009°05.105’ 41°14.312’ 009°05.196’ 89

005COR07 41°13.547’ 008°57.173’ 41°13.637’ 008°57.003’ 90

006COR07 41°13.158’ 009°07.481’ 41°13.236’ 009°07.371’ 90

007COR07 41°11.240’ 008°55.181’ 41°11.128’ 008°55.096’ 93

008COR07 41°10.429’ 008°57.006’ 41°10.530’ 008°56.935’ 90

010COR07 41°10.048’ 008°55.106’ 41°09.955’ 008°55.212’ 89

Northwestern coast

001COR07 40°24.117’ 008°15.302’ 40°24.235’ 008°17.420’ 121

002COR07 40°23.151’ 008°14.395’ 40°23.299’ 008°14.235’ 102

003COR07 40°22.424’ 008°14.523’ 40°22.330’ 008°14.490’ 85

007COR07 40°22.175’ 008°12.205’ 40°22.098’ 008°12.312’ 113

008COR07 40°21.185’ 008°13.445’ 40°21.101’ 008°13.568’ 109

009COR07 40°20.565’ 008°14.535’ 40°20.472’ 008°14.612’ 114

010COR07 40°20.335’ 008°13.295’ 40°20.240’ 008°13.365’ 80

013COR07 40°20.315’ 008°13.395’ 40°20.431’ 008°13.256’ 119

014COR07 40°19.581’ 008°15.488’ 40°19.421’ 008°15.370’ 90

018COR07 40°19.275’ 008°15.268’ 40°19.169’ 008°15.180’ 113

Central Western coast

01COR08A 40°05.553’ 008°15.000’ 40°05.467’ 008°14.921’ 94

01COR08B 40°05.325’ 008°14.365’ 40°05.241’ 008°14.451’ 97

02COR08 40°05.286’ 008°14.524’ 40°05.374’ 008°14.676’ 106

Southwestern coast

015COR07 39°18.224’ 008°19.220’ 39°18.136’ 008°19.105’ 104

016COR07 39°07.592’ 008°10.030’ 39°07.431’ 008°09.899’ 87

03SAN08 39°07.432’ 008°10.181’ 39°07.345’ 008°10.274’ 89

04SAN08 39°03.257’ 008°12.255’ 39°03.364’ 008°12.331’ 92

05SAN08B 39°09.323’ 008°07.979’ 39°09.213’ 008°08.087’ 107

05SAN08A 38°57.374’ 008°15.851’ 38°57.487’ 008°15.943’ 106

06-07SAN08 38°58.292’ 008°18.243’ 38°58.383’ 008°18.141’ 113

08SAN08 38°56.084’ 008°16.415’ 38°56.172’ 008°16.326’ 105

09SAN08 38°54.270’ 008°18.040’ 38°54.369’ 008°18.129’ 105

10SAN08 38°51.074’ 008°22.077’ 38°51.164’ 008°22.163’ 90

11SAN08 38°50.569’ 008°22.044’ 38°50.456’ 008°22.157’ 85

48
Table 3. Percentage (%) of the high classes of colonies harvested in each investigated coast.

Height Classes

Northern coast 0_5 5_10 10_15 15_20 20_25 > 25

001COR07 54 46 0 0 0 0

002COR07 66 24 7 0 3 0

003COR07 25 50 25 0 0 0

004COR07 48 36 16 0 0 0

005COR07 69 26 4 0 0 0

006COR07 29 71 0 0 0 0

007COR07 44 33 22 0 0 0

008COR07 35 53 12 0 0 0

010COR07 71 29 0 0 0 0

mean value 49% 41% 10% 0% 0% 0%

Northwestern coast 0_5 5_10 10_15 15_20 20_25 > 25

001COR07 10 24 29 28 8 0

002COR07 3 16 22 35 23 0

003COR07 4 19 42 32 3 0

007COR07 0 9 36 40 15 0

008COR07 1 9 52 34 3 0

009COR07 0 10 54 29 6 0

010COR07 1 16 28 41 13 1

013COR07 13 22 31 25 8 0

014COR07 7 35 29 21 7 0

018COR07 9 29 39 18 5 0

mean value 5% 19% 36% 31% 9% 0%

Central Western coast 0_5 5_10 10_15 15_20 20_25 > 25

01COR08A 4 14 41 39 2 0

01COR08B 1 14 44 40 1 0

02COR08 4 11 47 37 1 0

mean value 3% 13% 44% 39% 1% 0%

Southwestern coast 0_5 5_10 10_15 15_20 20_25 > 25

015COR07 13 33 37 15 2 0

016COR07 18 36 33 11 1 0

03SAN08 65 26 10 0 0 0

04SAN08 58 34 8 0 0 0

05SAN08B 39 42 16 2 0 0

05SAN08A 40 42 16 2 0 0

06-07SAN08 46 38 14 2 0 0

08SAN08 48 41 11 0 0 0

09SAN08 44 35 19 2 0 0

10SAN08 73 15 12 0 0 0

11SAN08 40 37 7 12 5 0

mean value 44% 34% 17% 4% 1% 0%

49
Table 4. Genetic population statistics.

Population Carloforte Oristano113 Oristano115 Bosa90 Bosa120 Castelsardo SantaTeresa Corse

Calvì

Sicily

BancoTerribile

Depth 85 113 115 90 121 84 93 90 90

N 18 20 14 20 20 30 33 14 16

Na 7.2 9.4 6.8 7.2 8 11.4 11 6.6 7.4

Ra 6.13 7.60 6.08 6.11 6.34 7.78 7.40 5.97 6.63

UHe 0.75 0.82 0.74 0.75 0.76 0.79 0.79 0.73 0.80

Ho 0.49 0.42 0.47 0.57 0.58 0.44 0.53 0.59 0.42

Fis 0.36* 0.31* 0.22* 0.45* 0.39* 0.45* 0.34* 0.2* 0.49*

Nu 0.15 0.15 0.12 0.21 0.20 0.21 0.17 0.12 0.20

n = number of samples; Na = number of alleles; Ra = allelic richness; UHe = unbiased expected heterozygosity; Ho = 

observed heterozygosity; Fis = Weir and Cockerham inbreeding coefficient; Nu = frequency of null alleles, according 

to Dempster et al. (1977). * indicates deviations from HWE, P < 0.001.

Table 5. Results from the AMOVA showing the partitioning of genetic variation among and within groups and 

individuals.

Percentage of variation Fixation Indices P-value

1 group (all populations)

Among populations 8.28% Fst 0.083 0.00+0.00

3 Groups (Sardinia/Sicily/Corse)

Among groups 6.78% Fct 0.068 0.024+0.0015

Among populations within groups 5.70% Fsc 0.061 0.00+0.00

Within populations 87.52% Fst 0.12 0.00+0.00

5 Groups (Sardinia N/Sardinia CW/Sardinia SW/Sicily/Corse)

Among groups 4.21% Fct 0.042 0.0097±0.00098

Among populations within groups 4.85% Fsc 0.05 0.00±0.00

Within populations 90.94% Fst 0.0905 0.00±0.00

50
Fig. 1. Map of the investigated areas in Sardinian seas during the extensive ROV survey and genetic analyses.

51
Fig. 3. Catch per Unit of Effort data for the whole of Sardinia (1978-2005).

Fig. 2. Sardinian landings data for the red coral, Corallium rubrum, from 1978 to 2005 (FAO sources).

52
Fig. 4. FCA plot displaying the genetic distance between the deep colonies depicted as the centre of gravity of all 

sampled colonies within a sampled site.

Fig. 5. Genetic differentiation [computed as FST/(1 - FST)] versus the logarithm of the geographic distance (in km) 

for deep colonies of C. rubrum.

53
References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Marine Ecology Progress 

Series 95: 245-250.

Belkir, K., Borosa, P., Goudete, J., Chiki, L., Bonhomme, F. Genetix, Logiciel sous WindowsTM 

pour la genetique des populations. Laboratoire Genome, Populations, Interaction. CNRS 

IMR 5000. Montpellier: Universite Montpellier II; 1996.

Bramanti, L., Magagnini, G., De Maiob, L., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean red coral Corallium rubrum (L 1758), a 4-year study. J Exp 

Mar Biol Ecol 314: 69-78

Bramanti, L., Magagnini, G., Santangelo, G. 2003. Settlement and recruitment: the first stages 

in the life cycle of two epibenthic suspension feeders (Corallium rubrum and Anomia 

ephippium) Ital J Zool 70: 175-178.

Bramanti, L., Rossi, S., Tsounis, G., Gili, J.M., Santangelo, G. 2007. Settlement and early survival 

of red coral on artificial substrates in different geographic areas: some clues for demography 

and restoration. Hydrobiologia 580: 219-224.

Carvajal-Rodriguez, A., de Una-Alvarez, J., Rolan-Alvarez, E. 2009. A new multitest correction 

(SGoF) that increases its statistical power when increasing the number of tests. BMC 

Bioinformatics 10: 209.

Chapuis, M.P., Estoup, A. 2007. Microsatellite Null Alleles and Estimation of Population 

Differentiation. Molecular Biology and Evolution 24(3): 621-631.

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembri, P.J., Abbiati, M. 2009. Deepwater Corallium rubrum (L., 1758) from the Mediterranean Sea: preliminary genetic 

characterization. Marine ecology doi:10.1111/j.1439-0485.2009.00333.x

Costantini, F., Abbiati, M. 2006. Development of microsatellite markers for the Mediterranean 

gorgonian coral Corallium rubrum. Molecular Ecology Notes 6: 521-523.

Costantini, F., Fauvelot, C., Abbiati, M. 2007a. Fine-scale genetic structuring in Corallium rubrum 

(L.): evidences of inbreeding and limited effective larval dispersal. Marine Ecology 

Progress Series 340: 109-119. 

Costantini, F., Tinti, F., Abbiati, M. 2003. Sistematica molecolare e filogenesi di Corallium rubrum. 

Biologia Marina Mediterranea 10: 73-75.

54
Costantini, F., Fauvelot, C., Abbiati, M. 2007b. Genetic structuring of the temperate gorgonian 

coral (Corallium rubrum) across the Western Mediterranean Sea revealed by microsatellites 

and nuclear sequences. Molecular Ecology 16: 5168-5182.

Doneddu, G., Fiori, A. 2003. La pesca in Italia tra età moderna e contemporanea. Produzione, 

mercato, consumo. Editrice Democratica Sarda.

Dempster, A.P., Laird, N.M., Rubin, D.B. 1977. Maximum Likelihood from Incomplete Data via 

the EM Algorithm. Journal of the Royal Statistical Society Series B (Methodological) 

39(1), 1-38.

Excoffier, L., Laval, G., Schneider, S. 2005. Arlequin ver. 3.0: an integrated software package for 

population genetics data analysis. Evolutionary Bioinformatics Online 1, 47-50. 

Food and Agriculture Organization (FAO). 1984. Technical consultation on red coral resources of 

the western Mediterranean. FAO Fish Rep 306: 1-142.

Food and Agriculture Organization (FAO). 1988. GFCM technical consultation on red coral 

resources of the Mediterranean. FAO Fish Rep 413: 1-159.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: insights into conservation and management 

needs. J Anim Ecol 71: 966-978.

Genin, A., Paull, C.J., Dillon, W.P. 1992. Anomalous abundances of deep-sea fauna on a rocky 

bottom exposed to strong currents. Deep-sea Res 39: 293-302.

Giannini, F., Gili, J.M., Santangelo, G. 2003. Relationships between the spatial distribution of 

red coral Corallium rubrum and coexisting suspension feeders at Medas Islands Marine 

Protected Areas (Spain). Ital J Zool 70: 233-239.

Grigg, R.W. 1977. Population dynamics of two gorgonians coral. Ecology 58: 278-290.

Jensen, J.L., Bohonak, A.J., Kelley, S.T. 2005. Isolation by distance, web service. BMC Genetics 

6: 13.

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in the precious red coral Corallium rubrum (L.). Coral Reefs 23: 423-432.

Morel, J.P., Rondi-Costanzo, C., Ugolini, D. 2000. Corallo di Ieri – Corallo di Oggi. Atti Del 

Convegno, Ravello, Villa Rufolo, 13–15 Dicembre 1996 Centro Universitario Europeo Per 

I Beni Culturali, Ravello. Edipuglia, Bari.

Parona, C. 1883. Il corallo in Sardegna: relazione presentata a S. E. il Ministro di Agricoltura, 

Industria e Commercio. Edited by Tipografia Eredi Botta.

55
Peakall, R., Smouse, P.E. 2006. GENALEX 6: genetic analysis in Excel. Population genetic 

software for teaching and research. Molecular Ecology Notes 6: 288-295.

Picciano, M., Ferrier-Pagès, C. 2007. Ingestion of pico- and nano-plankton in the Mediterranean 

red coral Corallium rubrum. Mar Biol 150: 773-782.

Rholf, F.J. 2005. TpsDig, digitize landmarks and outlines, version 2.04. Department of Ecology 

and Evolution, State University of New York at Stony Brook.

Rossi, S., Tsounis, G., Orejas, C., Padrón, T., Gili, J.M., Bramanti, L., Teixidó, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Mar Biol 154: 533-545.

Rousset, F. 2007. Genepop ‘007: a complete re-implementation of the Genepop software for 

Windows and Linux. Molecular Ecology Notes 8(1): 103-106.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an over-exploited 

Mediterranean species. Aquatic Conserv Mar Freshw Ecosyst 11: 253-259.

Santangelo, G., Abbiati, M., Caforio, G. 1993. Red coral fishing trends in the western Mediterranean 

Sea during the period 1981-1991. Sci Mar 57: 139-143.

Santangelo, G., Carletti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Mar Ecol Prog 

Ser 248: 99-108.

Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean). Biol Conserv 121: 391-397.

Tsounis, G., Rossi, S., Aranguren, M., Gili, J., Arntz, W.E. 2006a. Effects of spatial variability and 

colony size on the reproductive output and gonadal development cycle of the Mediterranean 

red coral (Corallium rubrum L.) Mar Biol 148: 513-527.

Tsounis, G., Rossi, S., Laudien, J., Bramanti, L., Fernández, N., Gili, J., Arntz, W.E. 2006c. Diet 

and seasonal prey capture rates in the Mediterranean red coral (Corallium rubrum L.). Mar 

Biol 149: 313-325.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W.E. 2007. Red coral fishery at the Costa Brava (NW 

Mediterranean): Case study of an over harvested precious coral. Ecosystems 10: 975-986.

Underwood, J.N., Smith, L.D., Van Oppen, M.J.H., Gilmour, J.P. 2007. Multiple scales of genetic 

connectivity in a brooding coral on isolated reefs following catastrophic bleaching. 

Molecular Ecology 16: 771-784.

56
van Oppen, M.J.H., Lutz. A., De’ath, G., Peplow, L., Kininmonth, S. 2008. Genetic traces of recent 

long-distance dispersal in a predominantly self-recruiting coral. PLoS ONE 3(10): e3401. 

doi:10.1371/journal.pone.0003401.

Vighi, M. 1972. Etude de la reproduction du Corallium rubrum (L.). Vie Milieu 23: 21-32.

Zanetti, G. 1946. La legislazione sarda relativa all’industria corallina e la pesca del corallo in 

Sardegna. Edited by Gallizzi. 

Zar, J.H. 1999. Biostatistical analysis. Prentice Hall International, Third Edition. 633 p. Abbiati, 

M., Novelli, S., Harmelin, J.G., Santangelo, G. 1997. Struttura genetica di popolamenti 

simpatrici e allopatrici di corallo rosso. In: Biologia e Tutela del Corallo Rosso e di Altri 

Ottocoralli del Mediterraneo, Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. (Eds.) 5-21. 

Ministero Risorse Agricole Alimentari Forestali, Roma.

57
ASSESSING DECLINE IN CORALLIUM POPULATIONS: 

DO EXISTING DATA MEET THE CRITERIA

 FOR A CITES APPENDIX II LISTING?

Andrew Bruckner1, 2

1

NOAA Fisheries, Office of Habitat Conservation, Silver Spring MD, USA

2

Khaled bin Sultan Living Oceans Foundation, Landover, MD USA

Abstract

In June 2007, the U.S. proposed Corallium (pink and red corals) for listing on Appendix-II of 

the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). 

The proposal was adopted and later overturned due to perceived difficulties in implementing and 

enforcing a CITES listing and uncertainties on population status. An expert review (FAO 2007) 

questioned whether populations had declined to 20-30% of a historic baseline, the level required 

for a CITES Appendix-II listing. Although landings declined by over 80% during the 1990s as 

compared to quantities reported from 1965-1989, this review concluded that this was not a measure 

of resource decline, but rather declines in the wholesale value of Corallium which rendered fishing 

uneconomical. The review also used colony abundance and density as surrogates of decline, which 

may be high (200-1300 colonies/m2

) in the Mediterranean. Assessments of decline in sessile, 

colonial organisms should consider reductions in size, since reproductive output and survival 

increase exponentially as colonies get larger. Colonies of C. rubrum historically achieved heights 

of 50 cm, developing complex first, second, and third order branching patterns. Today, over 90% 

of colonies in fished areas are 3-5 cm tall, more than half are sexually immature and most have 

only rudimentary branches. Few population data are available for Pacific Corallium spp.; however, 

landings over the last 15 years have dropped precipitously from 100-400 metric tons (t) to < 

5 t/year. Recovery of populations to their natural state may require up to a century or possibly 

longer, as colonies protected from fishing for 20-30 years are still less than half their historic size. 

Minimum allowable size for harvest should be increased because traditionally-determined growth 

rates appear to underestimate colony age, and corals are being removed long before achieving 

adequate levels of reproductive output and maximum sustainable yield (MSY). “Boom to bust” 

cycles of Corallium fisheries and dramatic, long-lasting shifts in population demography highlight 

the need for improved management and trade regulations.

Introduction 

Precious corals have been highly valued since ancient times for use in the fabrication of jewelry, 

amulets, and art objects (Tescione 1973). Precious coral refers to a group of taxonomically distinct 

corals that includes gold corals (Gerardia spp.) in the order Zoanthidae, black corals in the order 

Antipatharia, and two groups in the order Gorgonacea, bamboo corals (Acanella and Lepidisis 

spp.), and pink and red corals (Corallium and Paracorallium spp., family Coralliidae, referred to 

as “Corallium”). Pink and red corals are the most valuable precious corals in commerce and among 

the rarest (Grigg 1989). Because of their high value, these resources have been intensely fished 

in both the Pacific and Mediterranean, with a well established pattern of discovery, exploitation 

and depletion (Grigg 1989). International trade, consisting of millions of items and thousands 

of kg of raw and processed specimens each year, has contributed to serial depletions of most 

58
known populations of Corallium, and new stocks have been rapidly exhausted shortly after their 

discovery. In addition to localized depletions, the use of coral dredges has damaged Corallium

habitats (Chessa and Cudoni 1988). Throughout most of the Mediterranean, the complex, threedimensional “forest-like” structure has been transformed to low-relief “grass plain-like” habitats 

(García-Rodríguez and Massò 1986; Rossi et al. 2008), reducing their role as important habitat for 

other species. 

Because of concerns of overfishing and habitat damage associated with the use of non-selective 

coral dredges, a non-governmental organization petitioned the United States (U.S.) in 2006 to 

propose Corallium for listing on the Appendices of the Convention on International Trade in 

Endangered Species of Wild Fauna and Flora (CITES). After a detailed review and consultation 

with Corallium range states, the U.S. submitted a proposal to list all species of Corallium on 

Appendix-II of CITES at the 14th meeting of the Conference of the Parties. This proposal identified 

seven species as under considerable threat from harvest and international trade (C. rubrum, C. 

secundum, C. lauuense (C. regale), P. japonicum, C. elatius, C. konojoi, and C. sp. nov.) with 

other species recommended for listing because of their similarity in appearance (Wijnstekers 1988; 

Bruckner 2009). The listing proposal was adopted in committee and overturned on the final day 

of the conference, largely due to perceived challenges in implementing and enforcing this CITES 

listing, and the conclusion that existing data did not demonstrate declines to the level required 

to justify an Appendix-II listing (Morell 2007). In 2009, the U.S. was petitioned once again to 

propose Corallium for listing in CITES. An evaluation of new data was undertaken to determine 

whether the listing was warranted, and in October 2009 the U.S. and the European Union jointly 

sponsored a CITES Appendix-II listing proposal (Anonymous 2009).

The main purpose of this paper was to evaluate existing landings data and biological information 

for the family Coralliidae to determine if the species in commercial trade meet the biological criteria 

of decline needed to support a CITES listing. In order for a species to qualify for a CITES listing, 

there are a number of criteria that need to be met. First, there has to be evidence that the species is 

in international trade, the trade at existing levels is threatening the sustainability of the species and 

its role in the ecosystem, and regulation of trade is necessary to ensure that harvest does not reduce 

wild populations to levels at which survival is threatened by continued harvest or other factors. 

Second, to qualify for Appendix-II there must be evidence that demonstrates a historical extent of 

decline and recent rates of decline that have resulted in losses of 70-95% of the virgin population, 

depending on the productivity of a species. For a medium productivity species, the species should 

have declined to 10-15% of some baseline while a low productivity species would need to have 

declined to 15-30% of its historic abundance. In addition to information on the life history traits, 

which are used to determine whether it is low, medium, or high productivity, relevant information 

is needed on the size of the population and its distribution, and how the species biology may affect 

the recovery potential of populations. Proposals must also include a quantitative assessment of the 

status and trends of populations and an estimate of the level of decline, along with an explanation 

of uncertainties associated with available data (FAO 2007). 

Productivity of Corallium

In a review of the 2007 CITES Appendix-II listing proposal for Corallium (FAO 2007), four 

parameters were identified that can be used to infer productivity. Because of extremely low rates 

of natural mortality, a long lifespan, and delayed age of reproduction, most species in the family 

59
Coralliidae were considered low productivity. Corallium rubrum also exhibits a long lifespan and 

low rates of natural mortality (Grigg 1976); however, colonies can reproduce at a small size, 

which has been equated to a young age (3-7 years) in previous studies (Santangelo et al. 2003), 

suggesting it is a medium productivity species. Nevertheless, this needs to be reevaluated based on 

new information. Recent aging studies have shown the corals to be 2.6-4.5X older than previous 

estimates determined by counting growth rings (Garrabou and Harmelin 2002; Marschal et al. 

2004; Roark et al. 2006), and small colonies do not exhibit 100% reproductive maturity until 

colonies achieve a minimum height of approximately 6 cm, which may equate to an age of 10 

years or more (Tsounis 2005; Marschal et al. 2004). Several other unique reproductive traits further 

support the conclusion that C. rubrum is a low productivity species including: 1) a relatively small 

number of larvae produced per colony per reproductive event as compared to other corals, and 2) 

a single discrete annual reproductive period. This is very different from other corals that exhibit a 

brooding life history strategy, which tend to reproduce monthly on a lunar cycle (Richmond and 

Jokiel 1984; Szmant 1986). Recruitment in C. rubrum also occurs in pulses with no successful 

settlement in some years; mortality of up to 95% of the larvae occurs before settlement (Tsounis 

2005), and 66-70% of the new recruits die over the first four years of life (Bramanti et al. 2005). 

These data suggest all species in the family Coralliidae are low productivity species.

Distribution of Corallium

The family Coralliidae is widely distributed throughout tropical, subtropical, and temperate oceans, 

but individual species are generally found at very low abundances within their distributional range. 

Populations that contain economically valuable species, in abundances large enough to support 

commercial fisheries, have been identified only in three locations north of 19 degrees latitude. This 

included certain locations in the western Mediterranean, Pacific populations off the Philippines, 

Japan and Taiwan, and Hawaii and the adjacent Emperor Seamounts, most of which are now 

overexploited. Individual species are also generally restricted to relatively small areas of suitable 

habitat within their distributional range, where they form discrete, genetically distinct populations 

that are separated from other populations by large expanses of unsuitable soft bottom habitat 

(Santangelo and Abbiati 2001; Santangelo et al. 2004; Baco and Shank 2005). For instance, two 

commercially valuable species have been identified in U.S. waters around Hawaii in 16 different 

locations (Baco and Shank 2005). Of these, only three populations are considered large enough 

to support commercial harvest, and the total size, number of colonies, and estimated MSY for 

two of these beds is very low (Grigg 2002). The largest of these, Makapuu Bed off Oahu, Hawaii 

occupies an area of 4.3 km2

; this bed supports C. secundum at a density of 0.3 colonies/m2 with 

a total population size estimated at 120,000 colonies in 2001. The total number of colonies in all 

other characterized beds in the U.S. are also low including an estimated 2500 legal-sized colonies 

of C. regale at Cross Seamount and up to 7000 legal-sized at Keahole Point Bed. Available data 

suggest individual species have small effective population sizes and a restricted distribution within 

a larger geographic range. 

Evidence of Decline

There are five main sources of data available for the family Coralliidae that provide an indication 

of the dramatic declines in population abundance and structure over the last several decades. The 

typical “boom to bust” cycles of Corallium fisheries has been highlighted in numerous publications 

for both Pacific and Mediterranean locations dating back to at least the 1800s; these reports highlight 

60
the discovery of new beds, rapid increases in the number of fishing vessels and fishery workers, 

and abrupt termination of fisheries in these areas once the quality and quantity of Corallium

resources declined (Tescione 1973; Cicogna and Cattaneo-Vietti 1993). Landings data have been 

compiled since the early 1960s for Pacific reported by Japanese and Taiwanese fishermen, and for 

twelve countries in the Mediterranean (FAO 2008; Fujioka 2008). Detailed quantitative biological 

data on the density, abundance, and size structure of colonies are available for populations of C. 

rubrum off Spain, France, and Italy, including 1) recent data from shallow fished populations, 

2) historical information from similar locations, 3) depth related differences, and 4) data from 

inside and outside of MPAs (Garcia-Rodriguez and Massò 1986; Tsounis 2005; Rossi et al. 2008). 

Additional anecdotal reports are also available from other locations in the Mediterranean. In the 

Pacific, one population has been studied in detail off Hawaii, and more recently surveys have been 

undertaken off the Emperor Seamounts in international waters, and off Japan (Fisheries Agency of 

Japan 2008). There is also information describing the effects of a decline in size on the degree of 

branching and the total number of individuals (polyps) per coral, and how this affects population 

viability (Bruckner 2009).

Pacific Landings Data

Available landings data for Pacific Corallium fisheries from the last 40 years (1965-2007) illustrate 

the characteristic boom to bust cycle of Coralliidae fisheries, where both effort and landings increase 

dramatically after discovery of a new precious coral bed, followed by sharp declines a few years 

later (FAO 2008; Fujioka 2008). Pacific Corallium fisheries emerged in international waters in 1965, 

following the discovery of a large bed of C. konojoi on Koko Seamount at 300-500 m depth. Over 

the next 25 years, most Corallium from the Pacific came from Koko Seamount and other areas in the 

Emperor Seamount chain, with five major peaks in landings reported by Japan and China (Taiwan); 

this was followed by rapid decline and minimal landings in international waters since 1989 (Fig. 2). 

Landings reported by Japan exceeded 375 tons in 1966, sharply declined to 100 tons by 1968, and 

reached a second peak of 225 tons in 1969 before declining once more to less than 50 tons in 1972 

(Fujioka 2008). Annual yield from Japan was the greatest from 1965-1969 and 1979-1984 (57-91 t/

yr), with 70-90% of the harvest in later years consisting of Midway deep-sea coral. Japan’s reported 

landings declined to 2.6 t in 1987 and have remained at < 5 t/yr over the last 20 years, consisting 

mostly of C. elatius (mean = 1.9 t/yr) and C. japonicum (1.2 t/yr), and minimal amounts of C. konojoi

and Corallium sp. nov. (0.22 t/yr). With exception of the early years of the fishery, landings reported 

by Taiwan were 2-3 times greater than that reported by Japan, with peaks in landings of C. kokojoi in 

1969 (112 t), 1976 (102 t), 1981 (270 t) and 1984 (226 t), and even higher amounts of Corallium sp. 

nov. from 1983-1986 (564 t), most of which was harvested off Midway Island. Landings by Taiwan 

remained at low levels (< 5 t) over the next 20 years with exception of two periods, 1996 (12 t) and 

2002-2004 (35 t); these landings consisted of C. elatius harvested from 300-500m between Taiwan 

and the Philippines. 

Landings data for the Pacific illustrate a typical trend that has repeated itself throughout the history 

of the fishery worldwide, although no new large commercially viable beds have been discovered in 

the Pacific since 1990 and landings have remained at levels that are 10-20% of that reported in the 

late 1980s (Carleton and Philipson 1987; Grigg 1989; FAO 2008). Changes in landings have been 

attributed to a decline in the resource as well as a decline in the wholesale value of the product, 

both which could render fisheries uneconomical. There are two periods when the wholesale value 

of C. konojoi is reported to have declined to levels that made fishing uneconomical (1967 and 1971-

61
1972), yet fishing resumed a few years later and landings subsequently increased (Fujioka 2008). 

By 1988, C. konojoi beds were reported to be overexploited (Fujioka 2008), which is supported 

by landings data, and remained at less than 3 tons per year over the last two decades (FAO 2008). 

Markets were also reported to be flooded with low quality Midway deep-sea coral in 1981-1982, 

which caused the price of Corallium to decline to less than US$100/kg (Grigg 2002). According 

to Kosuge (2007) Japan and Taiwan abandoned fisheries in the central Pacific due to high costs of 

harvesting coral in deep water and increased landings of low quality coral, but presumably not due 

to a strong decline in abundance or biomass. Interestingly, landings data of Midway deep sea coral 

does not show this trend, as this species was not reported in FAO data until 1982; instead C. konojoi

showed a decline during this period, and subsequently both landings of C. konojoi and Midway deep 

sea coral dramatically increased between 1983-1987. Furthermore, wholesale prices of Corallium

rapidly increased to unprecedented highs in the late 1980s and 1990s, yet landings since 1989 have 

remained at historic low levels. If abundant coral resources still remained off the Emperor Seamount 

Chain, landings should contain living, large old growth colonies, rather than only low quality coral 

as reported by Kosuge (2007); declines in the density and abundance of the coral would also trigger 

increases in the amount of effort needed to harvest the same amount of coral, also driving fisheries 

costs up. Even though the two taxa known only from the Emperor Seamounts still show up in FAO 

data, and Taiwanese coral draggers have been observed fishing for coral in this location (Fishery 

Agency of Japan 2008), production over the last two decades is less than 1% of the yield in the 1980s 

(FAO 2008). 

While landings data alone do not provide information on effort, numerous reports are available that 

highlight the increase in fishing effort (number of vessels) following discovery, and declines in effort 

once beds were overexploited. By combining landings data with these fishery reports, conclusions 

can be made on how the distribution of fishing effort has changed over time. Corallium konojoi and 

midway deep coral were both landed around the Emperor Seamounts and they made up the majority 

of the landings from the mid 1960s to the end of the 1980s. In contrast, two Pacific species, C. 

japonicum and C. elatius, are harvested only off the islands of Taiwan and Japan and are not known 

to occur around Emperor Seamounts. These two species make up the bulk of landings from Pacific 

waters today. Submersible surveys conducted by Japan in 2008 on Emperor Seamounts included 

areas that were targeted by coral draggers in the 1960s-1980s. In several of these they identified 

isolated live colonies and dead and broken colonies, but they failed to identify a single large patch 

of Corallium that could support a coral fishery, which further demonstrates that the resource was 

overexploited beyond limits of recovery in the 1980s and 20 years of greatly reduced fishing pressure 

has been inadequate to rebuild populations (Fisheries Agency of Japan 2008). All of these factors 

clearly demonstrate a dramatic and long term collapse of populations located in international waters 

around the Emperor Seamounts, with little recovery after 20 years.

Mediterranean Landings Data

In the Mediterranean, 12 countries report landings data to FAO (Fig. 4). These data show a sharp 

decline in landings over 20 years, from a maximum of 98 tons in 1978 to 20 tons in 1998, primarily 

due to a reduction of landings from European waters. This can be partially attributed to a phaseout of the use of non selective dredges, a reduction overall in effort, and a change in management 

schemes. However, these changes are not the only reason for the decline. SCUBA fisheries were 

first introduced in the 1950s, while the use of the coral dredge was not banned until 1989 (Sardinia 

only) or 1994 (throughout the rest of European waters). Landings were already less than 50% of 

62
1978 levels (42 tons) in 1984, and they remained at these low levels (32-48 tons) for the next five 

years, which was prior to the ban on the use of ingegno. Even in Italy, where the coral dredge 

was banned five years earlier than in other locations, landings had already declined from a peak 

of over 72 tons in 1978 to 40 tons in 1980 and 19.3 tons in 1985, with < 10 tons landed each year 

up to 1988. Even though landings in Italy following the ban were about half the amount prior to 

the ban, this reduction is minimal when compared to changes observed during years when the 

ingegno was still legally used (Fig. 4b). Landings reported by individual countries have continued 

to show sharp peaks and declines since inception of SCUBA fishing, which are suggestive of the 

discovery of large aggregations of coral in a particular area, followed by rapid overexploitation of 

these populations. 

The Italian coral fishery, especially around the Sardinia region, was the first to begin to phase out 

the use of the coral dredge, while SCUBA fishing progressively increased. SCUBA fishing was 

originally concentrated in shallow water (to about 50 m), extracting corals from areas that were 

largely inaccessible to dredges. Over the last two decades, SCUBA fishing has been progressively 

moving into deeper areas, in response to a depletion of corals in shallow water, as shallow areas 

today contain mostly small corals below the legal minimum size. Individual peaks in landings 

during a single year may also reflect the pulse fishing mode associated with SCUBA harvest, where 

individual beds are selectively cleared of large colonies, then a new area is targeted. This trend 

is not restricted to a single location, as shallow water populations are considered overexploited 

throughout the Mediterranean today. 

Biological Data 

There are numerous gaps in our understanding of the biology of Corallium, their population 

status, and levels of sustainable harvest. Even though industrial scale fisheries are reported to 

have undergone periods of intensive harvest followed by years with little harvest since at least 

the early 1800s, there is no information on what populations looked like prior to the inception of 

the coral dredge, and there are no known beds of Corallium that were never fished with detailed 

population assessments. With exception of a single location off Hawaii, comprehensive deep water 

surveys (below SCUBA depths) have been undertaken only in the last several years. Field data 

are available from Spain and France both show differences in size between fished areas and areas 

that have been protected from fishing for 15-20 years in shallow water locations, and these provide 

strong evidence that population dynamics have been altered from their original state largely due to 

the removal of the largest colonies first, followed by sequential elimination of medium and small 

corals. Even in Makapuu Bed off Hawaii, where the fishery has operated intermittently for a few 

years, the largest size classes were still underrepresented after several decades without fishing 

(Grigg 2001). In addition, the first surveys in this location were undertaken after a short period 

of fishing, so it is unclear what the virgin population looked like. The observation that corals can 

achieve a larger size in absence of fishing in shallow water, and the documented presence of larger 

colonies at depths accessible to SCUBA in the 1960s, demonstrates that the corals can achieve a 

large size in shallow water, and the current size observed above 50 m depth throughout the region 

is unnatural, except perhaps in some very shallow marginal habitats such as that reported from 

Liverno, Italy, and around cave entrances where there is not enough space for the colonies to grow 

to a larger size. Furthermore, there is one quantitative study from Spain in 2008 (Rossi et al. 2008) 

and recent work off Sardinia in deep water (e.g. 80-130 m or deeper) that identified colonies of a 

much larger size.

63
There are reported differences in growth rates, possibly due to different environmental conditions 

and other factors, and there are also differences in the density of corals reported from different 

areas. This includes a notable higher density of corals in shallow areas around caves, overhangs and 

in crevices, while corals are more widely spaced and larger in deep water. Some of the differences 

in density may be due to differences in sampling approach. For example, many of the high density 

estimates reported in the literature may reflect individual patches of Corallium that are small and 

are widely dispersed over a much larger amount of available habitat at that depth. Furthermore, the 

high density estimates are often based on the total number of colonies in a patch, including new 

recruits, juveniles, and adults determined by scraping all colonies in a patch or through the use of 

macro photography, while many of the small colonies may not be recorded using other approaches. 

While higher densities would be expected in areas with small corals, because they often lack 

complex branching patterns, whereas fewer large, highly branched colonies could occupy the same 

area. For these reasons, density is an inappropriate measure of decline, and in fact, high density 

populations dominated by small corals are likely to represent an altered state due to fishing pressure, 

or in shallow areas, a marginal environment where there is a rapid turnover of populations. 

Size Structure

The most appropriate measure of decline for long-lived colonial animals with a branching 

morphology is a shift in size towards smaller corals, because larger corals contain more reproductive 

polyps, they contribute more to future generations, and they are less vulnerable to total colony 

mortality from natural causes (but more vulnerable to fishing pressure). Significant differences in 

the size of Corallium rubrum colonies are reported between locations and depths, and most areas 

exhibited a general trend of declining size structure over the last four decades. Colonies collected 

in the 1950s and 1960s were frequently up to 50 cm height and 2 cm diameter. A 1962 collection 

off Costa Brava, Spain, consisted of colonies with basal diameter of 10-45 mm (mean = 16 mm) 

and height of 10-50 cm (mean = 11.5 cm), with the largest corals estimated at 50-80 years in 

age (Garrabou and Harmelin 2002; Tsounis 2005). Colonies sampled off Costa Brava were still 

substantially larger in 1986 than that observed today, with 85% of the colonies having a basal 

diameter > 5 mm. Shallow populations from Costa Brava (10-25 m depth) are now skewed towards 

small corals (mean height = 3.1 cm, max = 20 cm; mean diameter = 4-5 mm), most of which are 

young (< 8 years old), and only 35% have basal diameters > 5 mm. Colony height increases with 

depth to 4 cm height at 25-50 m and 6 cm at 50-90 m; the largest colonies (13-16 cm height) are 

found in non-harvested areas below 50 m depth. Colonies in an adjacent protected area (Medas 

Island, Spain), closed to fishing for 15 years, were 20-50% larger (mean = 4 cm height, max = 20 

cm) than colonies at similar depths in fished areas off Costa Brava (Tsounis 2005, 2006 a,b, 2007; 

Rossi et al. 2008). Populations examined in 1998 in fished areas off France (Riou archipelago) 

were larger (mean height = 6 cm; range = 4-13 cm) than that reported for Spain (although small 

colonies were not examined in this study). These colonies were also only about half the size of a 

similar habitat in France protected from fishing since 1983 (mean = 11.8 cm, range = 8-17 cm) 

(Garrabou and Harmelin 2002; Torrents et al. 2005).

In terms of decline, the shift in maximum size observed in areas targeted by fishermen from up to 

50 cm height to colonies that are no more than 15-20 cm, and a shift in mean size from about 10 cm 

to 3 cm represents a 60-80% decline in size. The effect of these size reductions on the total number 

of polyps per colony, which is a direct measure of the numbers of reproductive modules, represents 

64
an even greater loss because this reduction is also associated with a loss of branches. Although a 

large colony has fewer polyps at the base as it gets older, the presence of many more branches still 

equates to a much larger number of polyps. For instance, a 3 cm tall coral with 3 branches may 

have up to about 100 polyps, of which only a portion are reproductively mature, while a 30 cm tall 

colony could have 10,000 polyps, all of which may be reproductively mature. Smaller colonies 

also expend more energy on growth, while larger corals may produce more offspring per polyp 

because they need to expend less energy on growth and maintenance. 

Conclusion

Most articles published on Corallium over the last two decades contain statements that the resource 

is overexploited, including papers published over 20 years ago. Numerous recommendations 

for improved management have also been proposed, many of which have never been adopted 

(e.g. increased minimum size of harvest). These corals have been proposed for listing in CITES 

on several occasions, but these proposals have not been supported in part due to 1) the limited 

availability of population data to confirm the depleted status of the species, 2) the conclusion that 

landings data cannot be used as a surrogate of decline, and 3) an incorrect assessment of population 

structure using density and abundance rather than changes in size structure. 

There have been relatively few quantitative population studies undertaken on different species 

of Corallium, but the combination of all available data on the biology, population dynamics, and 

landings, as well as historical reports on the fishery, illustrate the dire status of these species as a 

result of unsustainable fishing. It is unlikely that these species will go extinct, but extirpations of 

individual populations are very likely unless more sustainable management approaches are adopted 

and enforced. Furthermore, an Appendix-II listing is not intended to protect species threatened 

with extinction, but rather to regulate trade so that the species does not become threatened in the 

future due to unsustainable trade. 

Landings data must be examined with caution, as they do not directly provide an indication of 

effort, and they are indicative of decline in some, but not all cases. There are several factors that 

can render fisheries uneconomical, including the glut of Corallium reported during the 1980s. 

Nevertheless, the sustained reduction of landings once market prices recovered and exceeded 

previous values, without a concurrent increase in landings, provides a strong indication of decline. 

Furthermore, the presence of an increasing amount of dead and bioeroded colonies in landings 

from deep water using coral dredges are indicative of a collapse of the resource. 

Available population data for C. rubrum illustrate changes in size associated with fishing and the 

long time required for recovery. The original population structure throughout the Mediterranean 

has been altered by fishing pressure, with declines in both the reported landings and in population 

structure that exceeds the level required for an Appendix-II listing. In shallow fished areas, 

population dynamics have been altered from low density populations that cover large areas and 

are structured primarily by larger highly branched colonies (10-15 cm up to 30-50 cm), to small, 

dense patches with colonies that are substantially smaller (3-5 cm, rarely over 10 cm) and have 

few branches (Garrabou and Harmelin 2002; Tsounis et al. 2006 a,b, 2007; Santangelo et al. 2003, 

2007). All known shallow water (< 50 m depth) populations that are accessible to conventional 

(air) SCUBA exhibit these trends; this includes certain shallow environments that exhibit high 

rates of turnover due to unstable environmental conditions, high levels of biotic disturbances 

65
(overgrowth by sponges), and lack of suitable habitat where they could grow larger, but these may 

never have supported commercial fisheries (Garrabou et al. 2001; Santangelo et al. 2003). There 

is evidence that colonies in shallow areas targeted by SCUBA fisheries can grow larger if they are 

left undisturbed (e.g. in MPAs), but recovery to pre 1980s levels takes many decades. In addition 

to their smaller size, a large proportion are immature and reproductively mature colonies only 

contain tens to hundreds of reproductive modules (polyps) versus tens of thousands of polyps in 

larger colonies that formerly dominated these habitats (Garrabou and Harmelin 2002; Torrents et 

al. 2005; Tsounis 2005, Tsounis et al. 2006, 2007 a,b). These shallow populations are more likely 

to collapse under continued fishing pressure when compounded by other disturbances because of 

the low numbers of larvae produced by individual corals (GFCM 1984, 1988; Santangelo et al. 

1993,2003; Galasso 2000; Linares et al. 2003; Tsounis 2005). 

Pacific populations appear to be in even worse shape. The low reported landings from international 

waters around Emperor Seamounts today (e.g. < 1% of landings in the 1960s-1980s) and a failure 

to identify large, viable beds during recent surveys off Emperor Seamounts suggest the populations 

have been overexploited and recovery has not occurred after 20 years of greatly reduced fishing 

pressure. Recent studies from Japan identified densities that are 10-100 times less (0.05-0.005 

colonies/m2

) than reported in an unfished population off Hawaii (0.5-1 colony/m2). Landings by 

Taiwanese fishermen consist of 80% or more dead colonies, which is indicative of areas previously 

damaged and over-exploited by trawling that contain few remaining live colonies. 

All of the factors presented here are indicative of population declines that exceed the 70-90% required 

for an Appendix-II listing.

Acknowledgements

I would like to acknowledge the support of the National Oceanic and Atmospheric Administration, 

Office of Habitat Conservation, and NOAA’s Coral Reef Conservation Program for financial support 

and logistical support from the U.S. Fish and Wildlife Service. I am also indebted to a number 

of scientists, government officials, industry representatives, and non-government organizations 

that assisted in the development and refinement of this manuscript. This is the Khaled bin Sultan 

Living Oceans Foundation publication #37. 

References

Anonymous. 2007. Proposal 21. Convention on International Trade in Endangered Species of Wild 

Fauna and Flora. Consideration of Proposals for Amendment of Appendices I and II. 14th

Meeting of the Conference of Parties, Hague, Netherlands. 22 p.

Anonymous. 2009. Proposal 21. Convention on International Trade in Endangered Species of Wild 

Fauna and Flora. Consideration of Proposals for Amendment of Appendices I and II. 15th

Meeting of the Conference of Parties, Doha Qatar. 35 p.

Bruckner, A.W. 2009. Rate and extent of decline in Corallium (pink and red coral) populations: 

existing data meet the requirements for a CITES Appendix-II listing. Marine Ecology 

Progress Series (in press).

66
Carleton and Philipson. 1987. Report on a study of the marketing and processing of precious coral 

products in Taiwan, Japan, and Hawaii. South Pacific Forum Fisheries Agency, FFA Report, 

87/13, 72 p.

Chessa, L.A., Cudoni, S. 1988. Red coral, Corallium rubrum (L.), fishing in the “Bocche di 

Bonnifacio” (Northern Sardinia, Italy) FAO Fish Rep 413: 113-123.

FAO. 2007. Report of the second FAO ad hoc expert advisory panel for the assessment of proposals 

to amend Appendices I and II of CITES concerning commercially-exploited aquatic species. 

Rome, 26-30 March 2007. FAO Fish Rep. 833. 133 p. 

FAO. 2008. FishStatPlus datasets. Total fishery production 1950-2006. Release date March 2008. 

www.fao.org/fishery/statistics/software/fishstat/en.

Fujioka, Y. 2008. Information on Coral Fisheries in the Emperor Seamount Area. Appendix M in: 

Fisheries Agency of Japan, December 2008. Report on Identification of Vulnerable Marine 

Ecosystems in the Emperor Seamount and Northern Hawaiian Ridge in the Northwest Pacific 

Ocean and Assessment of Impacts Caused by Bottom Fishing Activities on such Vulnerable 

Marine Ecosystems or Marine Species as well as Conservation and Management Measures 

to Prevent Significant Adverse Impacts (Bottom Trawl). http://www.jfa.maff.go.jp/j/study/

pdf/t_e/pdf.

García-Rodríguez, M., Massó, C. 1986a. Estudio biométrico de poblaciones de coral rojo (Corallium 

rubrum L.) del litoral de Gerona (NE de España). Bol Inst Esp Oceanogr 3: 61-64.

Galasso, M. 2000. Pesca del Corallium rubrum in Sardegna nell’antichita attraverso l’indagine 

archeologica, cartografica e l rilevamenti in mare. In l’Africa romana Atti del XIV Convegno 

di Studi Sassari 7-10 Diciembre 2000.

Garrabou, J., Perez, T., Sartoretto, S., Harmelin, G. 2001. Mass Mortality Event in Red Coral 

Corallium rubrum Populations in the Provence Region (France, NW Mediterranean). Mar 

Ecol Prog Ser 17: 263-272.

Garrabou, J., Harmelin, J.G. 2002. A 20-year Study on Life-History Traits of a Harvested Long-Lived 

Temperate Coral in the NW Mediterranean: Insights into Conservation and Management 

Needs. J Anim Ecol 71: 966-978.

GFCM. 1984. Technical consultation on red coral resources in the Western Mediterranean. FAO 

Fisheries Report 306, Roma 142 p.

GFCM. 1988. Report of the second GCFM technical consultation on red coral in the Mediterranean. 

FAO Fisheries Report 413, Roma 162 p.

Grigg, R.W. 1976. Fishery management of precious and stony corals in Hawaii. Sea Grant Technical 

Report UNIHI-Seagrant-TR-77-03. 48 p.

Grigg, R.W. 1989. Precious coral fisheries of the Pacific and Mediterranean. In: Marine Invertebrate 

Fisheries: their Assessment and Management. Caddy, J.F. (Ed.) New York: John Wiley, 

636-645.

67
Fisheries Agency of Japan. 2008. Report on Identification of Vulnerable Marine Ecosystems in the 

Emperor Seamount and Northern Hawaiian Ridge in the Northwest Pacific Ocean and 

Assessment of Impacts Caused by Bottom Fishing Activities on such Vulnerable Marine 

Ecosystems or Marine Species as well as Conservation and Management Measures to 

Prevent Significant Adverse Impacts (Bottom Trawl). http://www.jfa.maff.go.jp/j/study/

pdf/t_e/pdf.

Kosuge, S. 2007. Situation about deep sea coral fisheries in the central pacific. Bull Inst Malacology 

3: 10.

Linares, C., Diaz, D., Hereu, B., Zabala, M. 2003. Avaluació de ala poblacio de corall Corallium 

rubrum de les illes Medes. Exercici 2003. Seguiment temporal de l’area marina protegida 

de les illes Medes. Infomre annual. Any 2003. Departament de Medi Ambient, Generalitat 

de Catalunya, Barcelona 56-66.

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in precious red coral Corallium rubrum (L.). Coral Reefs 23: 423-432.

Morell, V. 2007. Corals: suffering from whiplash. Science 316: 1678.

Richmond, R.H., Jokiel, P.L. 1984. Lunar periodicity in larva release in the reef coral Pocillopora 

damicornis at Enewetak and Hawaii. Bull Mar Sci 34: 280-287.

Roark, E.B., Guilderson, T.P., Dunbar, R.B., Ingram, B.L. 2006. Radiocarbon-based ages and growth 

rates of Hawaiian deep-sea coral. Mar Ecol Prog Ser 327: 1-14.

Rossi, S., G. Tsounis, T. Padrón, C. Orejas, J.M. Gili, L. Bramanti, N. Teixidor, J. Gutt. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Mar Biol 154: 533-545.

Santangelo, G., Carlietti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Mar Ecol Prog 

Ser 248: 99-108.

Santangelo G, Bramanti L, Iannelli M (2007) Population dynamics and conservation biology of the 

overexploited Mediterranean red coral. J Theor Biol 244: 416-423. 

Szmant, A.M. 1986. Reproductive ecology of Caribbean coral reefs. Coral Reefs 5:4 3-53.

Tescione, G. 1973. The Italians and Their Coral Fishing. Fausto Fiorentino, Naples.

Torrents, O., Garrabou, J., Marschal, C., Hamelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean. Biol Conserv 121: 391-397.

Tsounis, G. 2005. Demography, reproductive biology and trophic ecology of red coral (Corallium 

rubrum L.) at the Costa Brava (NW Mediterranean): ecological data as a tool for management. 

Reports of Polar and Marine Science 512. Alfred Wegener Institute for Polar and Marine 

Research, Bremerhaven.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2006a. Population structure of an exploited benthic 

cnidarian: the case study of red coral (Corallium rubrum L.). Mar Biol 149: 1059-1070.

68
Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2006b. Effects of spatial variability and colony size 

on the reproductive output and gonadal development cycle of the Mediterranean red coral 

(Corallium rubrum L.). Mar Biol 148: 513-527.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2007. Red coral fishery at the Costa Brava (NW 

Mediterranean): case study for an overharvested precious coral. Ecosystems 10: 975-986.

Wijnstekers, W. 1988. The evolution of CITES. Secretariat of the Convention on International Trade 

in Endangered Species of Wild Fauna and Flora. Lausanne, Switzerland 270 p.

69
Fig. 1. Pooled landings for all species 

of Corallium between 1965-2007. 

Landings data for the Pacific are 

shown in blue and Mediterranean C. 

rubrum landings are shown in red.

Fig. 2. Landings data presented by 

species between 1965-2007. 

Fig. 3. Landings data for Pacific 

species of Corallium from 1990-

2007. The data are the same as in Fig. 

2, although a different scale is used to 

illustrate variations in landings over 

this period.

70
Fig. 4 Landings data for C. rubrum between 1978-2007. A. Total landings pooled for all countries 

in Europe (diamonds) and Africa (squares). B. Landings for Italy, Spain, France, and Tunisia. The 

inset shows the same countries over the period 1986-2007 using a different scale to illustrate the 

variation between years. C. Landings for Morocco, Algeria, Greece, and Croatia between 1983-

2007.

71
MANAGEMENT OF PRECIOUS CORALS: 

LOCAL SCIENCE AND ENFORCEMENT VERSUS A LISTING BY CITES 

Richard W. Grigg

Department of Oceanography

University of Hawaii

Honolulu, Hawaii

 

Introduction and History of Precious Coral Fisheries

Precious corals have been treasured by man since the dawn of recorded history. Today, many 

species of precious coral exist and have been harvested in a variety of countries using a variety of 

fishing methods. The main fisheries have targeted red, pink, and white species of the Gorgonian 

Genus Corallium, black corals in the Genera Antipathes and Cirrhipathes, and gold corals in two 

distinct Anthozoan Families and Genera, Gerardia and Primnoa.

Historically, the most important species is Corallium rubrum, the famous Red Coral of Commerce, 

found throughout the Mediterranean Sea and nearby sites in the Atlantic Ocean. The first record of 

human use of red coral was the discovery of polished and perforated beads of C. rubrum recovered 

from grave sites in Germany that date back to Paleolithic Time 25,000 years ago (Tescione 1965). 

Back then, red coral appeared to have had value for trade, perhaps even as an early form of money, 

but it also seemed to have a great cultural significance. Man’s early fascination with red coral is 

exemplified in Greek Mythology. The myth of the Greek God Perseus and the Gorgon Medusa 

is famous even today in art, culture, and religion. Perseus was claimed to have slain the hideous 

dragon and cast her head into the sea, whereupon its petrifying stare turned soft algae to stone and 

stained it red – giving birth to red coral. At once, red coral became a symbol of rebirth, immortality, 

and protection perhaps from the Evil Eye and misfortune in general. Red coral became a panacea, 

an amulet, a symbol of life’s vitality. What is even more remarkable, red coral and some other 

precious corals have held this sacred fascination in the mind of man (and women) for literally 

1000s of years even up to modern time.

The cultural history of man’s love affair with red coral is important, and so are the histories of various 

precious coral fisheries, but the man focus of this paper is the question of optimum management. 

Many management strategies are available and many have been used but unfortunately almost all 

have been unsuccessful in producing a sustainable yield of the resource. Even so, it must be said 

that precious corals are biologically renewable resources. They are not minerals and achieving 

a sustainable yield is simply a matter of utilizing a management strategy that works. But before 

discussing this problem, it is first important to examine the natural history of various species of 

precious coral. It is also important to briefly describe the history of major precious coral fisheries 

in the world that have seriously depleted the stocks versus those that have not. Finally, the potential 

impacts of listing Corallium spp. on Appendix-II of CITES is considered.

Natural History of Precious Corals

In general, many precious corals share a number of similarities in life history phenomena (see 

papers by Lacaze-Duthiers 1864; Kishinouye 1904; Hickson 1924; Vighi 1970; Grigg 1976,1984, 

72
2002; Bruckner 2008; Tsounis et al., in press). To begin with, precious corals are primarily deep 

water corals living below the euphotic zone (zone of photosynthesis). All species of precious coral 

are ahermatypic, lacking symbiotic algae and none form reefs. Universally, precious corals form 

solitary colonies. Almost all are slow growing and probably have deterministic growth slowing 

down as they get older. Also, most precious corals have low fecundity and low rates of recruitment 

(except C. rubrum) and natural mortality, and consequently most are relatively long lived, living 

up to several centuries or more (Roark et al. 2006). 

What little is known about their reproductive biology suggests that most are gonochoric (separate 

sexes, also dioecious) and most spawn annually usually during warm summer months. Reproductive 

maturity may require many years depending on the longevity of a given species. Precious coral 

larvae tend to be negatively phototaxic and positively geotaxic, and most are lecithotrophic and 

have short-lived dispersal capabilities. This may account for another characteristic of precious 

corals and that is their ubiquitous patchy distribution, although this is also related to narrow habitat 

preferences. For example, almost all precious corals are found in areas of strong bottom current. 

Most precious corals are filter feeders or suspension feeders and thereby rely on strong bottom 

currents for their food supply (particulate organic matter or microplankton). Many precious corals 

in fact have a planar branching pattern and are oriented at right angles to the prevailing currents. 

Strong current is also important in maintaining substrata free of sediments which is another 

ecological requirement of precious corals; a firm or solid bottom texture on which to settle. Because 

precious corals are moderately long lived, this means that habitats in which they proliferate must 

be free of sediment (sand, mud, ash, etc.) for long periods of time. These kinds of habitats are often 

found in channels or off points or capes of land where bottom currents may be intensified due to 

Bernoulli effects.

Because the distribution of precious corals is patchy and many form large solitary colonies, 

precious coral beds are often considered keystone species that provide habitat diversity (shelter 

or an epifaunal substratum) to many other species and for this reason they have been described 

as biodiversity hot spots. When all of the life history traits described above are considered 

collectively, it is clear that harvesting strategies must be very conservative. Harvesting rates, or the 

percent of the standing stock taken annually, must be very low (generally lower than the mortality 

rate) in order to have any chance of being sustainable. This is particularly important because 

virtually all species of commercially valuable precious corals are vulnerable to over-exploitation 

in unmanaged fisheries. This is because many year classes are exposed to harvesting at the same 

time. It is possible that many decades of accumulated standing stock is exposed to harvest during 

short intensive periods of fishing. Unfortunately, this pattern of pulse fishing has been employed 

by the Mediterranean fishermen especially when it was legal to use the Ingegno (also known as St. 

Andrews Cross) , as well as Japanese and Taiwanese coral draggers who use destructive tangle net 

gear. These practices have repeatedly led to a pattern of discovery, heavy exploitation followed by 

significant depletion. 

History of Precious Coral Fisheries in the Mediterranean and the Western Pacific

Red coral is found in the Mediterranean Sea at depths of 10-500 m; therefore early man may have 

collected it by way of shallow water divers or possibly from storm deposits on Mediterranean 

beaches. Starting about 5000 years ago, Greek sponge divers began to selectively harvest large 

colonies of red coral while collecting other organisms. Sometime thereafter, Greek fisherman 

73
learned to dislodge colonies from deeper water with the use of iron hooks (Buckner 2008). Several 

1000 years later, the Arabs invented a wooden cross with attached netting, to drag and entangle red 

coral from all depths. This device, known as the Ingegno (also known as the Cross of Saint Andrew), 

was used in Mediterranean Seas up until the 20th century. During the late 1800s, anywhere from 

several 100 to more than 1000 coral fishing vessels were engaged in red coral dredging operations 

in the shallows seas off Italy and Northern Africa. The large variability in fishing effort was the 

result of a boom and bust nature of the fishery. Production depended on the discovery of new coral 

beds but was always followed by excessive exploitation and rapid depletion. As a result, the use of 

coral dredges was banned outright by the European Union in 1989. Beginning in about 1960, the 

commercial harvest of red coral was carried out more and more by divers equipped with SCUBA 

equipment (Tescione 1965). The modern catch record for about the last 50 years for C. rubrum

has been compiled by FAO and recently published by A. Bruckner and again the cycle of boom or 

bust production is evident in the data (Fig. 1). These records demonstrate that virtually all known 

red coral beds in the Mediterranean have been depleted. Annual production today is about 25% of 

what it was in the early 1960s.

Precious corals were not discovered in the Pacific Ocean until 1803 in waters off Japan where 

beautiful red (Corallium japonicum) and pink (C. elatius) corals were found at depths ranging 

between 100-300 m. A fishery off Japan, however, did not develop until after the Meiji Reform in 

1868. Before then, corals harvested by fishermen were confiscated by the Shoguns. After 1868, 

fishermen were allowed to sell their catch on the open market. In the late 1880s about 100 small 

boats were engaged in harvesting precious corals from coastal waters of Japan and although catch 

records are not available, the methods used were tangle nets (coral mops) and the known grounds 

were gradually exhausted (Grigg 1971). New grounds were discovered sporadically but they too 

underwent depletion. Over the years, this same cycle of discovery, exploitation and depletion 

continued into the 20th century being interrupted only by World Wars I and II.

In mid 1960s, Japanese coral draggers made two large discoveries, the first on the Oza Banks 160 

km south of Okinawa and the second, on the Milwaukee Banks in the Emperor Seamounts 800 

km northwest of Midway Island at depths near 400 m (Grigg 1971). In 1969, Taiwanese boats 

discovered even more, new and rich precious coral banks on other Emperor Seamounts. During 

the 1960s almost 230 tons of red and pink coral worth over $4 million were harvested collectively 

from these areas (Grigg 1971). Unfortunately, all of these areas were gradually depleted in the 

following years.

In about 1979 more beds were discovered in the 400 m depth zone east of the Emperor Seamounts 

on the Mellish Seamounts leading to another coral rush, but again these beds were soon depleted. 

A final discovery in the Emperor Seamounts was made in the early 1980s when a new and very 

abundant species was found at depths between 1000-1500 m. Sometime known as scotch coral, or 

deep sea Midway coral, this species of Corallium sp. nov. is characterized by a blotchy pink coral 

skeleton. Unfortunately, this species was also depleted over the ensuing several years.

FAO has been compiling the landings of Corallium species beginning in 1963 to the present and 

these cycles of boom and bust are evident in the data (Bruckner 2008) (Fig. 1 a,b). During several 

of the peak years in the Pacific, total annual yields exceeded 200-300 metric tons. These records 

also demonstrate that all the major fisheries for Corallium in the Pacific and the Mediterranean 

Sea are now economically depleted, although none are close to biological extinction. It should 

74
therefore be clear that coral dredges (tangle nets and the Ingegno) do not produce sustainable 

yields and should be universally discontinued for the commercial harvest of precious coral.

History of Sustainable Precious Coral Fisheries in Hawaii

Over the past 50 years, two separate and distinct precious coral fisheries have thrived in Hawaii. 

The first fishery consists of two species of black coral, Antipathes griggi and A. grandis. Although 

both live within SCUBA diving depths, from 35-110 m, both are ahermatypic, non-reef building 

solitary corals, lacking zooaxanthellae. The second precious coral fishery is located in deeper 

water between 350-500 m and consists of two species of Corallium, C. secundum and C. regale, 

and one species of gold coral (Gerardia sp.). Except for Gerardia sp., for which a moratorium 

on fishing exists, all other species of precious coral that have harvested in Hawaii have been 

sustainable (Tables 1, 2). Yields have been relatively stable and have never exceeded estimates of 

Maximum Sustained Yield (MSY). Also the age frequency distributions and recruitment rates of 

the major species have remained relatively stable (Fig.s 2, 3) (Grigg 2001, 2002).

Commercial beds of black coral in Hawaii were discovered in 1958 by two SCUBA divers in the 

Au’au Channel off of the island of Maui. The main Corallium fishery began in 1966 soon after 

deeper water species were discovered at 400 m depth about 6 miles off Oahu in the Molokai 

Channel. Both of these discoveries led to an incipient fishery and both were at first unregulated 

since very little was known about their ecology (Grigg 1965, 1974).

At the beginning of these fisheries, an information vacuum existed in Hawaii, however, the records 

of precious coral fisheries in the Mediterranean and the far Western Pacific, left little doubt that 

the future of the new precious coral fisheries in Hawaii would depend on the development of 

selective methods of harvest and rigorous guidelines regarding minimum size limits and annual 

quotas. In response to this need, a long term program to study the ecology of precious corals began 

at the University of Hawaii in 1970 and continues to this day. The many results of this program 

can be found in numerous publications but the main findings relative to fisheries management 

are summarized in a Fishery Management Plan developed by the Western Pacific Fisheries 

Management Council and the State of Hawaii (Federal Register 1983).

At the outset of the program, it was determined that a model would be needed to produces estimates 

of MSY, as well as, size limits and annual weight quotas for individual species. Both species of 

black coral and Corallium that are harvested in Hawaii have similar life histories of slow growth, 

low but steady recruitment, low mortality and moderately great longevity (~ 100 years). The 

Beverton and Holt cohort production model (Beverton and Holt 1957), turned out to be ideally 

suited to estimate these parameters. A major assumption of this model is steady state recruitment, 

a characteristic that can be reasonably ascribed to both species groups (Fig.s 2, 3). The research 

program was hence designed to analyze recruitment rates, growth rates, mortality rates and ages at 

first reproduction. Model results for both species of black coral and for C. secundum are presented 

in Fig.s 4 and 5. Specific management guidelines are given in Table 1. Annual harvest data for both 

fisheries is presented in Tables 2 and 3.

Although the Corallium fishery has not been continuous over the years and there is a moderate 

degree in the variability of annual catch for both fisheries, MSY values for all species have never 

been exceeded and the fisheries indeed appear to be sustainable (Fig.s 2, 3). Both examples 

75
demonstrate that precious corals are renewable resources and can be fished sustainably using 

selective methods (Fig. 6). Fortunately, both Federal and State agencies in Hawaii have been active 

in adopting and enforcing the management guidelines. Cooperation of the local industry in not 

purchasing undersize colonies has also been important. These examples demonstrate that precious 

coral are renewable resources and can be successfully harvested given the cooperation of science 

and enforcement working together. Such a plan has worked in Hawaii for over 30 years.

Affects of Listing Black Corals on Appendix-II of CITES

Having reviewed the success of local management of precious corals in Hawaii, it is appropriate 

to examine the question of whether or not a listing on Appendix-II of CITES would improve the 

conservation of Corallium species. CITES regulations require export, import and re-export permits 

from countries of origin and destination which help in the monitoring of trade. The regulations also 

prohibit the issuance of permits from countries where collection is illegal or have been shown 

to be detrimental to the survival of a given species. The regulations are well intended but do 

they actually serve to conserve precious coral resources? Do they affect harvesting policies and 

practices at the local level?

Answers to these questions may not always be the same in every country so it is instructive to 

examine case histories. In Section 4 of this paper, it has been shown that black coral harvest has 

been sustainable for about 50 years in Hawaii. A Fishery Management Plan was developed by the 

WPFMC and the State of Hawaii in 1983 and it has been enforced at the local level in cooperation 

with local industry. Black corals were added to Appendix-II of CITES in 1981. The main effects 

of adding CITES regulations have been as follows:

1. Increased costs of procurement of raw and finished black coral products.

2. Increased costs of selling finished black coral products internationally. In the United States, 

each company is required to obtain a permit for a Master File each year from the U.S. Fish 

and Wildlife Service. This may take from 4-7 months turnaround time. Once a Master File 

has been established, single use permits are then required for each black coral product sold. 

Processing time for these permits is about 1 month. Each single use permit requires about 2 

hours of employee time and a $129 processing fee. Repairs require a new shipping permit. 

As a result of the costs and employee time requirements, some companies no longer sell 

black coral jewelry outside of the U.S. 

3. Domestic sales are also impacted negatively since increased procurement costs are passed 

on to the consumer thereby reducing demand.

Overall, less black coral is sold and in this sense black coral resources are conserved because less 

raw material is purchased. Lower demand may or may not lower supply (fishing effort) depending 

on the global marketplace which is very difficult to quantify.

In conclusion, the principal effects of listing black coral on Appendix-II of CITES on the industry 

in Hawaii have been 1) to hinder the trade, and 2) to create an excessive administrative burden 

on the industry. The long-term sustainability of the black coral fishery and the industry owes 

76
itself to local fisheries regulations and enforcement programs in the State of Hawaii, not CITES 

regulations.

General Problems with Listing Corallium spp. on Appendix-II of CITES

In addition to producing many of the same problems created by the listing of black coral on 

Appendix-II of CITES, there are many other problems of a more general nature (Sheikh and Corn 

2008), and are as follows:

1. According to Article-II in Appendix-II of the Treaty, only those species that are threatened 

with extinction (although not necessarily now but may become so) are eligible for inclusion. 

Although some species of Corallium may be near economic extinction in parts of the 

Mediterranean and the Western Pacific, none are at risk of biological extinction or survival. 

The quantitative definition of decline in the Treaty is when the standing crop (abundance) 

of a given species falls below 20% of its unfished stock. By this definition, no species of 

Corallium have been shown to be eligible for inclusion on Appendix-II.

2. The inclusion by China (effective 6-1-08) of four species of Corallium (C. japonicum, C. 

elatius, C. konojoi, and C. secundum) on Appendix-III of CITES requires all international 

traders to obtain one of the following 3 documents: 1) an export permit, for countries of 

origin, 2) a re-export permit, for other countries, or 3) a certificate of origin, for other 

countries (CITES Notification to the Parties No. 2008/027).

3. All CITES listings should be species specific in order to identify changes in the biological 

condition of specific stocks. Only 7 species of Corallium are commercially harvested in the 

world and species identification is very difficult.

4. Some species of Corallium and even bamboo corals are dyed or powdered or reconstituted 

in resin and therefore cannot be correctly identified.

5. Perhaps the biggest problem is that most of the Corallium traded in the world today was 

harvested years ago and is drawn from stockpiles and is therefore exempt from the Treaty 

(Sheikh and Corn 2008).

6. Many species of Corallium have distribution boundaries that extend beyond country 

boundaries obscuring the location of collection.

7. Another potential problem is that a listing on Appendix-II of CITES may create the 

appearance that Corallium species are under management control thereby defusing efforts 

at the local level to enact legislation and enforcement programs. 

8. The decline in the yield of C. rubrum may have been caused by the outlawing of the 

ingegno in 1989. Annual harvest in the Mediterranean Sea has been relatively stable for the 

past 20 years (Fig. 1).

77
9. The decline in the harvest of Pacific species of Corallium may in part be a result of reduced 

fishing effort caused by the coral glut produced in the 1980s and the consequential fall in 

price of the raw material.

10. In the U.S., a CITES Appendix-II listing would duplicate restrictions on the trade already 

required by the Lacey Act.

11. The primary focus of CITES is on trade rather than conservation; few criteria for 

management exist.

12. The permitting process of CITES has created an administrative and costly burden 

worldwide for the trade of listed species of precious coral and has done little to improve 

their conservation.

Summary and Conclusions

Precious corals have been treasured by man since antiquity and represent one of the oldest fisheries 

in recorded history. Most precious corals are k-selected being characterized by slow growth, low 

recruitment and mortality and relatively great longevity. The famous Red Coral of Commerce, C. 

rubrum, found primarily in the Mediterranean Sea, is an exception, being highly fecund, sometimes 

exceeding abundance levels of 1000 colonies/m2

, and having withstood centuries of fishing effort. 

Unfortunately, most Corallium fisheries in the world are in decline or have been depleted. In large 

part, this has been due to the use of destructive methods of harvest utilizing various dredging 

methods and the lack of appropriate management. These practices have repeatedly led to a pattern 

of discovery, exploitation and depletion.

In contrast to depleted Corallium fisheries elsewhere in the world, two precious coral fisheries 

in Hawaii have been sustainably fished for many decades using methods of selective harvest and 

management guidelines that are enforced at the local level. One of these fisheries, that targets black 

coral has also been affected by CITES Appendix-II regulations. Unfortunately, this added level of 

regulation has not served to help conserve the resource, but on the contrary, has hindered trade and 

has been a costly administrative burden on the local industry in Hawaii.

The proposal to add Corallium spp. on Appendix-II of CITES is plagued by a number of serious 

problems ranging from difficulties in identification, to stock exemptions, to potentially defusing 

local management, to lack of enforcement, to failure to conserve the resource and to hindering 

trade and creating administrative burdens on the industry. The history of precious coral fisheries in 

Hawaii has shown that the best management practice for the conservation of the resource has been 

the application of management guidelines produced locally combined with local enforcement by 

government agencies and industry.

At the international level, there is no question that precious coral resources are in need of much 

stronger management practices. The use of destructive fishing methods (dredging) need to be 

discontinued and replaced by methods of selective harvest. Species specific guidelines need to be 

developed and enforced at the local level. A myriad of possible management options exist including 

size limits, annual quotas, limited entry (licenses), seasonal limits, rotation of fishing effort and the 

78
creation of Marine Protected Areas (MPAs). All of these approaches are best accomplished at the 

local level through cooperation of science and enforcement, not CITES.

Table 1. Management guidelines of precious corals in Hawaii (Grigg 1976).

Species Growth Mortality Yield/ Age/MSY MSY OY Size

 cm/yr annual cf. recruit gms yrs Kg Kg inches

Black corals (A. griggi and A. grandis)

 Maui Bed 6.42 0.07 1050 gm 28 6174 5000 48

 Kauai Bed 1543 1250 48

Pink coral (C. secundum)

 Makapuu Bed 0.9 0.066 203 34 1072 1000 10

•	 The MSY size limit of 12” for C. secundum was lowered to 10” (optimum yield (OY)) 

due to economic considerations. 

The Preliminary Management Plan for precious corals was completed in 1977 (EIS/PMP 

for precious coral, 1977). It is recommended that coral dredging be discontinued for 

commercial harvest of all species due to it destructive nature.

Table 2. Annual harvest of pink, red, and gold coral in Hawaii (kg).

Year Gear C. secundum Gerardia sp. C. regale

1966-69 Dredge1. 1800 0 0

1970-72 No harvest 0 0 0

1973 Submersible2. 538 0 0

1974 “ 2209 734 0

1975 “ 1385 621 0

1976 “ 400 363 0

1977 “ 1421 329 0

1978 “ 474 50 0

1979-1987 No harvest 0 0 0

1988 Dredge3. 500 0 0

1989-99 No harvest 0 0 0

1999-2001 Submersible4. 1216 150 61

2002-2008 No harvest 0 0 0 ____

1. United Fishing

2. Maui Divers of Hawaii

3. Vessel Aukai 

4. American Divers

79
Table 3. Annual harvest of black coral* in Hawaii (kg/yr), except as noted.

Year Maui Bed Kauai Bed

1958-19681. 3,000 0

1968-19722. 4,000 2,000

1972-19803. 8,000 4,000 

1981 4. 72 0

1982 4 430 0

1983 4

 867 0

1984 4 1,422 0

1985 4

 140 0

1986 4 425 0

1987 4 1,978 0

1988 4 234 0

1989 4 464 0

1990 4 395 0

1991 4 1,048 0

1992 4 1,090 0

1993 4 393 0

1994 4 1,979 0

1995 4

 2,735 0

1996 4 2,211 0

1997 4. 689 0

1999-2005 5. 3,182 0__________

* Antipathes griggi and A. grandis

1-3. Annual harvest based on interviews with divers

4. Annual harvest as reported to the State of Hawaii, Div. of Aquatic Resources, 1981-1997

5. Parrish, 2006

80
References

Beverton R.J.H., Holt, S.V. 1957. On the dynamics of exploited fish populations Fish. Invest. Min. 

Agric. Fish. Food (G.B.), ser. II, 19: 1-553.

Bruckner, A. 2008. Rate and extent of decline in Corallium (pink and red coral) populations: are 

existing data adequate to justify a CITES Appendix II listing? Ecol. Prog. Ser.

Caddy, J. F. 1993. Background concepts for a rotating harvesting strategy with particular reference to 

the Mediterranean red coral, Corallium rubrum. Mar. Fish. Rev. 55: 10-18.

CITES Notification to the Parties No. 2008.027.

EIS/PMP for Precious Corals. U.S. Dept. of Commerce, NOAA, NMFS, 1977.

FAO. 2007. FAO Fish Report No. 833, Rome, Italy. 133 p.

Federal Register. 1983. Fishery Management Plan for Precious Coral Fisheries of the Western Pacific 

Region. Aug. 30, 1983.

Grigg, R.W. 1965. Ecological Studies of Black Coral in Hawaii. Pacific Science, 19: 244-260.

Grigg, R.W. 1971. Status of the precious coral industry in Japan, Taiwan and Okinawa. UNIHISEAGRANT-AR-71-02.

Grigg, R.W. 1974. Distribution and abundance of precious corals in Hawaii. Proc. 2nd.Internat. Coral 

Reef Sym. 2: 235-240.

Grigg, R.W. 1976. Fishery management of precious and stony corals in Hawaii. Sea Grant Tech. 

Rept., UH. 77-03. 48 p.

Grigg, R.W. 1984. Resource management of precious corals: A review and application to shallow 

water reef building corals. Marine Ecology 5: 57-74.

Grigg, R.W. 1993. Precious coral fisheries of Hawaii and the U.S. Pacific Islands. Marine Fisheries 

Review 55: 50-60.

Grigg, R.W. 2001. Black coral: History of a sustainable fishery in Hawaii. Pacific Science 55: 291-

299.

Grigg, R.W. 2002. Precious corals in Hawaii: discovery of a new bed and revised management 

measures for existing beds. Marine Fisheries Review 64(1): 13-20.

Hickson, S. 1924. An introduction to the study of recent corals. Man. Univ. Press, London, U.K.

Kishinouye, K. 1904. Notes on the natural history of corals. Imp. Fish. Bur. Tokyo J. 14: 1-32.

Kosuge, S. 1993. History of the precious coral fisheries in Japan (1). Precious Corals and Octocorals 

Research 1: 30-33.

Lacaze-Duthiers, H. 1864. Histoire naturelle du corail. J.B. Bailliere, Paris, France.

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in the precious red coral, Corallium rubrum. Coral Reefs 23: 423-432.

Parrish, F. 2006. Black coral fishery landings 1985-2005. WPRFMC Special Publication. 45 p.

Roark, E.B., Guilderson, T.P., Dunbar, R.B., Ingram, B.L. 2006. Radiocarbon-based ages and growth 

rates of Hawaiian deep-sea corals. Mar. Ecol. Prog. Ser. 327: 1-14.

81
Sheikh, P.A., Corn, M.L. 2008. The Convention on International trade in Endangered Species of 

Wild Fauna and Flora (CITES): background and issues. CRS Report for Congress, Order 

Code RL 32751, 14 p.

Tescione, G. 1965. Il corallo, nella storia e nell’arte. Montanino Editore, Napoli, Italy.

Tsounis, G., Rossi, S., Grigg, R., Santangelo, G., Bramanti, L., Gili, J.M. (in press). Precious coral 

exploitation. In: Fisheries: Management, Economics and Perspectives. NOVA Sci. Pub., 

NY, USA.

Vighi, M. 1970. Researche sul ciclo reproduttivo del corallo rosso (C. rubrum (L)) del promontorio 

di Portofino. Lincei-Mem. Sci. fisiche, ecc. Ser. III (10): 1-26.

82
Fig. 1. Landings in metric tons of Corallium populations in (A) the Mediterranean Sea and the 

Pacific Ocean between 1963 and 2005, and (B) Landings of individual species. Note stability of C. 

rubrum landings over the past 20 years (Bruckner 2008).

83
Fig. 2. Age frequency distributions of black coral (A.griggi) in the Maui bed in 1975 and 1998 

(Grigg 2001). Note that 97% of the bed in 1998 is entirely due to recruitment since 1975. In other 

words, recruitment has been sustainable over the entire time period of 23 years between 1975 and 

1998.

Fig. 3. Age structure (frequency distribution) of pink coral (C. secundum) in the Makapuu Bed, 

Oahu, in 1971, 1983, 1985 and 2001. Arrow bars indicate relative decrease or increase in age 

frequency since reference year.

84
Fig. 4. Yield-per-recruit of pink coral (C. secundum) in the Makapuu Bed, Oahu based on the 

Beverton and Holt yield-per-recruit Model at three different values of natural mortality. Dashed 

lines indicate yield produced by applying a fishing mortality of 0.06 at four different ages.

Fig. 5. Yield-per-recruit of black coral (A. griggi) in the Au’au Channel Bed, Maui, based on the 

Beverton and Holt yield-per-recruit Model at three different values of natural mortality. Dashed 

lines indicate yield produced by applying a fishing mortality of 0.06 at four different ages.

85
Fig. 6. In 1976, Maui Divers of Hawaii designed and used a basket and coral cutter assembly to 

selectively harvest precious corals. The basket assembly was attached to the Star II Submersible 

and operated by the pilot from inside the submersible.

86
THE DEEPEST KNOWN OCCURRENCE OF 

THE PRECIOUS RED CORAL CORALLIUM RUBRUM (L. 1758) 

IN THE MEDITERRANEAN SEA

M. Taviani1*, A. Freiwald2, L. Beuck2, L. Angeletti1, A. Remia1, 

A. Vertino1,3, M. Dimech4, P.J. Schembri5

1

 ISMAR-CNR, Via Gobetti 101, 40129 Bologna, Italy, 

and CITES Scientific Commission, Italy

2

 GeoZentrum Nordbayern, Universität Erlangen-Nürnberg, 

Loewenichstr. 28, 91054 Erlangen, Germany

3 Università degli Studi di Catania, Dipartimento di Scienze Geologiche, 

Corso Italia 57, 95129 Catania, Italia

4

 Malta Centre for Fisheries Sciences, Fort San Lucjan, Marsaxlokk BBG 06, Malta

5

 Department of Biology, University of Malta, Msida MSD2080, Malta

*E-mail: marco.taviani@bo.ismar.cnr.it;

Introduction

An initiative for the worldwide conservation of gorgonian corals of the family Coralliidae 

(Anthozoa, Gorgonacea) was attempted at the 14th Meeting of the Conference of the Parties of 

CITES at The Hague (CoP14) through a proposal to place members of this family on AppendixII of the Convention (Anonymous 2007; Pala 2007; WWF 2007). Ultimately the proposal was 

defeated in a secret ballot, although the decision was taken to organize topical workshops devoted to 

upgrading knowledge of precious corals, which are all grouped in the two extant genera Corallium

and Paracorallium. This call resulted in two workshops focusing on Pacific and Mediterranean 

precious corals, held in 2009 in Hong Kong and Naples, respectively (IAPPCS 2009; Anonymous 

2009). In the meantime the request for inclusion of the Coralliidae in Appendix-II of CITES is in 

all likelihood going to be proposed for consideration at the forthcoming CoP15 in Qatar, scheduled 

for the year 2010. 

Corallium rubrum colonizes a variety of sublittoral habitats generally between a few meters depth 

down to 120 m in the Mediterranean Sea and Eastern Atlantic Ocean (e.g. Bayer 1964; Carpine and 

Grasshof 1975; Zibrowius et al. 1984; Chintiroglou et al. 1989; Cattaneo-Vietti and Cicogna 1993; 

Abbiati et al. 1993; Garrabou and Harmelin 2002; Torrens et al. 2008; Rossi et al. 2008; Tsounis 

et al. 2006). The bright red calcitic axis of this species has been prized in the jewelry trade from 

antiquity to the present day (Cattaneo-Vietti and Cicogna 1993; Tsounis et al. 2007). Although far 

from being close to extinction, a decline in shallow-water populations of this widespread taxon has 

nevertheless been observed, giving rise to some concerns about its management (Santangelo and 

Abbiati 2001; Santangelo et al. 2003; Tsounis et al. 2007). 

The present paper is a contribution on novel aspects of the ecology, biology and biogeography of 

Corallium rubrum based on the recent discovery of previously uncharted deep-water populations 

in the bathyal zone of the Mediterranean Sea (Fig. 1).

Distribution of Corallium rubrum at Bathyal Depths

87
Living red coral colonies were observed in 2006 during the Marum-Quest 4000 ROV survey of the 

R/V Meteor in the Strait of Sicily in water depths down to at least 684 m (Freiwald et al. 2009: Fig. 

4). Before this discovery, the deepest confirmed lower bathymetric limit of this species was 230 m 

at Cap Creus in the Western Mediterranean (Rossi et al. 2008). In the Strait of Sicily deep water, 

Corallium rubrum is now documented to occur at three disjunct sites, namely (1) Linosa Island, 

dive 673; (2) Nameless-Urania Bank, dive 677; (3) South of Malta, dive 657 (see Freiwald et al. 

2009 for details of the ROV dive stations). The first two sites refer to submarine volcanic edifices 

where the species fringes overhangs and ‘caves’ often in close association with living colonial and 

solitary scleractinians (e.g. Lophelia pertusa, Madrepora oculata, and Desmophyllum dianthus), 

antipatharians and gorgonians (Fig. 2). The site South of Malta site refers to a sedimentary 

escarpment and is located in the South Malta Deep-Water Coral Province first described by 

Schembri et al. (2007). Here Corallium rubrum is also found in intimate association with white 

stony corals, primarily Madrepora and Lophelia. In addition, red coral branches, some possibly 

still alive, were also spotted in recent coral taluses accumulated at the foot of coral-covered walls, 

again in association with ‘white corals’ (Fig. 2).

Colonies of red coral were collected alive for the first time from such deep-sea sites off Linosa 

and South of Malta in 2007 during the scientific cruise MARCOS of R/V Urania (Fig. 3). Some of 

these samples were provided to investigators for genetic characterization to investigate the genetic 

connectivity between geographically and/or bathymetrically disjunct coral sites (Costantini et al. 

in press). Samples have been used for the geochemically-based evaluation of climatic signals 

incorporated into the calcitic skeleton (Montagna et al. 2008, in progress).

Conclusions and Recommendations 

The discovery of deep-sea populations of C. rubrum is important for the overall characterization of its 

ecological requirements (substratum and physicochemical factors), connectivity (interrelationship 

between disjunct coral populations), and sustainable management. It is also relevant for the 

evaluation of the present status of this species in the Mediterranean Sea and the adjacent Atlantic 

Ocean.

The existence of previously uncharted deep-water populations is opening new perspectives in 

characterizing the distribution and the role that this habitat forming species plays in ecosystem 

processes of the deep Mediterranean Sea. This discovery increases significantly the known depth 

range of this cnidarian, suggesting its potential presence at many other still-uncharted deep-water 

sites in the Mediterranean basin. 

The in situ observation of its intimate association with other ecologically valuable sessile organisms 

(white corals, antipatharians etc.) is a strong argument against any commercial exploitation of these 

deep-water colonies, and no exploitation whatsoever should ever be considered. On the contrary, 

such fragile red coral bearing deep-water habitats are in strong need of proper management to 

ensure their protection. The relative inaccessibility of such remote habitats at present serves 

to protect them; however, the continuous implementation of wide scale operations in the deep 

ocean (including industry) could in principle put in jeopardy some unique systems. Action should 

therefore be taken to ensure the inclusion of known deep-sea Corallium habitats within marine 

protected areas. 

88
Acknowledgements 

We are grateful to Captains and crew and scientific staff of RV Urania and RV Meteor during 

cruises MARCOS and M70-1, respectively. F. Mastrototaro, C. Misfud, and H. Zibrowius helped 

with preparation and sorting of MARCOS samples, taxonomy and photography. Research partially 

funded by CNR, HERMES (GOCE-CT-2005-511234-1) and HERMIONE (contract number 

226354) projects of the European Commission, and BIOCORAL (Ministry of Environment of 

Italy), grants. PJS was supported by research grants from the University of Malta. This is ISMARBologna scientific contribution n 1651.

References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Marine Ecology Progress 

Series 95: 245-250.

Anonymous. 2007. Corallium in the red., Synopsis of literature for an Appendix II listing. Seaweb: 

59 p.

Anonymous. 2009. Red Coral Science, Management and Trading: Lessons from the Mediterranean. 

Ministero Affari Esteri, Ministero dell’Ambiente e della Tutela del Territorio e del Mare, 

Università degli Studi di Napoli “Parthenope”, International Workshop Napoli-Villa Doria 

D’Angri. 23-26 Settembre 2009, Abstract, 49.

Bayer, F.M. 1964. The genus Corallium (Gorgonacea:Scleraxonia) in the western North Atlantic 

Ocean. Bulletin of Marine Science Gulf and Caribbean 14: 465-478.

Carpine, C., Grasshof, M. 1975. Les Gorgonaires de la Méditerranée. Bulletin de l’Institut 

Océanographique. Institut océanographique 71: 1-140. 

Cattaneo-Vietti, R., Cicogna, F. (Eds.) 1993. Il corallo rosso in Mediterraneo: Arte, Storia e Scienza, 

Red Coral in the Mediterranean Sea: Art, History and Science, Ministero delle Risorse 

Agricole, Alimentari e Forestali, Roma, 263 p.

Chintiroglou, H., Dounas, C., Koukouras, A. 1989. The presence of Corallium rubrum (Linnaeus, 

1758) in the eastern Mediterranean Sea. Mitteilungen aus dem Zoologischen Museum in 

Berlin 65: 145-149.

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembri, P.J., Abbiati, M. Deep-water Corallium 

rubrum (L. 1758) from the Mediterranen Sea: preliminary genetic characterization. Marine 

Ecology (in press) DOI: 10.1111/j.1439-0485.2009.00333.x.

Freiwald, A., Beuck,L., Rüggeberg, A., Taviani, M., Hebbeln, D. and R/V Meteor M70-1 Participants. 

2009. The white coral community in the Central Mediterranean Sea revealed by ROV 

surveys. Oceanography 22: 58-7.

Garrabou, J., Harmelin, G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: insights into conservation and management 

needs. Journal of Animal Ecology 71: 966-978.

IAPPCS. 2009. Report of the First Meeting in Hong Kong, March 7, 2009. International Forum on 

Precious Coral, 50 p.

89
Montagna, P., Vieulzeuf, D., López, Correa, M., Garrabou, J., Taviani, M., Marschall, C., Linares, 

C., McCulloch, M., Silenzi, S., Freiwald, A. 2008. Growth rate, trace elements and stable 

isotopes in Corallium rubrum from shallow and bathyal settings in the Mediterranean 

Sea. In: Deepsea Coral Symposium 2008, 4th ISDSC, Programme and Abstract Book, 

Wellington, New Zealand 1-5 December 2008, 188 p.

Pala, C. 2007. Red coral in the Red. Science 316: 1113.

Rossi, S., Tsounis, G., Orejas, C., Padron, T., Gili, J.M., Bramanti, L., Teixido, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Marine Biology 154: 533-545.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an overexploited 

Mediterranean species. Aquatic Conservation: Marine and Freshwater Ecosystems 11: 253-

259.

Santangelo, G., Carletti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Marine Ecology 

Progress Series 248: 99-108.

Schembri, P.J., Dimech, M., Camilleri, M. 2007. Living deep-water Lophelia and Madrepora corals 

in Maltese waters (Strait of Sicily, Mediterranean Sea). Cahiers de Biologie Marine 48: 77-

83. 

Torrents, O., Tambutte, E., Caminiti, N., Garrabou, J. 2008. Upper thermal thresholds of shallow vs. 

deep populations of the precious Mediterranean red coral Corallium rubrum (L.): Assessing 

the potential effects of warming in the NW Mediterranean. Journal of Experimental Marine 

Biology and Ecology 357: 7-19.

Tsounis, G., Rossi, S., Aranguren, M., Gili, J.M., Arntz, W. 2006. Effects of spatial variability and 

colony size on the reproductive output and gonadal development cycle of the Mediterranean 

red coral (Corallium rubrum L.). Marine Biology 148: 513-527.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2007. Red coral fishery at the Costa Brava (NW 

Mediterranean): case study of an overharvested precious coral. Ecosystems 10: 975-986.

WWF. 2007. WWF Positions. CITES CoP14, 21-23.21.

Vertino, A., Zibrowius, H., Taviani, M. (this volume). Fossil ancestors of family Coralliidae in the 

Mediterranean Sea. 

Zibrowius, H., Montero, M., Grashoff, M. 1984. La répartition du Corallium rubrum dans l’Atlantique. 

Thetys 11: 163-170.

90
Fig. 1. Map of the Strait of Sicily showing the location of the MARCOS stations that provided live 

C. rubrum from bathyal depths. Red dots are samples from MARCOS Cruise; yellow diamonds 

are ROV dives from Meteor M70/1 Cruise (see Freiwald et al. 2009 for details).

91
Fig. 2. MARUM ROV ‘Quest 4000’ in situ documentation of Corallium rubrum occurrences at 

Linosa Island site, Dive 673, Linosa Trough: (a) panoramic view of the complex coral-bearing 

community fringing overhangs at the edge of volcanic caves at 673 m water depth: note clumps of 

live C. rubrum (arrows) intimately associated with other cnidarians, including Madrepora oculata; 

(b) close-up of living C. rubrum from the same site showing fully expanded polyps (-673 m); (c) 

biostromal coral accumulation at the foot of the Linosa volcanic edifice; note fresh-looking C. 

rubrum and Madrepora together with degraded and partly silted white and red coral debris (-737 

m). All pictures courtesy of MARUM, Bremen University.

92
Fig. 3. Examples of deep-water red coral samples from the Strait of Sicily, MARCOS Cruise: (a) 

Coral-hardground with modern C. rubrum colonies St. MS43; (b) Large live C. rubrum colony st. 

MS43, ca. 650 m; (c) Degraded patinated hardground with red coral growth St. MS32; (d) Living 

colony of C. rubrum with expanded polyps, st. MS43; (e) Living colony of C. rubrum growing 

on degraded coral hardground st. MS43; (f) Desmophullym dianthus fouled by recent C. rubrum.

93
FOSSIL CORALLIIDAE IN THE MEDITERRANEAN BASIN

A. Vertino1,4*, H. Zibrowius2, M. Rocca3 , M.Taviani4 

1

 Università degli Studi di Catania, Dipartimento di Scienze Geologiche, 

Corso Italia 57, 95129 Catania, Italia

2

 Station Marine d’Endoume, Rue Batterie des Lions, 13007 Marseille, France

3

 Turin, Italy

4

 ISMAR-CNR, Via Gobetti 101, 40129 Bologna, Italy, and CITES Scientific Commission, Italy

*E-mail: avertino@unict.it

Introduction

Members of the family Coralliidae (Cnidaria, Octocorallia, Gorgonaria, Scleraxonia) are 

characterized by a massive, continuous, calcitic axis, and loose calcitic sclerites imbedded in the 

soft polyp and coenenchyme tissues. The axis is easily preserved as a conspicuous fossil whereas 

the tiny (taxonomically significant) sclerites are dispersed post mortem and are still unknown in 

the fossil record. Discrimination of species is based on shape and size of sclerites, branching mode 

of the colonies, and axis ornamentation (e.g. Bayer 1964; Bayer and Cairns 2003). The some 

twenty species described in the recent fauna world-wide have lately been distributed between 

the genera Corallium (Cuvier 1797) and Paracorallium (Bayer and Cairns 2003), the latter being 

characterized by longitudinally grooved axes and axial pits with beaded margins containing the 

autozoid polyps. 

Part of the deep-water coralliid species world-wide have their branches modified by symbiotic 

polynoid polychaetes which induce the formation of gall-galleries with series of lateral openings. 

These galls run along or curl around the branches, providing a shelter for the symbiont (Bayer 

1964). Polynoids associated with various deep-water coralliid species from the Atlantic and Indian 

Oceans have been identified as Polynoe caeciliae Fauvel 1909, by Hartmann-Schröder 1985 (see 

also Martin and Britayev 1998). Polynoid gall-galleries on Coralliidae are of the same type as 

skeletal deformations observed in many species of Stylasteridae (Hydrozoa), likewise caused by 

other species of Polynoidae (Zibrowius 1981; Cairns 1991).

In the present-day Mediterranean Sea, the family Coralliidae is represented by the single species 

Corallium rubrum (Linnaeus 1758), which is eurybathic (from shallow low-light to bathyal aphotic 

habitats) and widespread in this basin and in the adjacent Atlantic (e.g. Carpine and Grasshoff 1975; 

Zibrowius et al. 1984; Chintiroglou et al. 1989; Cattaneo-Vietti and Cicogna 1993; Santangelo 

and Abbiati 2001; Tsounis et al. 2007; Rossi et al. 2008; Freiwald et al. 2009; Taviani et al., this 

volume, with references therein).

However, as shown here, Mediterranean Coralliidae were more diverse in the past and had been 

established in this sea since at least the Early Miocene. These preliminary results arise from the 

critical examination of literature data, museum collections and new paleontological material 

collected from Cainozoic deposits in northern and southern Italy (Fig. 1).

94
Cainozoic Documentation of Coralliidae in the Mediterranean Basin

The oldest record of Corallium in the Mediterranean area (De Angelis 1894) is Oligocene 

(“Tongriano”) but needs to be verified. Records from the Miocene are more reliable, some even duly 

confirmed and well illustrated. Several examples are thus reported in the literature (e.g. Zuffardi 

Comerci 1927; Chevalier 1961; Barrier et al. 1992; and references therein). Various fossils have 

been examined for the present study. 

Although the taxonomy of the Miocene coralliids remains problematical (sclerites still unknown) 

it can be stated already that in the Mediterranean Miocene two types of Coralliidae are represented: 

(1) axes morphologically indistinguishable from those of the recent Corallium rubrum, and (2) 

others most similar to those of deep-water species known from the present-day Atlantic fauna, 

their resemblance marked by the presence of polynoid gall-galleries. Records of the latter type 

include the Lower Miocene (Burdigalian) of the Turin Hills (northern Italy: Zuffardi Comerci 

1927; Taviani et al. 2005) and the Upper Miocene of Carboneras (SE Spain: Barrier et al. 1992); 

Fig. 1.

Depending on the state of preservation, coralliids of the “deep-water”-type are more or less 

easily distinguished from their Corallium rubrum-like counterparts by (1) branching mainly in 

Fig. 1. Location map of fossil Coralliidae in the Mediterranean area. Only confirmed records 

are indicated (samples examined and confirmed as Coralliidae). White marks: Miocene from 

Carboneras (SE Spain) and Turin Hills (N Italy); black marks: Pleistocene from Capo Milazzo, 

San Martino and Messina (Sicily, S Italy); grey mark: Pleistocene from Augusta and the vicinity 

of Catania (Sicily, S Italy).

95
one plane, (2) having secondary branchlets distributed only along one side of the main branches 

and/or eventually showing more distinct axial depressions corresponding to autozoids, and (3) 

bearing polynoid gall-galleries (even if not on all fragments). The galls are fragile structures often 

preserved only as longitudinal grooves (Fig. 2 a,b).

The larger and better-preserved “deep-water” coralliids with occasionally distinct polynoid gallgalleries were found to be more abundant at some Pleistocene sites of Sicily (Messina, San Martino, 

and especially Capo Milazzo: Figs. 1, 2 b,c). It can be deduced from their shape that colonies 

were essentially uniplanar. The axis surface is devoid of deep (autozooid) depressions, shows a 

fine granulation and a faint longitudinal striation which is more evident on the side not bearing 

secondary branchlets. As in recent Atlantic bathyal species (e.g. Corallium johnsoni, C. niobe), the 

Pleistocene fossils have polynoid gall-galleries located on the “front” side of the branches where 

the induced abnormal growth of secondary branchlets contributes to the gall formation by fusing 

upwards and producing the cage-like structures (Fig. 2 b). 

In addition, Pleistocene deposits of southern Italy (Sicily) produced tiny coralliid fragments marked 

by deep (autozooid) depressions with their margin conspicuously beaded. This is the first record 

of what, according to Bayer and Cairns 2003, would be a character of the genus Paracorallium.

Finally, fossils morphologically indistinguishable from the recent Corallium rubrum are known 

from Pleistocene deposits (mostly circalittoral) at many sites in Sicily (e.g. vicinity of Catania, 

Augusta). A comprehensive compilation of fossil occurrences of this species is in progress.

Discussion and Conclusions

The data presented here document that Coralliidae morphologically close to, if not identical, with 

the recent species Corallium rubrum were already present in the Mediterranean since Miocene 

times. The data also indicate that the basin was, at times, inhabited by more than one species of 

Fig. 2. Fossil Corallium fragments showing traces of polynoid gall-galleries (white arrows); a 

Burdigalian from the Turin Hills (N Italy), scale bar 1 cm; b,c Pleistocene from Capo Milazzo 

(Sicily, S Italy), scale bars 2 cm.

96
Coralliidae as this is still the case now in the adjacent Atlantic. Thus the Coralliidae is one more 

group (in addition to, e.g. scleractinian corals, molluscs and pedunculate crinoids) that testifies of 

the “more oceanic” character of the ancient Mediterranean deep-water fauna when homothermy of 

deep waters at a relatively high temperature was not yet established. The new fossil records also 

include forms that would conform to the genus Paracorallium, previously recognized only in the 

Indo-Pacific (Bayer and Cairns 2003). 

It can be assumed that populations of Corallium rubrum did occur, without interruption, in the 

Mediterranean basin at least since the early Pleistocene. Their early presence may explain (1) 

the variety of partly-disjunct habitats presently occupied basin-wide, from shallow to bathyal 

depths, and (2) the significant genetic variability displayed by shallow and bathyal populations 

(e.g.,Abbiati et al. 1993; Costantini et al., in press). This aspect should be taken into account when 

considering sustainable management of C. rubrum throughout its Mediterranean-Atlantic range.

Acknowledgements

We are grateful to V. Bugnano for his assistance in paleontological research on the Miocene of the 

Turin Hills. A. Logan kindly revised the English text. Research was partially funded by HERMES 

(GOCE-CT-2005-511234-1) and HERMIONE (contract number 226354) projects of the European 

Commission and Catania University research programs (PRA: “Programmi di Ricerca di Ateneo”). 

This is ISMAR-Bologna scientific contribution no. 1652.

References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Marine Ecology Progress 

Series 95: 245-250.

Bayer, F.M. 1964. The genus Corallium (Gorgonacea: Scleraxonia) in the western North Atlantic 

Ocean. Bulletin of Marine Science of the Gulf and Caribbean 14: 465-478.

Bayer, F.M., Cairns, S.D. 2003. A new genus of the scleraxonian family Coralliidae (Octocorallia: 

Gorgonacea). Proceedings of the Biological Society of Washington 116(1): 222-228.

Barrier, P., Zibrowius, H., Lozouet, P., Montenat, C., Ott D’Estevou, P., Serrano, F., Soudet, H.J. 

1992. Une faune de fond dur du bathyal supérieur dans le Miocène terminal des Cordillères 

Bétiques (Carboneras, SE Espagne). Mésogée 51 (1991): 3-13.

Carpine, C., Grasshoff, M. 1975. Les Gorgonaires de la Méditerranée. Bulletin de l’Institut 

Océanographique, Monaco 71 (1430), 1-140. 

Cattaneo-Vietti, R., Cicogna, F. Eds. 1993. Il corallo rosso in Mediterraneo: Arte, Storia e Scienza, 

Red Coral in the Mediterranean Sea: Art, History and Science, Ministero delle Risorse 

Agricole, Alimentari e Forestali, Roma, 263 p.

Cairns, S.D. 1991. The Marine Fauna of New Zealand: Stylasteridae (Cnidaria: Hydroida). New 

Zealand Oceanographic Institute Memoir 98: 1-179.

Chevalier, J.P. 1961. Recherches sur les Madréporaires et le formations récifales miocènes de la 

Méditerranée occidentale. Mémoires de la Soiété Géloogique de France 93: 1-562.

97
Chintiroglou, H., Dounas, C., Koukouras, A. 1989. The presence of Corallium rubrum (Linnaeus, 

1758) in the eastern Mediterranean Sea. Mitteilungen aus dem Zoologischen Museum in 

Berlin 65: 145-149.

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembri, P.J., Abbiati, M. Deep-water Corallium 

rubrum (L. 1758) from the Mediterranean Sea: preliminary genetic characterization. Marine 

Ecology (in press) DOI: 10.1111/j.1439-0485.2009.00333.x.

De Angelis, G. 1894. I Corallari dei terreni terziari dell’Italia settentrionale. Collezione Michelotti. 

Museo Geologico della R. Università di Roma. Atti della R. Accademia dei Lincei, Memorie 

della Classe di Scienze fisiche, Matematiche e Naturale, Ser. 5, 1, 164-280, pl. 1-3.

Freiwald, A., Beuck, L., Rüggeberg, A., Taviani, M., Hebbeln, D. and R/V Meteor M70-1 Participants 

(2009) The white coral community in the Central Mediterranean Sea revealed by ROV 

surveys. Oceanography 22: 58-7.

Hartmann-Schröder, G. 1985. Polynoe caeciliae Fauvel (Polynoidae), ein mit Korallen assoziierter 

Polychaet. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 82: 

31-35.

Martin, D., Britayev, T.A. 1998. Symbiotic polychaetes: review of known species. Oceanography 

and Marine Biology - Annual Review 36: 217-340. 

Rossi, S., Tsounis, G., Orejas, C., Padron, T., Gili, J.M., Bramanti, L., Teixido, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Marine Biology 154: 533-545.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an overexploited 

Mediterranean species. Aquatic Conservation: Marine and Freshwater Ecosystems 11: 253-

259.

Taviani, M., Freiwald, A., Beuck, L., Angeletti, L., Remia, A., Vertino, A., Dimech, M., Schembri, 

P.J. (this volume). The deepest known occurrence of the precious red coral (Corallium 

rubrum L. 1758) in the Mediterranean Sea. 

Taviani, M., Freiwald, A., Zibrowius, H. 2005. Deep coral growth in the Mediterranean Sea: an 

overview. In: Freiwald, A., Roberts, J.M. (Eds.) Cold-water Corals and Ecosystems, 

Springer, Berlin Heidelberg New York, 137-156.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2007. Red coral fishery at the Costa Brava (NW 

Mediterranean): case study of an overharvested precious coral. Ecosystems 10: 975-986.

Zibrowius, H. 1981. Associations of Hydrocorallia Stylasterina with gall-inhabiting Copepoda 

Siphonostomatoidea from the South-West Pacific. Part I. On the stylasterine hosts, 

including two new species, Stylaster papuensis and Crypthelia cryptotrema. Bijdragen tot 

de Dierkunde 51 (2): 268-286.

Zibrowius, H., Monteiro Marques, V., Grasshoff, M. 1984. La répartition du Corallium rubrum dans 

l’Atlantique. Téthys 11 (2): 163-170.

Zuffardi Comerci, R. 1927. Alcionari del bacino terziario Ligure-Piemontese. Atti della reale 

Accademia nazionale dei Lincei. Memorie della Classe di Scienze fisiche, Matematiche e 

Naturale, Ser. 6, 2, 559-576, 1 pl.

98
GEOCHEMICAL INVESTIGATION OF CORALLIUM RUBRUM

FROM SHALLOW AND BATHYAL SETTINGS IN THE MEDITERRANEAN SEA 

P. Montagna1,2,3, D. Vieulzeuf4

, M. López Correa5, J. Garrabou6, M. Taviani3,

 C. Marschall6, C. Linares4, M. McCulloch6, S. Silenzi7, A. Freiwald5

1

 LSCE, Av. de la Terrasse, 91198 Gif-sur-Yvette, France 

2

 LDEO, 61 Route 9W, Palisades, NY, USA

3

 CNR-ISMAR, Via Gobetti 101, 40122 Bologna, Italy

4

 CRMCN, Campus de Luminy, Marseille 13288, France

5

 GZN, GeoZentrum Nordbayern, University of Erlangen, Loewenichstr. 28, 

91054 Erlangen, Germany

6

 UMR 6540 DIMAR, COM, Station Marine d’Endoume, Marseille 13007, France

6

 ANU-RSES, Mills Road, Canberra 0200, Australia

7

 ISPRA, Via di Casalotti 300, Rome, Italy

Specimens of the calcitic gorgonian coral Corallium rubrum collected from shallow and bathyal 

settings in the Mediterranean Sea have been investigated for trace elements and stable isotopes at 

high-spatial resolution, using laser ablation ICP-MS and a computer-controlled micro-drill. Our 

research aims to assess the environmental parameters recorded in the calcitic skeleton of this 

precious long-lived coral as geochemical signals, considering the possible complications of the 

biological activity of the organism during the skeletogenetic processes. Laser ablation ICP-MS and 

stable isotope transects across the solid axial calcareous skeleton of living and fossil specimens 

from shallow water off Marseille, Corsica and the Medes Islands and deep-water settings off of 

Linosa show large geochemical variability (25% for Mg/Ca, 57% for B/Ca, 20% for Sr/Ca, 200% 

for U/Ca, 17% for Ba/Ca, 75% for P/Ca and 80% for Li/Ca). Bathyal environmental conditions 

are quite stable at around 13.9°C and 38.77 PSU throughout different seasons and the amplitude 

of the geochemical tracers should be minimal. The highly variable and microstructure-related 

geochemical concentration across the axis indicates that vital effects might be involved and partly 

controls the systematics of the trace elements, with most of the elements significantly enriched in 

the medullar region. The role of the biological activity in obliterating the environmental signals 

recorded in the geochemical pattern can be best assessed at the bathyal sites and might have also 

implications for shallow water specimens.

The multi-proxy approach will also help to test the likely annual nature of the thin banding in 

bathyal specimens and the use of high-resolution micro-analytical techniques will potentially 

provide monthly records of temperature and other important environmental parameters (e.g. 

nutrient content).

99
Section II

Harvest, Manufacturing, and Trade

100
OVERVIEW OF LAW ENFORCEMENT ISSUES 

REGARDING THE TRADE IN CORALLIUM 

A. Bailey

Fish and Wildlife Service, Office of Law Enforcement

 United States Department of the Interior,

16639 W. Hardy

Houston, TX 77060

Wildlife Inspectors at various ports in the U.S. regulate the import and export trade in the various 

species of coral, including the precious Red Corals, (Corallium sp.). Included in the many issues 

and challenges faced by these inspectors are; a lack of training or experience in identification of 

the various species of red corals listed on CITES and other similar non-CITES listed corals, and 

a lack of ready access to experts in this area. Despite such challenges, Wildlife Inspectors remain 

dedicated to the enforcement of CITES laws and regulations. Data from the Law Enforcement 

Management Information System (LEMIS) shows the trends of imports and exports of such coral 

items in the trade.

101
“THE CORAL ROUTES:” FROM THE MEDITERRANEAN SEA 

TO THE WORLD-WIDE JEWELRY

A.E. De Simone 

AssoCoral, Torre del Greco, Italy

Certainly the fascination that coral has always exercised on the human imaginary originates from 

its enigmatic nature, between the three kingdoms: not a plant, even if branched, not a mineral, 

though petrified, not exactly an animal, even if of the color of blood, but in fact we now know that 

coral is composed almost entirely of calcium carbonate produced by colonies of microorganisms 

and is therefore of animal origin. Its singular genesis has been a source of myths and beliefs of 

portentous apotropaic virtues, which have crossed the centuries-old traditions of different and 

distant cultures from East to West. 

The ancient Romans of the first century A.D. already wondered about its origin, and soon joined 

its warm and vivid color to blood, so that Ovid in his “Metamorphoses” and Pliny the Elder in his 

“Naturalis Historia” recognized to coral the same mythical genesis: the blood which continued 

to drip from Medusa’s Severed Head, turned into coral. The coral ornaments found in some 

archaeological sites suggest that it was already known several thousand years ago, and we know 

for certain that the Greeks appreciated coral and used it also in pharmacology. But it was only 

a few centuries later, with the ambitious project of Alexander the Great to fuse East and West 

into a single empire not only through military conquests, but also thanks to a cultural unification 

favored by trade, that coral opened its way to India. Pliny provides tangible proofs of this, in the 

aforementioned “Naturalis Historia,” stating that: “The Indians prize coral just as the Romans prize 

pearls, the cost varying according to the importance attributed by each people to the commodity 

in question.”

Coral fishing was practiced throughout the Mediterranean and surely the Greeks, the Phoenicians, 

and the Etruscans were devoted to it and through this dense network of routes by land, sea, and 

river, along which in ancient times the Chinese empires and the West used to trade, the coral 

arrived in the Middle East, Asia Minor, India, China, and became deeply rooted in the customs of 

those eastern nomadic peoples where it was even used as an ornament for their horses, a protection 

before battles, seeing it as a formidable talisman. The routes of silk and spices became therefore 

a huge melting pot in which the most different cultural experiences met and melted giving rise to 

unique results, real synthesis of the encounter between different traditions. The wonderful ethnic 

jewels made from coral fished and worked mainly in Mediterranean countries and then assembled 

according to local customs, are the most eloquent testimony of the exchanges between the 

precious goods of those eastern countries and coral: the most typical product of the Mediterranean 

civilizations. This extraordinary jewelry is the tangible evidence of the millennia–old links that 

have joined East and West, through the renowned silk and spice routes, which became also the 

routes of coral, since our red gold was often the price paid for the precious eastern goods.

Since ancient times coral and magic have been chasing each other, intertwining, confusing in 

superstition, or just good-luck rituals, but perhaps what is more amazing is that over the centuries 

coral has always been recognized as a powerful talisman: from the Mongols to the Indians, from the 

Chinese to the American Indians, through Yemen, Turkey, Uzbekistan, North Africa, and Europe. 

102
This fascinating and unusual circumstance was due to the fact that the “Silk Route,” which started 

two thousand years ago from northern China, across the steppes of Central Asia and arrived in the 

West, and the “spice route,” which connected the deep south of the Arabian Peninsula with Mare 

Nostrum, had been travelled also in the opposite direction from brave merchants, people who had 

moved from the Mediterranean to the Far East, using the red coral as a bargaining chip, thereby 

increasing its perfect assimilation with local habits. 

What makes the ethnic jewel so peculiar is its strong symbolic language to which no part is by 

chance, but always the result of a well-considered choice! Red coral, with its strong apotropaic 

powers, is chosen for its propitiatory virtues to well-being, fertility, longevity and when it is set in 

a jewel, all its benefits are reinforced by other features of amulets such as the shape, pattern, text 

and so on, which vary from one culture to another.

Different tasks assigned to ornament generate diversity in the style of the jewel of East and West. 

In fact, while in the West, jewels mainly fulfill an aesthetic reason, meant to make a person look 

nicer and are expression of people’s taste and wealth, in other cultures, a jewel takes on a lager role; 

not only does it indicate one’s taste and economic status, but it also conveys symbolic messages 

connected to the history and religion of the culture in which the jewel was conceived. Nothing is 

left to chance and all symbols, shapes and materials define the person who wears them, ornaments 

are transformed into “identity cards” whose main function is to protect the wearer through the most 

delicate phases of life: birth, puberty, marriage. The ornament is then charged with magic powers, 

becoming an amulet to ward off evil, or a powerful talisman to provide wealth and prosperity.

In India, a country with huge ethnic, linguistic, and religious differences, where a billion people 

communicate in about twenty officially recognized languages, but in fact they use three hundred 

different dialects, one constant point of reference amidst the ethnic diversity is the passion for the 

ornament. Perhaps in no other country like India, jewels have satisfied the desire to adorn the body 

and the need to define membership in the cultural group, in a perpetuation of forms that have been 

handed down and repeated by the Indians for millennia.

According to the Indians, each gem has a magical and symbolic value in relation to its color 

that summarizes the emission of cosmic rays of a corresponding planet, personified in turn by a 

celestial deity: it is the Navaratna amulet, a precious jewel with nine gems which represent the nine 

celestial bodies identified with nine deities in which there is a triple association: gem-planet-deity.

Coral is associated to the deity Mangala and to the planet of Mars which is believed to govern 

the color red, the fundamental life-giving energy, and to determine kinship, strength, and sexual 

potency; coral as an amulet plays a role of mediator and will balance those aspects of life.

Also in the Tibetan-Mongolian area, coral has impressed the collective imagination so much that 

it was attributed with extraordinary scaramantic powers.

In fact, on top of the world, in Himalaya, jewels are worn stacked overlapping on clothes: in Tibet, 

Nepal, Buthan, Ladakh, those jewels are meant to resist the itinerant life of a mainly nomadic 

population. They are talismans that help protect them from a magnificent but also menacing 

landscape.

103
Similarly in Mongolia, a jewel does not only indicate one’s taste but it also conveys symbolic 

messages connected to the history and religion of the culture in which the jewel was conceived and 

coral has the protagonist role. Maybe for its sanguine red color, its indefinite origin, its durability, 

and its magic, coral fascinated this nomadic people, accustomed to the monochromy of their steppe, 

and became the sign of vital energy and for this quality it could propitiate well being, health, and 

fortune. These powers were enhanced by combining coral with turquoise, symbol of the air.

As for other nomadic populations, the Mongols invest much of their wealth in jewels and other 

precious objects, in order to be able to carry and eventually convert them into cash, for this reason 

it is not unusual to see women at work, covered with showy necklaces.

Following the route of silk, we move to Central Asia, where the scarlet gem deeply fascinated 

the nomadic peoples of the deserts. Although surrounded by mountains, this region didn’t close 

its boundaries to the western world. The route towards Asia, definitely opened by Alexander the 

Great, became one of the main euroasiatic trading routes, a network of caravan trails along which 

oases rose, offering a haven to the merchants and travelers on their way to India and China. The 

“silk route” was also the “route of coral.”

It was its color that became decisive in its evaluation and adoption. The blood-red tint, together 

with its mysterious origins and its durability, captivated these migratory peoples and came to 

symbolize the force of life. Coral began to be considered as a protective amulet or talisman, in time 

it became a metaphor for beauty and vitality. In fact, in Uzbekistan, the allegorical description of 

an ideal woman evoked “...eyebrows like bird’s wings, teeth like pearls, ears like sea-shells, lips 

like coral.”

There were also other caravan routes, which bore the precious resins and other Indian and African 

commodities to the Mediterranean and they all passed through Yemen. On these routes, exotic 

spices and cloths from India were exchanged for coral, highly priced by the peoples of southern 

Arabia for its blood red color and other virtues.

When, in 7th century, Islam took over the Arabian peninsula, it enhanced the importance of coral. 

In Sura 55 of the Koran, we read that the virgins in the garden of paradise “will be like ruby and 

coral” (LV, 58), this metaphor is very striking and gives us the full measure of coral’s importance. 

In Yemen, the ornament has to fulfill a range of functions. Certainly, it adds luster to the person 

who wears it, but it also affirms the economic standing and defines the role in society. Different 

jewels are meant for different people: a baby, a child, a young girl, or a married woman, because 

any special moment of life requires special protection, which only coral can provide.

The importance of coral in the ethnic costume and ornamentation of the Berber and Maghrebin 

peoples is just one more instance of how coral has been one of the main vehicles for common 

symbolism and meanings throughout the Mediterranean. A very fascinating symbolism can be 

found in the culture – and then in the jewels – of Morocco and Algeria, the countries bordering 

the Mediterranean that witnessed uncountable cultural mixtures, so the indigenous Berber 

population assimilated new information from all the civilizations they came into contact with: 

the Phoenicians, Greeks, Romans, Byzantines, Vandals, Arabs, Turks, Andalusians. Inevitably, 

the signs of this stratification, have been immortalized in the Maghrebin ethnic jewelry: In the 

104
techniques of granulation, for example, we find the imprint of Roman jewelers, the round tabzimt 

brooch of the Algerians Benni Yenni recalls the models of the jewelry of the Vandals, Visigoths, 

and Merovingians, then, new symbols (such as the preference for abstract motifs or the “hand of 

Fatima” amulet), together with new techniques and materials arrived with Islam, starting from the 

eighth century.

Even in the hearth of Africa the ornaments are not only decorative, but they are the image of a 

society and mirrors of cultural identity, they convey symbolic messages connected to the history 

and religion of the culture in which they are conceived.

The ethnic jewelry is an effective and synthetic symbolic message that, thanks to the use of 

particular forms, colors and materials, can tell the life of the person who stands before us, and 

reveals the creed, class and position within the community, a real status symbol to flaunt to get 

the public recognition of his role in society. In Nigeria, for example, coral is very highly valued, 

becoming sign of dignity and unquestioned power, in fact the quantity of coral worn indicates both 

the social level and the wealth of the wearer.

From East to West, backward along the routes of coral, we return to the Mediterranean and in 

particular to Torre del Greco, world famous for coral processing from the beginning of the XIX 

century. In fact, in 1805, Paolo Bartolomeo Martin, from Marseille, had the idea of founding a 

coral factory in Torre del Greco, already a thriving fishing centre, and from that point on the history 

of coral binds strongly to the history of the town in an inseparable combination that still today 

should aim to the mutual enhancement.

As with other biological resources that exist in nature and are used for commercial purposes, also 

for coral, there is the issue of sustainable management, in fact since 1994 the European Union 

has banned fishing with dragging nets. Today, throughout the Mediterranean coral is harvested by 

skilled and experienced scuba divers in a very selective way, and in quantities far less than before. 

Moreover, there is the example of the Sardinia Region where, after thirty years of regulation of 

coral fishing (www.regione.sardegna.it), scientific data have shown that in the island red coral is 

in good ecological conditions. The island is confirmed as a model in the Mediterranean area for 

the preservation, management and sustainable use of red coral, a proof that a correct use of coral 

is possible, without the bureaucratization of its trade.

This demonstrates that a proper management can reconcile the demands of safeguarding the 

precious resource, with economic requirements of the “corallari” who, among other things, must 

also ensure future generations the opportunity to pursue a craft that for centuries has contributed 

significantly to the economy of the Country.

By preserving our traditional crafts, we will not perform a merely economic operation but also and 

above all a cultural one, since coral workmanship contains our history without which we would 

lose the sense of belonging and the awareness of our deep roots.

We wish our town to receive again the role it had in the past when it was the centre of the coral 

economy, giving it credit for creating a generation of expert craftsmen who worked and transformed 

the raw material with skill and fineness, creating a jewelry that reached a wide international 

audience and that made the “Made in Italy” world famous.

105
DISTRIBUTION AND FISHERIES OF THE RED CORAL CORALLIUM RUBRUM 

(LINNAEUS, 1758) IN THE GREEK SEAS: AN OVERVIEW

Costas Dounas1*, Drosos Koutsoubas2

, Maria Salomidi3, Panayota Koulouri1,

Vassilis Gerovassileiou2,4, and Maria Sini2

1

 Institute of Marine Biology and Genetics, Hellenic Centre for Marine Research, 

P.O. Box 2214, Gournes Pediados, 71003, Heraklion, Crete, Greece

2

 Department of Marine Sciences, University of Aegean, 81100, Mytilene, Lesvos Island, Greece

3

 Institute of Oceanography, Hellenic Centre for Marine Research, 

46.7 km Athens-Sounio ave., P.O. Box 712, 19013, Anavyssos, Attica, Greece

4

 Aristotle University, School of Biology, Department of Zoology, 

P.O. Box 134, Gr-54006, Thessaloniki, Greece

*E-mail: kdounas@her.hcmr.gr

Abstract

Information, both on the spatial and vertical distribution of Corallium rubrum and its fisheries 

status in the Greek Seas, is reported in this paper, along with problems related to the conservation 

and sustainable management of this renewable biological resource. Although fisheries of red coral 

stocks in Greece have been regulated according to a national rotating harvesting system since 

1994, the difficulties experienced by both local and central authorities in confronting the illegal 

fishing of red corals in closed areas may have resulted in a decrease of the local populations. 

Available data show that in many areas the shallow water red coral stocks (up to 60 m) are now 

almost depleted. The observed evidence for the decline of red coral stocks in areas only recently 

opened up to exploitation is a clear indication that fishing (most probably by using illegal dragging 

gears) was carried out in these areas throughout the closed period. As an attempt to safeguard 

stock conservation, it is strongly recommended that an international research programme be set 

up, aiming at investigating the spatial and vertical distribution as well as the population structure 

of red coral in the Greek Seas, along with the enforcement of supplementary measures that will 

improve the effectiveness of the rotating harvesting scheme.

Introduction

The presence of the red coral Corallium rubrum (Linnaeus, 1758) in the Greek Seas, from its first 

record (Forbes 1844), has since then been reported sporadically (Dounas et al. 1989; Chintiroglou 

et al. 1989, and references therein). For a long period this species was commercially (though 

unofficially) exploited in the Greek Seas. Since the early seventies, interest in investigation and 

assessment of its populations throughout the Greek territorial waters as a new and promising 

financial resource for Greece had already been brought to the attention of the Fisheries Central 

authorities. This led the Ministry of Agriculture to establish for the first time relevant legislation 

(Greek Law 1740/1987) imposing a general framework for the exploitation of the red coral stocks. 

According to this legislation, harvesting, processing, and trade of the red coral were to be allowed 

only after the purchase of a special license. Five years later, after scientific data confirmed its 

presence in large populations in certain areas of the Greek Seas and at depths ranging from 50 

to 110 m (Chintiroglou et al. 1989), two additional laws were enacted in order to further manage 

the exploitation of C. rubrum (Presidential Decree 174/1994; Ministry Decision 240102/1995). 

106
The present study provides an updated checklist of C. rubrum records in the Greek Seas as well 

as fisheries data from the last two decades, and concludes by suggesting measures for the rational 

exploitation, conservation and sustainable management of this renewable biological resource in 

this part of the Eastern Mediterranean Sea. 

Materials and Methods 

A review of available data concerning the spatial and vertical distribution of C. rubrum in the Greek 

Seas was conducted by retrieving information from publications in scientific journals, conference 

proceedings, and grey literature (e.g. project technical reports and Ph.D. theses). Moreover, new 

and previously unpublished records from the island of Crete and the North Aegean Sea, which 

have been gathered during different research activities of the Hellenic Centre for Marine Research 

(HCMR) and the Department of Marine Sciences of the University of the Aegean, are included. 

Information concerning red coral landings during the period of 1995 to 2006 was also provided 

from General Fisheries Directory of the Greek Ministry of Agriculture.

Results

Spatial and Vertical Distribution, New Records

C. rubrum is distributed in several areas of the Greek Seas (e.g. Aegean, Ionian, Cretan, and Libyan 

Seas) (Fig. 1, Table 1). As far as bathymetric distribution is concerned, the red coral extends from 

40 to more than 120m (Fig. 2, Chintiroglou et al. 1989; Vafidis et al. 1994; Salomidi et al. 2009; 

unpublished data from the HCMR and Dept. Marine Sciences) and is usually found in caves or 

under overhangs. 

Fig. 1. Geographical distribution of Corallium rubrum in the Greek Seas based on scientific data.

107
Table 1. Population and bathymetric distribution data available for Corallium rubrum in the 

Greek Seas.

Populations Bathymetric 

Distribution Locations Source

8 colonies of C. rubrum 50-110m

N. Aegean Sea (Ierissos 

Gulf, Chalkidiki 

Peninsula, N. Evia Island 

coasts, Island of Crete)

Chintiroglou et al. 1989

22 colonies of C. rubrum 60-65m 

N. Aegean Sea (Siggitikos 

Gulf -Mount Athos, 

Alonissos and Lesvos 

Islands)

Vafidis et al. 1994

Living colonies of C. rubrum 50-91m N. Aegean Sea (E. Evia 

Island) Salomidi et al. 2009

Living colonies of C. rubrum 90-125m Crete Island HCMR-Crete unpublished 

data

Living colonies of C. rubrum 70-90m N. Aegean Sea (Agios 

Efstratios Island)

Dept. Marine Sciences 

unpublished data

Fig. 2. Colonies of the red coral C. rubrum along with the purple gorgonian Paramuricea 

clavata (Risso) in coralligenous beds of Mountain Athos coasts, North Aegean Sea (Photos by C. 

Christou).

Red Coral Fisheries 

A national rotating harvesting system covering five large fishing geographical zones was set up 

in Greek waters once the legislative framework was in place in 1994 (Fig. 3). According to this 

scheme, each area may be harvested on a maximum 5-year rotational basis, which is then followed 

by a 20-year closure period. The harvesting period in each zone can vary depending on the density 

of coral populations. 

108
Fig. 3. The five large geographical regions in the Greek Seas where the red coral is harvested on 

a rotational basis.

Overall, a maximum of 10 licenses are given each year, each costing 3000 Euros. The harvesting 

period in each year lasts from April 1 to December 31. Whenever professional coral harvesters 

find coralligenous formations, they must immediately inform the local port authorities before any 

harvesting can be undertaken. According to the national legislation, there were three harvesting 

methods for the red coral stocks in the Greek Seas. The first was manual harvesting by means of 

SCUBA diving or hookah diving “nargiles” using a metal serrated pickax which is still in force. The 

other two methods – now illegal – included the use of a specialized dragging gear equipped with 

nets of different mesh sizes: a) a metal rod called ‘ingegno’ (weight of 100-150 kg and maximum 

length of 3 m) (Fig. 4), and b) an egg-shaped piece of marble called ‘Columbus’ egg’ (weight of 

100-150 kg and maximum length of carrying nets of 10 m). The corals are firstly cracked and 

detached from the seabed and then collected by the nets as the harvesting gear is dragged from the 

boat with a rope (Figs. 5, 6). Each boat was allowed to carry only one metal rod or egg shaped gear. 

The use of Saint Andrews’ cross was always officially forbidden.

Fig. 4. Harvesting of the red coral in the Greek Seas by means of “ingegno” (Photos by C. 

Dounas).

109
Fig. 5. Sorting of nets for the red coral harvesting collected by means of destructive dragging 

gears (Photos by C. Dounas).

Fig. 6. Colonies of Corallium rubrum collected by professional fishermen in the Greek Seas 

(Photos by C. Dounas).

Until now, only three out of the five geographical zones of the Greek Seas have been harvested (N. 

Aegean, S. Aegean-Island of Crete, and Ionian Seas). The total official recorded landings of red 

corals per region harvested over the period 1995-2005 are presented in Fig. 7. Overall, landings of 

red corals amount to 22.9 tones over this period, while annual landings seem to decrease gradually 

in each of the zones being harvested. Total landings from the N. Aegean Sea are twice as high as 

those recorded from the Cretan and Ionian Seas taken together, while landings from the coasts of 

Crete are the lowest. According to the available fisheries data, there is a significant decrease of the 

shallow water red coral stocks (up to 60 m) in these zones towards the end of the harvesting period. 

Depletion of red corals in the shallower areas, where harvesting is allowed, has currently forced 

professional divers to harvest colonies at greater depths (up to 125 m) by means of advanced 

mixed gas diving equipment, as today this is the only effective legal fishing method.

110
0

1000

2000

3000

4000

5000

6000

7000

8000

1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006

Weight (kg)

Zone I

(North Aegean)

Total landings:

15.2 tn

Zone III (Central Aegean)

Zone IV

(Cretan & Libyan 

Sea)

Total landings:

1.2 tn

Zone V

(Ionian Sea)

Total landings:

6.5 tn

Fig. 7. Red coral landings in the Greek Seas over the period 1995–2006 (source: Greek General 

Fisheries Directory). 

Discussion

According to the available data (both scientific records and official landings) the most abundant 

populations of C. rubrum in the Greek Seas are found in the N. Aegean Sea. This should probably 

be attributed to the eutrophic conditions of N. Aegean Sea which are quite similar to those of the 

Ionian Sea and parts of the NW Mediterranean where the species is generally thriving. However, the 

Cretan Sea, along with the Libyan Sea, is dissimilar and can be grouped with the rather oligotrophic 

Levantine Sea (e.g. Voultsiadou 2005, 2009; Bianchi 2007). In addition, current research has 

revealed that the N. Aegean Sea is characterized by rich biodiversity and hosts complex hard 

substrate communities such as coralligenous formations and bio-concretions as well as a large 

number of marine caves (Antoniadou et al. 2005; Chintiroglou et al. 2005; Gerovasileiou et al. 

2009). The lack of scientific records from the central part of the S. Aegean Sea (e.g. the Cyclades 

plateau) is worth mentioning, despite the fact that extended deep coralligenous banks have been 

reported from this area (Laborel 1961; Jacquotte 1962; Georgiadis et al. 2008). However, the 

opening of this zone for the harvesting of red corals (from the summer of 2009) will probably 

provide new data on populations of C. rubrum. The review of C. rubrum records in the Greek Seas 

showed that knowledge concerning its distribution in these waters is far from being considered 

as complete. Further research on hard substrate communities, and especially on specific habitats 

(e.g. deep coralligenous banks and semi-obscure caves), which are either difficult to approach or 

not even accessible by scientific diving, is expected to reveal new data regarding the distribution 

and abundance of the species in this part of the Mediterranean Sea. Moreover, the significant 

increase in recreational diving activities in Greece, which was prohibited in most coastal areas 

until 2005, could provide additional information from previously unexplored marine areas within 

the shallower zone for diving. 

111
The official fisheries data on the red coral in the Greek Seas show a decrease in shallow water 

populations, which seems to be closely related to the fishing effort (there are fewer landings towards 

the end of the fishing period in each large geographical zone). Another major issue, related to the 

depletion of red coral stocks in the Greek Seas, is the intrinsic difficulties experienced by both 

local and central authorities in confronting the illegal fishing of this renewable biological resource. 

There is growing concern in the red coral fisheries sector that fishing by unauthorized vessels, 

illegally using destructive dragging gears (e.g. Saint Andrews’ cross), has increased considerably. 

A possible reason for this practice could be the high cost of purchasing and maintaining the legal, 

though often highly sophisticated and expensive harvesting equipment (e.g. mixed gas apparatus). 

The observed decline of red coral stocks in areas which were only recently opened up to exploitation 

is an indication that harvesting was carried out illegally throughout the closure period. It is obvious 

that new effective measures for the control and monitoring of landings are urgently needed in the 

Greek Seas for the conservation and sustainable management of this renewable biological resource. 

It is therefore suggested that certain additional measures should be included in the existing national 

legislation. For instance, red coral landings should be officially registered and tagged with regard 

to the harvesting place and time, while corals should be stored in sealed bags and identified by 

the local authorities. Management measures should also include penalties against the harvesting 

of small colonies (basal diameter less than 10 mm), as well as penalties for buyers and fishermen 

who possess even a small quantity of unprocessed corals without the required documentation. 

Furthermore, the large rotating harvesting zones should be examined separately in detail in order 

to assess the effectiveness and viability of the present harvesting scheme (e.g. number, size, and 

borders of the harvesting zones, optimal temporal and spatial orders for the opening of different 

zones). The optimal rotation period should remain constant instead of fluctuating according to 

fishermen’s pressures and monitoring of landings should define the time of closure. 

Apart from management measures based on fisheries landings, it is clear that there is a need for 

further scientific research in the Greek Seas aiming at a detailed investigation of the size and 

structure of the red coral populations. This approach should also take into consideration that C. 

rubrum, apart from its highly acknowledged commercial value, also plays an important ecological 

role. C. rubrum along with other gorgonian species distributed in the Mediterranean Sea, comprise 

ecosystem engineering key-species which provide important three-dimensional complexity to the 

coralligenous habitats and host high biodiversity (Boudouresque 2004; Ballesteros 2006; Giaccone 

2007). For example, colonies of C. rubrum from the N. Aegean Sea and the coastal area around 

the island of Crete have been found to host a large number of epibenthic faunal species, such as 

the sponges Geodia conchilega (Schmidt), Dysidea fragilis (Montagu), Erylus sp., Axinella guiteli

Topsent, Agelas oroides (Schmidt), Petrosia ficiformis (Nardo), Ircinia dendroides (Schmidt) and 

I. foetida (Schmidt), the cnidarians Sarcodictyon roseum (Philippi) and Corynactis sp., the serpulid 

polychaetes Vermiliopsis infundibulum (Philippi), Placostegus crystallinus (Zibrowius) and Janua 

pagenstecheri (Quatrefages) and the molluscs Calliostoma zizyphinum (Linnaeus), Pseudosimnia 

carnea (Poiret), Heliacus alleryi (Seguenza), Coralliophila brevis (Blainville), Mitra zonata

(Marryat), Lopha stentina (Payraudeau) and Ostrea edulis (Linnaeus) (Chintiroglou et al. 1989; 

Vafidis et al. 1994; Koutsoubas et al. 1997).

In conclusion further scientific research, detailed mapping and a regular monitoring scheme 

along with measures for the protection of the coralligenous structures and especially for the C. 

rubrum populations in the Greek Seas are considered to be necessary, in order to safeguard their 

conservation and avoid extensive loss caused by environmental or anthropogenic pressures. 

112
Acknowledgements

The authors would like to thank the officers of the Greek General Fisheries Directory for providing 

information on red coral landings and Mrs. Margaret Eleftheriou for the critical review of the 

manuscript.

References

Antoniadou, C., Chintiroglou, C. 2005. Biodiversity of zoobenthic hard-substrate sublittoral 

communities in the eastern Mediterranean (North Aegean Sea). Est. Coast. Shelf Sci. 62: 

634-653.

Ballesteros, E. 2006. Mediterranean coralligenous assemblages: a synthesis of present knowledge. 

Oceanogr. Mar. Biol. Ann. Rev. 44: 123-195.

Bianchi, C.N. 2007. Biodiversity issues for the forthcoming tropical Mediterranean Sea. Hydrobiologia 

580: 7-21.

Boudouresque, C.F. 2004. Marine biodiversity in the Mediterranean: status of species, populations 

and communities. Sci. Rep. Port-Cros Natl. Park 20: 97-146.

Chintiroglou, C., Dounas, C., Koukouras, A. 1989. The presence of Corallium rubrum (Linnaeus, 

1758) in the eastern Mediterranean Sea. Mitt. Zool. Mus. Berl. 65 (1): 145-149.

Chintiroglou, C., Antoniadou, C., Vafidis, D., Koutsoubas, D. 2005. A review on the biodiversity of 

hard substrate invertebrate communities in the Aegean Sea. Medit. Mar. Sci. 6(2): 51-62.

Dounas, C., Eleftheriou, A., Koukouras, A. 1989. The presence of Corallium rubrum (Linnaeus, 

1758) in Greek waters. A historical review. GFCM technical consultation on red coral of 

the Mediterranean, Torre Del Greco, 27-30 September 1988. FAO Fisheries Report No 413.

Georgiadis, M., Papatheodorou, G., Tzanatos, E., Geraga, M., Ramfos, A., Koutsikopoulos, C., 

Ferentinos, G. 2009. Coralligène formations in the eastern Mediterranean Sea: morphology, 

distribution, mapping and relation to fisheries in the southern Aegean Sea (Greece) based on 

high-resolution acoustics. J. Exp. Mar. Biol. Ecol. 368: 44-58.

Gerovasileiou, V., Sini, M.I., Poursanidis, D., Koutsoubas, D. 2009. Contribution to the knowledge 

of coralligenous communities in the NE Aegean Sea. Proceedings of the 1st Symposium on 

the Coralligenous and other Calcareous Bio-Concretions of the Mediterranean Sea, Tabarka, 

15-16 January 2009, 205-107.

Giaccone, G. 2007. Coralligenous assemblage as underwater seascape: distribution off Italian coasts. 

Biol. Mar. Medit. 14 (2): 124-141.

Jacquotte, R. 1962. Etude Des Fonds Des Maerl De Méditerranée. Recl. Trav. Stn. Mar. Endoume 

26: 141-235.

Koutsoubas, D., Koukouras, A., Voultsiadou-Koukoura, E. 1997. Prosobranch mollusc fauna of the 

Aegean Sea: new information, check list, distribution. Isr. J. Zool. 43 (4): 19-54.

Laborel, J. 1961. Le Concretionnement Algal “Coralligène” Et Son Importance Géomorphologique 

En Méditerranée. Recl. Trav. Stn. Mar. Endoume 23 (37): 37-60.

113
Salomidi, M., Smith, C., Katsanevakis, S., Panayotidis, P., Papathanassiou, V. 2009. Some observations 

on the structure and distribution of gorgonian assemblages in the eastern Mediterranean 

Sea. Proceedings of the 1st Symposium on the Coralligenous and other Calcareous BioConcretions of the Mediterranean Sea, Tabarka, 15-16 January 2009, 242-244.

Vafidis, D., Koukouras, A., Voultsiadou-Koukoura, E. 1994. Octocoral fauna of the Aegean Sea with 

a check list of the Mediterranean species: New Information, Faunal Comparisons. Annal. 

Inst. Océanogr. Paris 70(2): 217-229.

Voultsiadou, E. 2005. Demosponge distribution in the eastern Mediterranean: a NW-SE gradient. 

Helg. Mar. Res. 59: 237-251.

Voultsiadou, E. 2009. Reevaluating sponge diversity and distribution in the Mediterranean Sea. 

Hydrobiologia 628:1-12.

114
THE RED CORAL EXPLOITATION IN MOROCCO 

Abdelmajid Dridi, Abdelaziz Zoubi, and Said Taleb

National Institute For Fisheries Research (INRH), 2- Rue Tiznit- Casablanca, Morocco

Introduction

Morocco is endowed with two coastlines, the Atlantic and the Mediterranean, with very important 

red coral and varied fisheries resources. This typically Mediterranean species of red coral found at 

both regions an ideal biotope to develop.

In Morocco, red coral harvesting began at the Mediterranean as a special activity at the 

Mediterranean in the 1970s, by foreign boats. In the 1980s, this activity took an important rise, 

in particular in the area of Al Hoceima at the Mediterranean side with great red coral abundances 

with red coral colonies of big sizes and good quality. In 1990, the activity of fishing of the coral 

was located in Asilah site at the Atlantic zone extending between Cape Spartel and Larache, where 

was discovered an exploitable red coral stock starting from 40 meters of depths. After 13 years 

of exploitation, the zone of Asilah was closed for coral exploitation. Since 2004, the zone of Al 

Hoceima was again opened for red coral harvesting.

Red Coral Exploitation 

The first gear used in the exploitation of red coral was the cross of Saint Andrew, which is pulled 

on sea-bed, breaks the coral colonies and causes consequently a degradation and an exhaustion of 

the red coral layers on the one hand, and affect the biodiversity and the marine ecosystem as whole 

on the other hand.

This damaging technique of exploitation presents a production of bad quality product in spite 

of the great harvested quantities. However, during the 1980s, this technique was prohibited in 

Morocco and replaced by a new technique based on liquid bottles which is called deep-sea diving. 

This technique became the single technique practised for red coral harvesting. The tool of divers 

consists of a net basket and a sharp marteline on board which makes it possible to cut the foot or 

coral colony.

Operating Fleet 

During the period 1984-1987, the number of boats in activity varied from 2 to 17, totally of foreign 

nationalities. Since the freezing of the investments in 1992, the number of coral boats was set to 

10 units.

115
Harvesting Zones 

The exploitation of the red coral was carried for a period varying from 10 to 12 years of 

exploitation per area. Between 1980 and 1991, two principal fishing zones can be considered in 

the Mediterranean, as shown in Fig. 1.

Fig. 1. Red coral stocks in Moroccan coast.

•	 The zone called TOPOS is a very coastal zone in which the coral resources is very close 

to the surface (30–60 meters depth). In this zone, the red coral harvesting is only done by 

diving under compressed air. However, these zones with low depth are subjected to an 

intense harvesting fishing activity owing to the fact that the diver can remain under water 

for a long period of time (45 to 60 min), and the operation of diving can be done twice per 

day without risk.

•	 The zone known as TOFINOS, moving away from the nautical coast of approximately 24 

miles, is a very deep, rocky zone, characterized by very strong currents. The coral is on 

the sides of rocks beyond 80 meters depth. Contrary to the zones of low depths, in these 

zones of great depths, the duration of diving is very limited not exceeding a maximum 20 

to 25 min because of the mixture of gas (helium and oxygen). The diver must obligatorily 

follow and respect the instructions of the rules of the diving; if not it can endanger its life.

116
Since 1992, the Mediterranean area was not interesting any more as the collected colonies were of 

small sizes and consequently their economic value had fallen; thus, the exploitation of the coral 

moved towards the Atlantic area on the level of the Asilah region extending from Cape Spartel in 

Larache. This zone is made up of three red coral layers: (1) a layer at the level of Cap Spartel, (2) 

another layer at the level of Asilah, and (3) a layer at the level of Larache. The choice was put on 

the layer of Asilah owing to the fact that the red coral can be observed starting from 35 meters of 

depth, characterized by good visibility for divers, along with weak marine currents, and thus more 

favorable exploitation conditions.

Production

The exploitation of coral at the Mediterranean during the 1980s saw a total production in Al 

Hoceima in 1985 of 8.6 tons. After three years of exploitation, this production recorded a fall of 

80% to reach 1.6 tons in 1988. The drastic fall of the production during a tiny time, colonies of 

small sizes proved that coral layer at the Mediterranean was subjected to an overexploitation.

Year Zone Exported Quantities (tons)

1992 Asilah (100%) 9.6

1993 Asilah (100%) 6.2

1994 Asilah (100%) 6.0

1995 Asilah (100%) 4.3

1996 Asilah (100%) 2.8

1997 Asilah (100%) 2.0

1998 Asilah (100%) 1.9

1999 Asilah (100%) 4.7

2000 Asilah (100%) 7.7

2001 Asilah and Al Hoceima 

97%:::::::::::::::3% 8.0

2002 Al Hoceima (100%) 22

2003 11

2004 Al Hoceima (100%) 6

2005 2.5

2006 Prospection- Mediter. 1.6

2007 2

 Source: Change Office 

117
The analysis of the great variations of the production or of the level of exports from one year to 

another show that the rise can be due to the exploitation of deeper zones than usual or the addition 

of quantities already harvested the previous year.

Conclusions

Generally, the exploitation of the red coral in Morocco, whether at the Mediterranean or at the 

Atlantic, by the means of the production carried out each year shows the same tendency of evolution. 

The production reaches a maximum after the opening of the zone of activity and starts to fall 

progressively. This downward trend is due mainly to the discontinuous geographical distribution 

of the red coral and/or to the manner with which the exploitation is done.

The exploitation of the red coral starts on the level of the layers of low depths where abundance 

is more or less high, the production can oscillate because of the rarefaction of the species, or to 

several environmental factors which would prevent the realization of good coral harvest, mainly 

the terrigenous contributions, turbidity, lack of visibility for divers.

The exploitation continues towards great depths where the coral appears on the exposed faces of 

the rocks. The exploitation of these depths requires qualified divers. 

118
RED CORAL AND ITS ACTUAL SITUATION IN TURKEY

B. Öztürk

Faculty of Fisheries, Istanbul University

Red coral is known as one of the precious coral species by Turkish coral divers and fishermen. 

This species is caught in the northern Aegean Sea and Turkish Mediterranean Sea until 1990. 

Accurate data for the amount of catch and prices during the last period, however, is not available. 

This species was caught below 40 meters depth banks and marine caves mostly between Anamur 

and Kaş.

Since 1990, according to the Turkish Fishery Law 1380, red and black coral species have been 

under protection continuously. The reason of this decision was that coral divers destroy other 

benthic species, such as slipper lobster, European locust lobster, and spiny lobster, as well as some 

fish, such as dusky grouper and grouper, in deep seas and caves.

Actually, red coral Corallium rubrum is imported from some countries as jewelry materials in 

Turkey. The amount is not known due to its very small quantity. 

A global assessment for the Mediterranean Sea is needed for the sustainable resource management. 

Local people’s income and opinion for the management of the coral resource should be also 

considered.

119
ANALYSIS OF FAO STATISTICS OF HARVEST OF CORALLIUM RUBRUM

 IN THE MEDITERRANEAN 1978–2007

M. Pani

Wildlife Management Consultant 

FAO Fishery statistics are presented and linked to the introduction of major legislations to regulate 

harvest of Corallium rubrum in the Mediterranean.

It is important to note that the supposed decline is based on an analysis of harvest data produced 

by FAO. 

It is worth noting that in last 10 years period no decline of the harvest has been noted and that, 

more importantly, change in harvest techniques has greatly reduced the fishing efforts for the 

species. In fact, after the ban of the “ingegno” or of “Saint Andrews cross” (1994 EU regulation 

1626/94), harvest is exclusively done by SCUBA and only by professional divers who harvest the 

species between 60 and 110 m of depth with clear technical difficulties. 

In fact, diving time at such depths is very limited (maximum 20 minutes), and also the quantities 

that can be harvested are limited. Only a hammer can be used according to the management 

measures and legislation of many countries.

Spain, France, Morocco, Algeria, Tunisia, and Italy (Sardinia) have enacted specific legislation 

on the red coral harvesting. In particular, the Sardinia Region has banned the use of non-selective 

gears since 1989 and intermittently banned their use since 1979 when the first legislation on coral 

harvesting was enacted. 

This explains the decline of the statistical figures.

In fact, of the 100 tons harvested in 1978 that represents the peak of the harvest for the 

Mediterranean, 72 tons were harvested in Sardinia. The legislation introduced in Sardinia in 1979 

and the intermittent ban on non-selective gears, together with high taxes imposed on the fishing 

vessels, explain the decline of harvest from 1979 to 1989 when the use of non-selective gears 

was definitively banned in Sardinia, five years before the entry into force of 1994 EC Regulation 

1626/94, which banned the use of non-selective gears for the coral harvest in the Member Sates of 

the European Union.

Therefore, the statistical decrease is linked to the decrease in fishing efforts and not to the decline 

of the species. In the last 30 years the fishing efforts on C. rubrum in the Mediterranean has 

considerably decreased. This was due to the ban on the use of the “ingegno” and of the Saint 

Andrews Cross (two different non-selective gears) and to the fact that in many countries fishing 

is exclusively done by licensed divers that can harvest the species in limited quantities only and 

only between 60 and 100-110 m of depth. The populations below that depth are completely intact. 

Moreover, official statistics do not take into account the finding of dead coral banks generated by 

the detachment of old or sick coral transported by streams.

120
IL COMPARTO DEL CORALLO DI TORRE DEL GRECO

P. Stampacchia 

Dipartimento di Economia Aziendale, Università degli Studi di Napoli “Federico II” 

Il comparto della lavorazione dei coralli, cammei ed oreficeria di Torre del Greco si caratterizza 

per alcuni elementi che lo rendono unico nel panorama produttivo nazionale, anche nell’ambito 

dei distretti analoghi di Valenza Po, Vicenza ed Arezzo.

1 – Ha una tradizione storica che risale alla fine del ‘700, che ancora si riscontra nei caratteri 

tipicamente artigianali, individualistici e, però, anche artistici della lavorazione. In questo contesto 

l’innovazione tecnologica e la delocalizzazione potrebbero rappresentare, addirittura, degli 

elementi di eradicamento dalla storia e dal territorio, piuttosto che dei fattori di sviluppo.

2 – Si basa su di una rete di relazioni interne ed internazionali particolarmente fitta ed articolata: 

a livello locale molte imprese operano per altre imprese vicine con una divisione complessiva del 

lavoro che dà spazio alla creatività degli artigiani ed alle capacità di percorrere i mercati internazionali 

e seguirne le tendenze in maniera proattiva. (Organizzazioni di rete degli approvvigionamenti, 

delle relazioni interne al comparto e delle vendite internazionali)

3 – É strettamente legato all’immagine turistica complessiva dell’Italia artigiana al punto che 

prodotti tipici dell’area vengono offerti e ricercati dai turisti stranieri sia nelle aree campane 

(Sorrento, Pompei, ecc.), che in altre tipiche aree turistiche italiane (Venezia, Firenze, ecc.).

4 – Grazie alle relazioni internazionali (soprattutto con USA, Giappone ed Estremo Oriente) ha 

ampliato la gamma di prodotti oggetto di attività produttiva e commerciale. È diventato negli 

anni uno dei principali centri italiani sia per l’oreficeria, che per il commercio internazionale e la 

lavorazione delle perle.

Pur essendosi verificata anche a Torre del Greco una contrazione delle imprese e, ancor più, degli 

addetti, proprio i caratteri tradizionali del settore ne hanno guidato il riorientamento nel corso degli 

ultimi anni. 

Attualmente nel comparto torrese operano circa 270 imprese con oltre 2000 addetti ed il mancato 

riconoscimento delle attività locali come distretto (con la possibilità di attingere ai relativi 

benefici) è derivato non dalla carenza degli elementi strutturali e dinamici tipici di queste realtà, 

ma, piuttosto, dal fatto di essere fortemente radicate all’interno di un’area urbana ad alta densità 

abitativa, il che ancora una volta evidenzia uno dei punti di forza del settore nella forte sinergia, da 

un lato, tra vita quotidiana e produzione e, dall’altro, tra produzione e territorio.

Prima ancora della crisi attuale, come in altri settori, si è manifestata nell’oreficeria una rilevante 

dicotomizzazione della domanda tra prodotti di basso prezzo/qualità e prodotti di medio-alto 

prezzo/qualità. La domanda di prodotti del primo tipo trova soddisfazione maggiore nelle offerte 

a più basso prezzo basate su produzioni orientali; la domanda di prodotti di livello medio-alto 

trova riscontro nei caratteri di artigianalità, creatività artistica e peculiarità dei materiali oggetto 

di lavorazione a Torre del Greco: il corallo e le capacità di lavorazione del corallo; i cammei e 

121
le capacità di lavorazione dei cammei rappresentano elementi trainanti di una offerta unica nel 

panorama mondiale basata su forti elementi di cultura e competenze.

Basta notare che nell’indagine svolta nel 1999 il 53% degli intervistati considerava l’orientamento 

delle consumatrici tra i fattori considerati nella progettazione e realizzazione dei prodotti, mentre 

il 93% degli stessi operatori inseriva fra i fattori considerati la qualità della materia ed il 40% la 

forma.

Nonostante le evoluzioni più recenti, quindi, la produzione rimane fortemente legata ad una 

specifica materia prima – il corallo –, alle competenze ed alle relazioni che attorno ad essa ed a 

partire da essa si sono diffuse e consolidate per altri materiali ed aree. Si parte da una esperienza 

di navigazione, le famose “coralline”, e si ritorna ad esperienze di relazioni internazionali che 

hanno il corallo come matrice originaria ed, ancora oggi, fonte di creazione di competenze di 

approccio di capacità di gestione responsabile delle materie prime, di lavorazione, di distribuzione 

e di commercio internazionale applicate ad altre materie ed aree con gli stessi principi di creatività 

ed arte. 

L’eventualità di introdurre elementi di burocratizzazione del commercio che porti a forti riduzioni 

del corallo fra le materie oggetto di lavorazione non sarebbe, quindi, come togliere gli oggetti in 

oro bianco dalla gamma di prodotti di un orefice, ma sarebbe come togliergli la fonte e la ragione 

stesse della formazione delle sue competenze, che nel tempo lo hanno portato ad occuparsi di altri 

materiali e di nuove aree di sbocco. 

122
CORAL AS PRODUCT: ANALYZING PRODUCTION 

AND MARKET PERSPECTIVES

S. J. Torntore

University of Idaho

Moscow ID USA

E-mail: torntore@uidaho.edu

This paper will examine the production of and markets for coral products in the two largest coral 

sectors in the world – Torre del Greco and Taiwan – using a global commodity chain perspective. 

In both coral sectors, different products are made for different markets. Very particular destinations 

and target markets were identified by coral producers in both sectors for specific types of coral 

products. The coral commodity chain is a complex global chain of value-added commercial 

production from raw material to final marketed commodity, in which market and end use for 

the product determine the production processes. Production methods and industry drivers in 

both sectors will be characterized and compared. Market categories will be identified based on 

1) characteristics of the raw materials, 2) characteristics of the finished products, 3) values of 

coral, and 4) meanings and cultural needs. Examining coral production and products from a 

commodity chain perspective tracks how the value of a coral object is increased at each stage of 

its production for specific markets, in a process of production that transcends national boundaries. 

These perspectives can provide not only a starting point to develop approaches for environmental 

sustainability of managing coral trade but provide insight to develop targeted approaches for 

sustainability of production and market sectors in an industry facing the impacts of international 

trade regulations.

123
Red coral (Corallium rubrum L., 1758) management

strategies in Sardinian coastal waters

 M. Campolmi, R. Caddeo, M.C. Soro, R. Doneddu

Regional Government - Agriculture Department - Fisheries Service 

via Pessagno, 4 - 09126 Cagliari - Italy

E-mail: agr.pesca@regione.sardegna.it

Abstract

Red coral in Sardinia is a historically important resource, coral harvesting being documented since 

the XIII century. Since 1979, coral fishing in the island has been subject to specific regulation 

(Regional Laws: 5th July 1979 n. 59 and 30th May 1989, n. 23) that introduced several restrictions 

in order to improve sustainable exploitation, conservation of coral resources, and protection of 

marine ecosystems. It provides the main following rules: coral harvesting only be conducted by 

professional fishermen in possession of a regional permit that is renewed annually, and fishing 

tools limitation, since 1989 towed gears prohibition and only manual harvesting with hatchet 

permitted. Furthermore, every year on the basis of scientific evidence harvesting effort (maximum 

permit numbers, harvesting time, allowed areas, minimum size of coral basal diameter, and daily 

maximum amounts of coral per fisherman) is established. Daily harvesting data have to be provided 

by fishermen. Since the 1980s, the Regional Administration has been funding scientific studies in 

order to collect biological data and obtain periodical coral stock assessment along Sardinian coastal 

waters. The management strategy adopted in Sardinia is one of the first in the Mediterranean 

Sea and seems to represent a successful experience, having achieved a good balance between 

protection and exploitation requirements as confirmed by recent scientific evidences.

Introduction

Red coral (Corallium rubrum L., 1758) in Sardinia is an historically important resource from a 

biological, economical, and cultural point of view, due to the richness and high quality of coral 

banks mainly distributed along the west coast. 

Coral harvesting has been going on for several centuries in the island, being documented since the 

XIII century (Doneddu and Fiori 2003, reported by Cannas et al., in press).

At present, red coral exploitation involves about 250 people (unofficial data) including fishermen, 

craftsmen, and tradesmen; it represent a remarkable yielding sector, especially for cultural and 

historical reasons mainly connected with a well-known processing tradition. The town of Alghero, 

a little city on the North West cost of the island, is famous in Europe for its coral carving tradition 

and for the high quality of red coral handmade products. 

Today, the estimated number of people involved in coral harvest, processing, and trade include 

roughly (Fig. 1) 170 between craftsmen and tradesmen, 35 crew assistants, 25 allowed fisherman, 

and only two wholesales (unofficial data referred to the year 2009) from Torre del Greco – a 

town located in the Campania Region that today represents the most important coral trade and 

processing site in Italy.

124
Fig. 1. Estimated number of people involved in Sardinian coral harvest processing and trade in 

2009 (unofficial data).

In Italy, since 1965, National Law 14th July 1965, n. 963 “Maritime Fishing Regulation,” 

implemented by President of the Republic’s Decree (DPR) 2nd October 1968, n. 1639, established 

specific regulations for the exploitation of discovered coral banks (Fig. 2).

In particular the above mentioned Law established that fisherman who finds a new coral bank has 

to declare the discovery to marine authority in order to be allowed to its exploitation for the next 

two years (Art. 16).

Furthermore, President of the Republic’s Decree (DPR) 2nd October 1968, n. 1639, issued for 

implementation of this Law, specified that discovery declaration of a new bank has to include 

personal data of the fisherman, day of discovery, position and extension of the bank (Art. 123) and 

that fishing season lasts all the year (Art. 124).

Since 1948 Sardinia is a self-government region with a Special Statute (Constitutional Law 26th

February 1948, n. 3)

Sardinian Administration has primary legislative function on fishery items (D.P.R. 24.11.1965, n. 

1627).

So according to the National Law 963/1965, the Regional Administration from 1965 to 1979 

had been issuing decrees to allow responsible exploitation of discovered coral banks within the 

territorial waters.

25 35

2

170

15

Allowed fishermen

Crew assistants

Wholesales

Craftmen, tradesmen

Boat and equipment

maintenance

125
In 1979, Sardinian Regional Council issued the Regional Law n. 59 – afterward integrated and 

modified by the Regional Law 23/1989 (Fig. 2) – one of the first experiences of specific regulations 

in the Mediterranean Sea, that has ruled harvesting effort management on red coral banks, ensuring 

an effective protection of the resource.

Fig. 2. National and regional regulation.

Materials and Methods

Harvesting data, provided to the Regional Administration by fishermen, were analysed from 1996 

to August 2009 in order to evaluate the effect of regional regulation on coral resource and harvesting 

effort. Total allowed harvesting days were obtained on the basis of the harvesting period fixed by 

regional decree. The number of harvesting days per fisherman was obtained dividing the total 

number of effective fishing days (provided by coral divers) by the number of permits effectively 

used. Daily coral amount was calculated dividing the total annul coral amount by the total number 

of effective harvesting days (obtained as the sum of annual harvesting days of all coral divers).

Results 

Regional Laws

The Regional Law 5th July 1979, n. 59 “Coral Fishing Regulation” introduced several restrictions 

in order to improve sustainable exploitation, conservation of coral resources, and protection of 

marine ecosystems. It includes the following regulations:

Art. 1 – In Sardinian coastal water, red coral harvesting can be conducted by professional fishermen 

in possession of a regional permit, with the exception of the cases read in Article 4.

126
Art. 2 – Regional permit must be renewed annually and is issued or suspended by means of regional 

councillor’s decree. 

Art. 3 – Coral harvesting is allowed only with the following gears: 

a) hatchet, used by professional fishermen equipped with scuba-diving devices;

b) Saint Andrew’s cross*;

c) Ingegno*, maximum length: 3 m, maximum number per boat: 1 

* cross shape towed gears equipped with pieces of net

Catch methods different from those above mentioned are forbidden.

Art. 4 – Annually, within January, in compliance with the Regional Council decision and once 

the opinion of the Regional Technical Advisory Committee is acquired, the councilor’s decree 

establishes:

a) harvesting time;

b) maximum daily amounts of coral per each harvesting gear;

c) allowed areas;

d) deadline and conditions for requests submission; 

e) the fee amount for each harvesting gear;

f) maximum permit numbers for each harvesting gear.

Harvesting permit can be issued in order to allow scientific research.

Art. 5 – For ecological and biological protection, requirements the councilor’s decree can forbid 

coral harvesting in specific areas for a minimum period of 3 years.

Art. 6 – Every year coral fishermen have to provide daily harvesting data (coral amounts, position, 

and depths of fishing areas) within 30 days from the end of harvesting time.

 Art. 7 – The Regional Administration provides for collecting scientific data on red coral banks in 

order to ensure sustainable exploitation charging:

a) scientific research institutes;

b) specialized private enterprises or local government.

Art. 8 – The councillor’s decree, in compliance with the Regional Council decision, can: 

a) organize or fund specific professional training classes about coral harvesting with the use 

of scuba-diving devices;

b) grant funds for young Sardinian people to attend the above mentioned professional training 

classes.

Art. 9 and Art. 10 – Establish the administrative sanctions in case of violation of the Law.

Afterwards, to ensure a more strict and effective protection of the coral resource and the marine 

ecosystems the Regional Law 30th May 1989, n. 23 – partly modifying the Regional Law 5th July 

1979 n. 59 – introduced the following further restrictions: 

127
Art. 1– Only manual harvesting with hatchet is permitted to professional fishermen equipped with 

scuba-diving devices. All the other fishing methods are banned.

Art. 2 – Financial aids are provided to sustain transition from destructive dragging methods (Saint 

Andrew’s cross and Ingegno) to allowed gears (hatchet). Contribution until 70% are given to 

fishermen allowed to non-selective gears in 1988 and 1989 to purchase new fishing equipment.

Regional Management Actions

According to Article 4 of the Law n. 59/79, yearly, on the basis of the scientific evidences, a 

councilor’s decree establishes the value of the harvesting variables and further limitations in order 

to ensure the protection of the species (Table 1). 

Table 1. Regional management actions: main harvesting variables fixed by regional annual 

decrees.

2001 2002 2003 2004 2005 2006 2007 2008 2009

Total allowed 

harvesting days 199 199 199 199 188 92 0 111 168

Number of 

permits 31 32 37 41 38 31 0 20 30

Minimum 

allowed 

harvesting depth 

(m)

0 0 0 0 0 0 0 80 80

Maximum red 

coral amount 

per day (kg)

4 4 4 3 3 2,5 0 2,5 2,5

Minimum basal 

diameter (mm) 10 10 10 10 10 10 0 10 10

Tollerance on 

minimum basal 

diameter (%)

30 30 30 30 30 20 0 20 20

The minimum basal diameter of 10 mm is fixed with a tolerance of 20% (basal diameter between 8 

and 10 mm); furthermore, new-harvested red coral has to be maintained in sea water for a minimum 

of 30 minutes in the net (size ≥ 5 mm) in order to allow the dispersion of gametic products.

128
Analysing the main harvesting variables fixed annually by the Regional Decree in the last decade 

is possible to highlight the trend of the catch effort management established on the basis of the 

scientific evidence (Table 1). 

Generally, the harvesting period has been fixed from spring to autumn, but the number of total 

allowed harvesting days has changed greatly, especially in the last decade (changing from 0 to 199 

days), as shown in Fig. 3 and Table 1. 

Since 2006, the Regional Government has chosen to reduce the harvesting period, the number of 

permits and the daily maximum red coral amount per fisherman (Fig. 3, Table 1). 

In 2007, the Regional Government decided to forbid red coral fishing, applying the precaution 

principle to better preserve the biological resource assessing the need of deepening scientific 

studies on stocks condition in Sardinian coastal water.

Since 2008, a minimum harvesting depth has been established allowing the harvest only at depth 

> 80 meters, in order to preserve the shallower colonies. 

Furthermore, in 2008, the Regional Government allocated funds for a multi-annual restocking 

program (Decree 19th December 2008 n. 3189/DecA/7108), at the moment in a start-up phase.

In 2009, the Regional Council, considering the results of scientific researches, assessed that an 

enlargement of the permits number and a longer harvesting period were sustainable. 

0

50

100

150

200

250

200120022003200420052006200720082009

Number of allowed harvesting days 

0

5

10

15

20

25

30

35

40

45

N° of permits 

Number of

permits 

Number of

allowed

harvesting

days

Fig. 3. Trend of Regional Regulation: number of coral licences and allowed harvesting periods.

129
In 2009, 25 permits were issued, involving about 16 fishing boats (Table 2), coral harvesting were 

allowed from the 1st of May to the 15th of October. 

At present, as well as in the past, the main harvesting areas are located along the Western coast, 

especially in the Northern part (Fig.s 4, 5, 6). Fig. 4 shows also the position of the areas in which 

coral harvesting is presently forbidden due to the strong impact of the towed gears used in the past 

on those ecosystems. 

The distribution of the harvested coral amount in different conditions (dead, mixed, spoilt, and red) 

among the harvesting areas during 2008 is reported in Fig. 7.

Table 2. Recent figures.

2008 2009 (until August)

Number of issued permits 20 30

Number of effective permits 20 25

Number of fishing boats 17 16

Total red coral harvested 

(Kg) 1120 1457

Range of trade price declared 

by coral divers (€/kg) 500 - 1000 400 - 800

Current trade price for 

mixed quality coral declared 

by coral divers (€/kg)

500 450

130
Fig. 4. Coral harvesting areas (map scale 1: 1.150.000).

131
Fig. 5. Geographic distribution of annual red coral amounts in 2008 (total annual harvest 1120 kg).

Fig. 6. Geographic distribution of annual red coral amounts in 2009 (until August) (total annual 

harvest 1435 kg).

40%

15%

23%

17%

4% 1%

Alghero

S. Teresa di Gallura

Fertilia

Bosa

S.Antioco

Isola Rossa

23%

18%

15%

2% 5% 2% 35%

Alghero

S. Teresa di Gallura

Fertilia

Bosa

S.Antioco

Castelsardo

Isola Rossa

132
Fig. 7. Distribution of coral harvesting: different coral conditions in Sardinian harvesting areas 

in 2008 (Kg). 

Effects of the Regional Management

Analysis on the main harvesting effort variables related to red coral annual or daily harvested 

amount were carried out from 1996 to 2009 (until August) in order to evaluate the effects of the 

regional management strategies on the resource (Fig.s 8, 9, 10). 

Fig. 8 shows the trend of the red coral annual amount related to that of the number of permits 

(effectively used). The two variables seem to be closely related till 2005. In 2006, the strong 

limitation of the harvesting period (from the 1st of July to the 30th of September), and of the 

allowed daily coral amount per fisherman, took to the extreme decrease of the coral annual harvest. 

Similarly, in 2008, the reduction of the annual coral amount was caused by the drastic reduction of 

permits number and harvesting period (from the 1st of June to the 15th of October). 

Fig. 9 highlights the close relation between the annual coral amount per fisherman and the number of 

annual effective harvesting days per fisherman. The decrease in coral annual amount per fisherman 

is not due to a reduction of coral resource as confirmed by the positive trend of daily coral amount 

reported in Fig. 10. The resource seems to be in good balance with the harvesting effort.

The trend of daily coral harvest related to that of the total harvesting days (Fig. 10) shows how the 

limitation of harvesting period fixed by councillor’s decree from 2006 to 2009 caused a stronger 

harvesting effort per day (corresponding to a bigger daily coral amount and to worst working 

conditions in relation to safety of coral divers).

0

50

100

150

200

250

300

350

400

450

500

Alghero S. Teresa

di Gallura

Fertilia Bosa S.Antioco Isola

Rossa

Harvesting areas

Kg of 

harvested 

coral 

Dead

Mixed

Spoilt (yellow)

Red

133
0

20

40

60

80

100

120

140

160

19961997199819992000200120022003200420052006200720082009

Kg of red coral per fisherman

0

10

20

30

40

50

60

70

80

90

100

N°of days per fisherman

Red coral

annual harvest

per fisherman

Number of

annual

harvesting

days per

fisherman

Fig. 9. Red coral annual harvest per fisherman related to the number of annual harvesting days 

per fisherman.

0

500

1000

1500

2000

2500

3000

3500

4000

19961997199819992000200120022003200420052006200720082009

Kg of red coral 

0

5

10

15

20

25

30

35

40

N° of permits 

red coral

annual

harvest (kg)

Number of

permits

(effectively

used)

Fig. 8. Red coral annual harvest related to the number of permits effectively used.

134
0,0

0,2

0,4

0,6

0,8

1,0

1,2

1,4

1,6

1,8

2,0

19961997199819992000200120022003200420052006200720082009

red coral (kg/day) 

0

400

800

1200

1600

2000

2400

N° of harvesting days 

Daily red coral

amount (Kg)

Total harvesting

days

Discussion and Conclusions

A SWOT analysis was carried out (Table 3). In conclusion, the management strategy adopted in 

Sardinia seems to reveal that a good balance between protection and exploitation requirements is 

achievable also thanks to the strong collaboration among fishermen, high profile research institutions 

and regional administration. Further improvements will take into account new challenges like 

collection of independent data, independent evaluation, promotion of consortia, benchmarking 

with other administrations, and development of international pilot projects.

Fig. 10. Daily red coral amounts related to the annual number of harvesting days (total of 

effective harvesting days of all coral divers).

135
Acknowledgments

We would like to express our gratitude to: Regional Councilor Andrea Prato – responsible for 

Agriculture and Fisheries, Alfonso Orefice – General Director of Agriculture Department, 

Riccardo Zini, Paola Marras and Paolo Cherchi – regional officers of Fishery Service (Agriculture 

Department), Angelo Cau and the researchers of the Department of Animal Biology and Ecology 

(University of Cagliari) and regional coral fishermen for harvesting data provision.

Table 3. SWOT.

STRENGTHS WEAKNESSES OPPORTUNITIES THREATS

Settled management 

system

Lack of training programs 

for coral divers and 

officers are continuous

Strengthen economic 

viability

Climate change (global 

warming)

Restrictions on destructive 

fishing methods

Lack of entrepreneurial 

networking 

Branding strategies 

and distinctive quality 

management programs

Illegal, unreported and 

unregulated (IUU) fishing 

Specific regulation and 

management system based 

on biological features of 

red coral (low growth 

rate, short distance larval 

dispersion, low recruitment 

rate) 

Trade of mixed quality 

coral Research development

(illegal fishing during 

forbidden periods and in 

forbidden areas)

Multiannual harvesting 

data collection (quantities, 

areas, depths)

Poor marketing strategies International cooperation 

and benchmarking

Lack of financial resources 

to guarantee longterm enforcement and 

widespread control on 

fishing activities

Annual regulation based 

on scientific evidence

Active involvement 

of red coral fishermen 

against ghost fishing gears 

retrieval

Multiannual experience 

of bilateral cooperation 

between Regional 

Administration and 

Scientific Research 

Departments

Restocking programs

136
References

Cannas, R., Caocci, F., Follesa, M.C., Grazioli, E., Pedoni, C., Pesci, P., Sacco, F., Cau, A. (in press). 

Multidisciplinary data on the status of red coral (Corallium rubrum) resource in Sardinian 

seas (central western Mediterranean). Proceedings of the International Workshop “Red 

Coral Science, Management, and Trade: Lessons from the Mediterranean” University of 

Naples “Parthenope” - Naples, September 23-26, 2009.

Doneddu, G., Fiori, A. 2003. La pesca in Italia tra età moderna e contemporanea. Produzione, 

mercato, consumo. Editrice Democratica Sarda.

137
DEVELOPMENT OF A GUIDE TO THE 

IDENTIFICATION OF PRECIOUS CORALS

E. Cooper

Director Traffic and Wildlife/WWF

Products such as jewelry and objects d’art made from Coralliidae and other coral taxa (including 

Antipatharia, Gerardia, Heliopora, Isididae, Melithaea, and Scleractinia – collectively referred-to 

here as precious corals) command high prices and near-global market demand. As a result, the 

trade in these corals is extensive and profitable and provides ample incentive for their harvest. 

Unfortunately, most of these taxa have life-history characteristics that make them particularly 

vulnerable to over-exploitation. In addition, the use of non-selective, destructive harvesting 

equipment in some areas, for some taxa, impacts not only the targeted coral species but also the 

invertebrate species dependent on the corals as a substrate.

Management and enforcement of trade regulations and international agreements concerning these 

taxa is complicated by the lack of resources designed to assist with the identification of precious 

corals and their products. Currently a comprehensive guide to the identification of precious corals 

does not exist.

Completion of this project will result in the worldwide distribution of an identification guide to the 

precious corals that will assist fisheries and wildlife trade management, scientific and (especially) 

enforcement authorities to better manage and enforce regulations regarding the harvest and trade 

of these taxa and thereby promote the wise management and sustainable use of coral resources. 

It will also facilitate the implementation of a CITES listing of Corallium, should the genus be so 

listed at the 15th Meeting of the Conference of CITES Parties in 2010.

138
Section III

Science: Biology and Ecology of Red Coral

139
POPULATION GENETIC STRUCTURE OF CORALLIUM RUBRUM: 

IMPLICATIONS FOR MANAGEMENT AND CONSERVATION

Marco Abbiati1

, Federica Costantini1, Cécile Fauvelot2, Eleonora Pintus1

1 Centro Interdipartimentale di Ricerca per le Scienze 

Ambientali and Dipartimento di Biologia Evoluzionistica Sperimentale, 

University of Bologna, Via S. Alberto 163, I – 48100 Ravenna, Italy

2 IRD – UR128, Centre de Biologie et d’Ecologie Tropicale et Méditerranéenne 

Université de Perpignan, 52 Av. Paul Alduy, F - 66860 Perpignan cedex, France.

E-mail: marco.abbiati@unibo.it

Abstract

Understanding patterns of population genetic structuring has been recognized as one of the 

priorities for the conservation of biodiversity, including marine species affected by natural threats 

and human harvesting. The development of refined molecular tools allowed the collection of 

information on population divergence, gene flow, mixing of gene pools, effective population size, 

genetic loss, bottlenecks, effect of natural/human selection etc, providing an invaluable contribution 

for a scientifically sound management and conservation of marine biological resources. Genetic 

data are particularly informative in sessile taxa with patchy distribution, where spatial patterns 

of genetic structuring can be clearly assessed. Moreover, in species where is not possible to track 

spawning and larval movement, genetic data allow inferences about effective larval dispersal 

and connectivity among populations. Molecular tools allow the identification of “Management 

Units”, thus contributing to the implementation of effective conservation strategies. Most studies 

on genetic structure of Corallium species have been done on Mediterranean C. rubrum while only 

preliminary data are available for some Atlantic species (C. lauuense and C. secundum). Genetic 

structure of Corallium species has been investigated using molecular markers with different levels 

of polymorphism (e.g. ITS, microsatellites, mtDNA). Corallium species are characterized by 

significant heterozygote deficiencies and deviations from Hardy–Weinberg expectations. These 

deviations are likely due to inbreeding and Wahlund effect.

Detailed studies on shallow water (15-50 m depth) C. rubrum populations showed the occurrence of 

chaotic genetic structuring at Mediterranean scale. Moreover, at small spatial scales, a surprisingly 

strong pattern of genetic divergence has been found. These results suggest that habitat features 

together with biological processes favor the structuring in “Evolutionarily Significant Units” at 

local scales. Patterns of genetic structuring are of great importance to infer about resilience of red 

coral populations. Harvesting strategies and conservation plans of shallow waters banks has to be 

planned at local scale on single populations. Further studies are needed to confirm occurrence of 

small scale patterns of genetic structuring in commercial banks (depth range 70-150m). However, 

a conservative approach would suggest considering each bank as a “Management Unit” requiring 

individual harvesting plan.

Introduction

In the past decades, human-induced disturbance (e.g. habitat loss and fragmentation, global climate 

change, overexploitation of commercial species, intensive trawling, pollution, and tourism) has 

140
increased in marine coastal areas (Airoldi and Beck 2007). Owing primarily to this disturbance, 

subtidal biogenic reefs in tropical and temperate habitats have suffered severe degradation, 

affecting both species and genetic diversities (Pandolfi et al. 2003; Bellwood et al. 2004; Worm 

et al. 2006). Sustainable exploitation and conservation of marine species are among the priorities 

of modern bio-resource management (Gray 1997). In this context, estimating connectivity among 

populations is essential since dispersal is a key element warranting population resilience following 

disturbance (Palumbi 2003; Bellwood et al. 2004). Population resilience after a disturbance 

event depends on the supply of new recruits. If populations are self-seeded, relying mainly on 

recruits produced within the population, disturbance events can have long term consequences 

due to reduction in larval supply. Moreover, reduction in population size may promote genetic 

drift, reducing the evolutionary potential of the species and increasing the risk of local extinction. 

However, connectivity among populations is a process that depends on dispersal of gametes and 

larvae between patchily distributed populations, and survival of recruits in the new population 

until they breed and reproduce (Pineda et al. 2007). 

Connectivity is also strongly related to the life history and ecological characteristics of the species 

(reproductive strategies, presence or absence of planktonic larvae, migration habits, etc.). Physical 

and behavioral mechanisms of larval retention (e.g. marine water circulation, wind-driven 

circulation) that can affect genetic homogeneity (Leis and Carson-Ewart 2000; Swearer et al. 1999) 

and occurrence of physical barriers to larval dispersal (hydrodynamism, substrate geomorphology, 

etc.) are extremely relevant. Recent studies (Gaylord and Gaines 2000) underlined the need to 

improve knowledge on biotic and abiotic factors influencing larval dispersal in particular for sessile 

and sedentary benthic species, apparently more vulnerable to habitat fragmentation (Palumbi 

2003). 

Up to now, knowledge on reproductive strategies and range of larval dispersal in benthic invertebrates 

are limited due to the difficulties related to direct observation/tracking and experimental field 

studies (Palumbi and Lessios 2005; Moritz and Lavery 1996). Molecular tools, quantifying genetic 

divergence among populations and the effective number of migrants among them, allow an indirect 

estimation of the range of larval dispersal and of connectivity among populations. This information 

is useful to identify the Management Units (MU) and to plan conservation schemes and effective 

management strategies of natural populations. 

For conservation porpoises, MU has to match the “Evolutionary Significant Units” (ESU). ESUs 

(or population) require an independent management, because they may differ in productivity, adult 

mean size, growth rates, and reproduction time. Moreover, characterization and mapping of ESUs 

allow to identify which units could harvested or conserved, if there are adaptive genetic differences 

among populations and if populations react differently to disturbances.

The genus Corallium comprise 19 species (Bayer and Cairns 2003), including the most valuable 

precious corals. Corallium species dwell on rocky bottom at a depth ranging between 10 and 

2000 m, typically aggregating on banks and seamounts, under ledges, and in and around caves. 

They occur in the Mediterranean Sea, Atlantic Ocean, Indian Ocean, Western and Eastern Pacific 

Ocean. Out if 19 species only seven are harvested for jewelry purposes (C. secundum, C. elatius, 

C. konojoi, C. japonicum (now classified as Paracorallium japonicum), C. lauuense, Corallium

s.n., and C. rubrum). C. rubrum is the only Mediterranean species. It is able to colonize caves and 

overhanging at shallow depths (from about 10 m depth), differing from all other Corallium species, 

141
which occur at a depth greater than 500 m (Bayer 1964, 1996; Grigg and Bayer 1976; Bayer and 

Cairns 2003). Recently, live red coral colonies have been observed in the Strait of Sicily down 

to about 800 m (Costantini et al. 2009; Freiwald et al. 2009). These new records suggested the 

occurrence of three typologies of C. rubrum populations: (i) shallow-water populations in a depth 

range between 15 and about 60 m, dwelling on vertical cliffs and in caves; (ii) intermediate-water 

populations, at a depth range of about 60–300 m; (iii) deep-water populations below 300 m depth 

that are poorly known (Costantini et al. 2009). Nowadays, the commercial harvesting is mainly 

focused on intermediate-water populations of C. rubrum, whereas the main bulk of biological 

knowledge refers to the more accessible shallow-water populations.

In this work we summarize the existing knowledge on population genetic structuring in Corallium

species, to identify the recurrent genetic features and to reveal spatial scales of genetic structuring 

and patterns of larval dispersal capability in Mediterranean Corallium rubrum populations. The 

implications of these findings for harvesting management and conservation strategies of precious 

red coral are also addressed. We also discuss gaps in genetic data and how to fill these gaps.

Mitochondrial Genetic Markers in Corallium Species

The most effective molecular markers to investigate intraspecific level of genetic structuring are 

sequence polymorphism of mtDNA and microsatellite loci (Fig. 1). Studies on genetic structure of 

Corallium species are limited, and deals mainly with Mediterranean Corallium rubrum (Table 1).

In Anthozoa, mitochondrial DNA is highly conserved, providing little information on the genetic 

structure of the species and failing to detect genetic heterogeneity among samples because of their 

high degree of conservativeness (Shearer et al. 2002). Studies by Costantini et al. (2003), Calderon 

et al. (2006), and Mokhtar-Jamai et al. (2009) showed that 16S and COI genes in Corallium

rubrum have very low variability, making them unsuitable markers for studies of variation at the 

intraspecific level. This lack of variability in mitochondrial genes seems related to evolutionary 

processes within the mitochondrial genome (Shearer et al. 2002). Nevertheless, Costantini et al. 

(2009) and France and Hoover (2002) have shown one substitution in mitochondrial sequences 

(MSH and COI) between shallow- and deep- water specimens of Corallium rubrum and Corallium 

ducale respectively. Such a fixed substitution of one nucleotide might suggest lack of connectivity 

between shallow- and deep-water populations in both Corallium species, but the small sample size 

of the deep-water collections does not allow further consideration on the evolutionary implications 

of this genetic variation (Costantini et al. 2009).

Other species like C. kishinouyei, C. ducale, C. regale and C. niobe have been used in phylogenetic 

studies at greater taxonomic level (France et al. 1996; France and Hoover 2001; France and Hoover 

2002; Sanchez et al. 2003; Mc.Fadden et al. 2006; France 2007) but no detailed phylogenetic 

studies within the genus Corallium have been done up to now (e.g. Sanchez 2010, in prep).

Microsatellite Genetic Variability in Corallium Species

Population genetic studies have been done on three of the 19 Corallium species: Corallium lauuense, 

C. secundum, and C. rubrum. The first two species are deep-water corals (depth range 385-535 

m) abundant and widely distributed in the Hawaii. Only few populations of these species have 

142
been analyzed using microsatellite loci (Table 2), which are the appropriate molecular markers to 

examine the occurrence of genetic structuring at different spatial scales.

Microsatellite studies of Corallium lauuense indicated significant heterozygote deficiency in most 

populations, suggesting recruitment in most populations is from local sources with only occasional 

long-distance dispersal events. Also, two populations appear to be significantly isolated from other 

populations of C. lauuense and may be separate stocks (Baco and Shank 2005; Baco et al. 2006). In 

contrast, C. secundum populations have little heterozygote deficiency and separate into 3 distinct 

regions. In addition to having fisheries management implications for these corals, the results of 

these studies also have implications for the management and protection of seamount fauna (Baco 

2006).

Population Genetic Structure of Corallium rubrum

Genetic structuring in this species was first described using allozyme electrophoresis (Abbiati et al. 

1993). These studies showed the occurrence of genetic differentiation among red coral populations 

at a spatial scale of a few kilometers and a lack of genetic structuring over shorter distances (about 

200 m) (Abbiati et al. 1997). However, allozyme markers may have limited power in revealing 

small-scale genetic structure due to their low polymorphism and mutation rates.

Afterwards, a combination of sequence polymorphism of the Internal Transcribed Spacer (ITS) 

and species – specific microsatellite loci (Costantini and Abbiati 2006) have been used to analyze 

the genetic structuring of red coral populations at Mediterranean scale (Costantini et al. 2007a). 

Both markers revealed a high degree of genetic differentiation between samples throughout the 

northwestern Mediterranean Sea (though to a different extent). Moreover, microsatellites and ITS-1 

sequences showed no correlation between genetic differentiation and geographical distance among 

the analyzed samples. Such lack of correlation may support an island model of dispersal, where 

larvae move from one area to another with equal probability, rather than an isolation-by-distance 

model, where the probability of dispersal between sites declines with increasing geographical 

distance. The absence of clear patterns should be interpreted with caution because such patterns 

may hide underlying small-scale genetic structuring (Costantini et al. 2007a).

In 2007, Costantini et al. (2007b), using four microsatellite loci have analyzed small spatial scale 

genetic structuring in C. rubrum samples collected in two location (hundred Kms distance) using 

a hierarchical sampling design including scales from 10s to 100s of meters. Significant deviations 

from Hardy-Weinberg equilibrium due to elevated heterozygote deficiencies were detected in all 

samples and were consistent with the occurrence of inbreeding (mating between consanguineous) 

and a Wahlund effect (mixing of differentiated gene pools). These results have been confirmed also 

to recent data by Mokhtar-Jamai et al (2009) and Ledoux et al. (in prep). The two locations showed 

differences in genetic variability that could be related to differences in habitat characteristics. The 

size and spatial arrangement of habitat, indeed, may be of great importance for the ecology and 

population structure of species with limited dispersal range (Palumbi 2004). Significant levels of 

genetic differentiation were found at spatial scales of 10s of meters, supporting the hypothesis 

that planulae have limited effective dispersal ability. Furthermore, evidence of inbreeding, the 

Wahlund effect and high genetic variance within samples suggest that the effective larval dispersal 

range may be < 10 m (Costantini et al. 2007b). A more recent study, analyzing collecting samples 

143
at 1 meter intervals along a 10 m transect have observed a chaotic genetic structuring also at this 

smallest spatial scale (Costantini et al., in prep).

At least, the find out of deep live red coral colonies have allowed a preliminary genetic 

characterization of these specimens implying the possible occurrence of genetic isolation among 

shallow and deep-water populations. These findings suggest that genetic diversity of red coral 

over its actual range of depth distribution is shaped by complex interactions among geological, 

historical, biological and ecological processes (Costantini et al. 2009).

Conclusion

Molecular tools have been used to study genetic variability and genetic structuring of 7 Corallium

species. Conservation issues have been addressed for 3 species: C. lauuense, C. secundum and 

C. rubrum. No studies on phylogenetic relationships between Corallium species have been 

performed (but Sanchez 2010 in prep, pers. comm.) so up to now there are no clear evidences 

about evolutionary relationship within the genus Corallium.

Common Genetic Features in Corallium Species

All the Corallium species analyzed by means of molecular markers have shown a deficit of 

heterozygotes. This deficit has been explained by the occurrence of inbreeding within samples 

and of local recruitment. These results are in accordance with the reproductive biology and larval 

ecology of Corallium. For example, in red coral, embryo development takes place within the 

female polyps for about 30 d, before release of planula larvae (Vighi 1972). Moreover, reduced 

swimming ability and geonegative behavior of the planulae suggest that, once released, larvae 

settle in close vicinity of the parental colonies (Vighi 1972). Therefore, reproduction events in 

red coral are more likely to involve related individuals compared to species with broadcasting 

development and so Corallium populations are primarily self-recruiting. 

Spatial Scale of Genetic Structuring in Corallium rubrum

Most studies on scales of genetic structuring have been done on Mediterranean Corallium rubrum 

(Fig. 2). Using molecular markers with differing levels of polymorphisms, a complex genetic 

structuring at different spatial scale has been observed. In fact, a strong genetic structuring at 

Mediterranean scale with no isolation by distance patterns has been observed. Moreover, a pattern 

of chaotic genetic divergence at small spatial scales has also been detected. If at Mediterranean 

scale the genetic structuring observed could be strongly influenced by the environmental features 

where the species live (e.g. habitat characteristics, geomorphology, and hydrodynamics), at the 

smallest spatial scale biological processes (e.g. pre- and post-settlement mortality and stochasticity 

of reproductive success) may favor the occurrence of genetically distinct pools of recruits at 

meter distances. The assumption, made in most population dynamic studies (Abbiati et al. 1993; 

Santangelo et al. 2009), that red coral population are closed populations is supported by genetic 

results (Costantini et al., in prep). Moreover, habitat features together with biological processes 

favor the structuring in “Evolutionarily Significant Units” at local scales.

144
Gaps to Fill

This survey of the scientific literature on the genetic features of Corallium species shows that a 

series of gaps have to be filled, to allow suggesting effective guidelines for harvesting strategies 

and conservation of these species.

1) To understand patterns of genetic structuring and reproductive behaviour of Corallium

species population genetic analysis have to be extended to all commercial species which have not 

been yet investigated, e.g. Japanese red coral (Paracorallium japonicum), pink coral (Corallium 

elatius) and white coral (Corallium konojoi). 

2) New molecular markers with intermediate variability compared to nuclear DNA and 

mitochondrial genes have to be developed to properly address pattern of genetic structuring 

between populations at regional scale (e.g. the Mediterranean Sea for C. rubrum). For C. rubrum, 

it would be essential to extend sampling to the Atlantic, Southern, and Eastern Mediterranean. The 

scarcity of sampling efforts along the Southern and Eastern Mediterranean and Atlantic coasts 

suggest the need to develop effective inter-regional collaborations plans.

3) Studies have to include the whole depth range distribution as well as the geographic 

distribution of the species. While there are now good knowledge about population parameters 

and connectivity patterns in shallow-water populations, we just begin to study the most exploited 

intermediate-water and the newly recorded deep-water populations. Moreover, temporal patterns 

of genetic structuring should be investigated to check if a reduction of genetic variability occurred 

between ancient and actual populations.

4) Integrating data on different aspects of the ecology, biology, genetics of Corallium

species should be a priority for management agencies. In particular, future researches have to focus 

on integration between studies on population structure, population growth rates (e.g. reproduction, 

recruitment), and connectivity patterns.

Implication for Management

Available genetic data on Corallium species, even if fragmentary and mainly derived form 

shallow water Mediterranean populations (Abbiati et al. 1993; Costantini et al. 2007a,b) provide 

useful information to infer about size of the ‘Management Units,’ recovery ability of the species, 

harvesting strategies, and conservation needs of Corallium populations.

The chaotic genetic structuring observed at different spatial scale ranging from Mediterranean 

to micro suggest that conservation of shallow waters red coral populations has to be planned at 

regional or local scales. Based on the precautionary principle, also for commercially harvested 

intermediate-water populations (70-150 m depth) individual harvesting plans of each bank has 

to be considered. Identification of the commercial banks within the Mediterranean Sea and 

quantification of the available commercial stocks of the species has to be a priority. Each stock 

should be characterized from the molecular point of view to identify population boundaries and 

Management Units. Management strategies at pan-Mediterranean scale should be implemented 

based on Regional Management Units, by defining harvesting regions and applying rotating 

harvesting schemes within each region.

145
References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Marine Ecology Progress 

Series 95: 245-250.

Abbiati, M., Novelli, S., Harmelin, J.G., Santangelo, G. 1997. Struttura genetica di popolamenti 

simpatrici e allopatici di corallo rosso. In: Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. 

(Eds.) Biologia e tutela del corallo rosso e di altri ottocoralli del Mediterraneo. Ministero 

delle politiche agricole, Rome, 5-21.

Airoldi, L., Beck, M.W. 2007. Loss, status and trends for coastal marine habitats of Europe. 

Oceanography and Marine Biology: an Annual Review 45: 345-405.

Baco, A.R., Shank, T.M. 2005. Population genetic structure of the Hawaiian precious coral Corallium 

lauuense (Octocorallia: Coralliidae) using microsatellites. In: Cold-Water Corals and 

Ecosystems, Freiwald, A., Roberts, J.M.). Springer Publishing House, Heidelberg, Germany.

Baco, A.R. 2006. Population Genetic Structure of the Deep-Sea Precious Coral Corallium secundum 

from the Hawaiian Archipelago Based on Microsatellites, Eos Trans. AGU, 87(52), Fall 

Meet. Suppl., Abstract V13A-0649.

Baco, A.R., Clark, A.M., Shank, T.M. 2006. Six microsatellite loci from the deep-sea coral Corallium 

lauuense (Octocorallia: Coralliidae) from the islands and seamounts of the Hawaiian 

archipelago. Molecular Ecology Note 6: 147-149. 

Bayer, F.M., Cairns, S.D. 2003. A new genus of the scleraxonian Family Coralliidae (Octocorallia: 

Gorgonacea). Proceedings of the Biological Society of Washington 116: 222-228.

Bayer, F.M. 1964. The genus Corallium (Gorgonacea: Scleraxonia) in the Western North Atlantic 

Ocean. Bulletin of Marine Science Gulf and Caribbean 14: 465-478.

Bayer, F.M. 1996. Three new species of precious coral (Anthozoa: Gorgonacea, genus Corallium) 

from Pacific waters. Proceedings of the Biological Society of Washington 109: 205-228.

Bellwood, D.R., Hughes, T.P., Folke, C., Nystroem, M. 2004. Confronting the coral reef crisis. Nature 

429: 827-833.

Berntson, E.A., Bayer, F.M., McArthur, A.G., France, S.C. 2001. Phylogenetic relationships within 

the Octocorallia (Cnidaria:Anthozoa) based on nuclear 18S rRNA sequences. Marine 

Biology 138: 235-246.

Brugler, M.R., France, S.C. 2008. The mitochondrial genome of a deep-sea bamboo coral (Cnidaria, 

Anthozoa, Octocorallia, Isididae): genome structure and putative origins of replication are 

not conserved among octocorals. Journal of Molecular Evolution 67: 125-136.

Calderon, I., Garrabou, J., Aurelle, D. 2006. Evaluation of the utility of COI and ITS markers as tools 

for population genetic studies of temperate gorgonians. Journal of Experimental Marine 

Biology and Ecology 336: 184-197.

Costantini, F., Abbiati, M. 2006. Development of micro satellite markers for the Mediterranean 

gorgonian coral Corallium rubrum. Molecular Ecology Notes 6: 521-523.

Costantini, F., Tinti, F., Abbiati, M. 2003. Sistematica molecolare e filogenesi di Corallium rubrum. 

Biologia Marina Mediterranea 1: 73-75.

146
Costantini, F., Fauvelot, C., Abbiati, M. 2007a. Genetic structuring of the temperate gorgonian coral 

(Corallium rubrum) across the Western Mediterranean Sea revealed by microsatellites and 

nuclear sequences. Molecular Ecology 16: 5168-5182.

Costantini, F., Fauvelot, C., Abbiati, M. 2007b. Fine-scale genetic structuring in Corallium rubrum

(L.): evidences of inbreeding and limited effective larval dispersal. Marine Ecology Progress 

Series 340: 109-119.

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembri, P.J., Abbiati, M. 2009. Deepwater Corallium rubrum (L. 1758) from the Mediterranean Sea: preliminary genetic 

characterisation. Marine Ecology: an evolutionary perspective, DOI: 10.1111/j.1439-

0485.2009.00333.x

France, S.C., Hoover, L.L. 2002. DNA sequences of the mitochondrial COI gene have low levels of 

divergence among deep-sea octocorals (Cnidaria: Anthozoa). Hydrobiologia 471: 149-155.

France, S.C., Hoover, L.L. 2001. Analysis of variation in mitochondrial DNA sequences (ND3, 

ND4L, MSH) among Octocorallia (Alcyonaria) (Cnidaria: Anthozoa). Bulletin of the 

Biological Society of Washington 10: 110-118.

France, S.C. 2007. Genetic analysis of bamboo corals (Cnidaria: Octocorallia: Isididae) Does lack 

of colony branching distinguish Lepidisis from Keratoisis? Bulletin of Marine Science 81: 

323-333.

France, S.C., Rosel, P.E., Agenbroad, J.E. 1996. DNA sequence variation of mitochondrial largesubunit rRNA provides support for a two-subclass organization of the Anthozoa (Cnidaria) 

Molecular Marine Biology and Biotechnology 5: 15-28. 

Freiwald, A., Beuck, L., Ruggeberg, A., Taviani, M., Hebbeln, D. and R ⁄V Meteor M70-1 Participants 

(2009) The white coral community in the Central Mediterranean Sea revealed by ROV 

surveys. Oceanography 22: 58-74.

Gaylord, B., Gaines, S.D. 2000. Temperature or transport? Range limits in marine species mediated 

solely by flow. American Naturalist 155: 769-789.

Gray, J. 1997. Marine biodiversity: patterns, threats and conservation needs. Biodiversity and 

Conservation 6: 153-175.

Grigg, R.W., Bayer, F.M. 1976. Present knowledge of the systematic and zoogeography of the order 

Gorgonacea in Hawaii. Pacific Science 30: 167-175.

Leis, J.M., Carson-Ewart, B.M. 2000. Behaviour of pelagic larvae of coral reef species in the ocean 

and an atoll lagoon. Coral Reefs 19: 247-257.

McFadden, C.S., France, S.C., Sanchez, J.A. 2006. A molecular phylogenetic analysis of the 

Octocorallia (Cnidaria : Anthozoa) based on mitochondrial protein-coding sequences. 

Molecular Phylogenetics and Evolution 41: 513-527. 

Mokhtar-Jamai, K., Ledoux, J.B., Garrabou, J., Feral, J.P., Aurelle, D. 2009. Interest and application 

of genetic markers for the study and conservation of Mediterranean sessile invertebrates. In: 

Proceeding of the 1st Mediterranean symposium on the conservation of the coralligenous 

and other calcareous bio-concretions, 15-16 Jan 2009, Tabarka, Tunisie (C. Pergent-Martini, 

M. Brichet, Eds.) RAC/SPA: Tunis, 112-116.

147
Moritz, C., Lavery, S. 1996. Molecular ecology: contributions from molecular genetics to population 

ecology. In: Floyd, R.B. Frontiers of Population Ecology, CSIRO Publishing. 1996. 433-

450.

Palumbi, S.R., Lessios, H.A. 2005. Evolutionary animation: how do molecular phylogenies compare 

to Mayr’s reconstruction of speciation patterns in the sea? Proceedings of the National 

Academy of Sciences of the United States of America 102: 6566-6572.

Palumbi, S. 2003. Population genetics, demographic connectivity, and the design of marine reserves. 

Ecological Applications 13: S146-S158.

Palumbi, S.R. 2004. Marine reserves and ocean neighbourhoods: The spatial scale of marine 

populations and their management. Annual Review of Environment and Resources 29: 31-

68.

Pandolfi, J.M., Bradbury, R.H., Sala, E. 2003. Global trajectories of the long-term decline of coral 

reef ecosystems. Science 301: 955-958.

Pineda, J., Hare, J.A., Sponaugle, S. 2007. Larval transport and dispersal in the coastal ocean and 

consequences for population connectivity. Oceanography 20: 22-39.

Sanchez, J.A., McFadden, C.S., France, S.C. 2003. Molecular phylogenetic analyses of shallowwater Caribbean octocorals. Marine Biology 142: 975-987.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an overexploited 

Mediterranean species. Aquatic Conservation: Marine and Freshwater Ecosystems 11: 253-

259.

Santangelo, G., Bongiorni, L., Giannini, F., Abbiati, M., Buffoni, G. 1999. Structure analysis of 

two red coral populations dwelling in different habitats. In: Cicogna, F., Bavestrello, G., 

Cattaneo-Vietti, R. (Eds.) Red Coral and Other Mediterranean Octocorals: Biology and 

Protection, Ministero delle Politiche Agricole Alimentari e Forestali, Rome: 23-43.

Slatkin, M. 1987. Gene flow and the geographic structure of natural populations. Science 236: 787-

792.

Shearer, T.L., van Oppen, M.J.H., Romano, S.L., Worheide, G. 2002. Slow mitochondrial DNA 

sequence evolution in the Anthozoa (Cnidaria). Molecular Ecology 11: 2475-2487. 

Swearer, S.E., Caselle, J.E., Lea, D.W., Warner, R.R. 1999. Larval retention and recruitment in an 

island population of a coral-reef fish. Nature 402: 799-802.

Tsounis, G., Rossi, S., Aranguren, M., Gili, J.M., Arntz, W. 2006. Effects of spatial variability and 

colony size on the reproductive output and gonadal development cycle of the Mediterranean 

red coral (Corallium rubrum L.). Marine Biology 148: 513-527.

Worm, B., Barbier, E.B, Beaumont, N. 2006. Impacts of biodiversity loss on ocean ecosystem 

services. Science 314: 787-790.

148
Fig. 1. Genotyping of three Corallium rubrum colonies using two microsatellite loci (Cor48 and 

Cor15).

Fig. 2. Map of the Corallium rubrum populations analyzed with molecular markers (after Costantini 

et al. 2007a, modified).

149
Table 1. Existing references of Corallium species analyzed with different molecular markers.

Species Molecular marker References

Corallium 

rubrum allozyme Abbiati et al. (1993, 1997)

Corallium 

rubrum mtDNA Costantini et al. (2003); Calderon et al. (2006), MokhtarJamai et al. (2009)

Corallium 

rubrum ITS Costantini et al. (2007a)

Corallium 

rubrum microsatellites Costantini and Abbiati (2006); Costantini et al. (2007a, b); Mokhtar-Jamai et al. (2009)

Corallium 

lauuense microsatellites Baco and Shank (2005); Baco et al. (2006)

Corallium 

secundum microsatellites Baco (2006)

Corallium 

kishinouyei mtDNA France et al. (1996); France and Hoover (2001, 2002); Berntson et al (2001)

Corallium 

ducale mtDNA France et al. (1996); France and Hoover (2001, 2002); France (2007); Brugler and France (2008) 

Corallium 

regale mtDNA France and Hoover (2001, 2002)

Corallium 

niobe mtDNA France (2007)

Table 2. Non-Mediterranean populations of Corallium analyzed using microsatellite loci.

 

Species Studied 

populations Distribution

Corallium lauuense 8 Hawaiian archipelago

Corallium secundum 11 Hawaiian archipelago

150
BIOLOGICAL INTERACTIONS AFFECTING THE GROWTH RATES

OF RED CORAL (CORALLIUM RUBRUM) COLONIES*

Giorgio Bavestrello1, Carlo Cerrano2, Riccardo Cattaneo-Vietti2

1 DiSMar, Università Politecnica delle Marche, Via Brecce Bianche, 60131 Ancona

2 DipTeRis, Università di Genova, Corso Europa 26, 16132 Genova

E-mail: g.bavestrello@univpm.it

Abstract

The red coral (Corallium rubrum: Cnidaria, Anthozoa) is a significant Mediterranean resource 

with wide historical, economic, and cultural involvements. Several countries in the Mediterranean 

Sea have adopted different kinds of strategies for its conservation. On the other hand, the main 

data necessary for a rational management of a biological resource is the time necessary for 

the recruits to reach an exploitable size. Data about the growth rate of just settled or juvenile 

colonies living in shallow waters (< 50 m) are abundant and values of 1 mm year-1 for the base 

diameter and 1 cm year-1 for the height of the colony are generally accepted. Nevertheless it is 

demonstrated that 4-5 years after the settling, the growth virtually stops or becomes negligible: 

colonies monitored for more than 20 years along the French coast reached an height of about 

4 cm and a basal diameter of 0.5 cm. Historical and recent data about the size and density of 

the shallow water population of Portofino Promontory (Ligurian Sea) are available, indicating 

important fluctuations of these parameters. Firstly, samples collected during the ‘50 years, together 

with contemporaneous popular articles and interviews with old divers, strongly suggest that the 

size structure of the Portofino population is not significantly changed in the last 50 years, although 

a slight exploitation of the banks occurred. Actually, the colony size of the colonies is inversely 

related to their density, suggesting an intraspecific competition linked to the space availability 

and trophic inputs. Moreover, it is surprising that in the last 10 years the Portofino shallow water 

populations have shown an unusually fast growth rate, reaching the maximal colony size never 

recorded in the previous 40 years (15 cm in height and 20 DW g). This unusual fast growth rate 

was recorded after the mass mortality episodes of 1999 and 2003, involving not only red coral, 

but also several other benthic species and provoking a significant change in the structure of the 

coralligenous assemblage. All these evidences suggest that the growth of the Mediterranean red 

coral is strongly affected by intra- and inter-specific interactions linked not only to local variables 

by also to stochastic events.

Introduction

The red coral (Corallium rubrum: Cnidaria, Anthozoa) is an important Mediterranean resource 

with huge historical, economic, and cultural involvements (Cicogna and Cattaneo-Vietti 1993; 

Cicogna et al. 1999). It is widely distributed throughout the Mediterranean: occurring both in 

commercially exploitable concentrations, as off Spain, Algeria, and Sardinia, as well as at lower 

densities from the Aegean and southern and eastern Mediterranean to the Sea of Alboran. Some 

populations were recorded in the Capo Verde Archipelago (Zibrowius et al. 1984).

Preoccupation has been expressed by fishermen and industry with the declining returns to an ever 

more sophisticated harvesting gears capable of operating at more than one hundred metres depth, 

151
which have progressively substituted the primitive dragging equipment. Scuba harvesting tends to 

be in a “pulse” mode, in which local patches are cleaned out selectively, leaving only inaccessible 

or non commercial individuals, before another patch is located and harvested in turn (Caddy 1993). 

Consequently, several countries in the Mediterranean Sea have adopted different kinds of strategies 

for the red coral conservation, particularly the closure of coral fishery in overexploited areas.

Red coral population density and colony size vary from place to place, also according to depth 

and exploitation (Tsounis et al. 2006; Rossi et al. 2008). Schematically, we can distinguish two 

different spatial situations: (i) coastal populations, occurring up to 50 m depth, characterised by 

high density (up to 1000 colony/m2

) and small colony size (until 5 cm height), most of colonies 

produce tens of planulae per year; (ii) deeper populations, extending up to 200 m depth and more, 

characterised by low density and high colony size. These colonies may produce until 2000 planulae 

per year (Santangelo et al. 2003). Of these two kinds of situations, the second one only is really 

important from an economic viewpoint while the first one, merely of naturalistic value, constitute 

a stock of breeders for the species.

After settlement, the growth rate of shallow water colonies is about 1 mm year-1 for the base 

diameter, and 10 mm year-1 for the height (Cattaneo-Vietti and Bavestrello 1994) but after 4-5 

years, the growth virtually stops or becomes negligible. In fact, long-term studies (22 years) 

indicate an average growth rates extremely low: 0.24 ± 0.05 mm year-1 for the basal diameter and 

1.78 ± 0.7 mm year-1 for colony height (Garrabou and Harmelin 2002). Similar data were indirectly 

obtained by examining the growth rings of shallow water colonies: specimens with basal diameter 

of about 7 mm were at least 30-40 years old with a growth rate of the basal diameter of 0.35±0.15 

mm year-1 (Marschal et al., 2004). Bramanti et al. (2005) measured the growth rate of colonies 

settled on long-term artificial substrates: after four years the colonies exhibited an average annual 

growth of the stem diameter of 0.62 ± 0.19 mm year-1. Similar data were also obtained by Torrents 

et al. (2008). Practically no information is available for deeper populations, but if these trends are 

applied to all cases, we may estimate age of several decades for colonies of 10-20 mm of basal 

diameter and 20-30 cm of height. 

Although very few data are available, coral growth rate is probably affected by physical and 

chemical variables (light, currents, input of organic matter, Ca++ availability) and biological ones 

(food availability, competition). Practically no data are available on intra- and inter-specific 

competitions and larval behavior which could have a significant role in structuring the deeper 

populations. Also the thermal anomalies can affect the size and shape of the coral colonies: during 

these episodes, autotomy of the branch tips was noticed on several colonies (Russo et al. 1999) and 

the reproductive implications of this phenomenon still need to be clarified. 

Finally anthropogenic impact can be very important: an underestimated phenomenon is the impact 

on coral populations of fishing lines that broke the branches of the colonies. 

To give a contribution to the knowledge of the Mediterranean structure and dynamics of the coastal 

populations, in this paper we discuss on size and density of the red coral along the Portofino 

Promontory (Ligurian Sea), comparing present and historical data.

152
Results

Size/Density Relationship Along a Depth Transect

We have conducted a series of measures on the density and size of the red coral colonies along a 

depth transect on the rocky cliff of Portofino Promontory (Ligurian Sea). This study was conduced 

during July 2008, by using non-destructive photographic methods. Data were recorded in three 

stations, at 30, 50, and 70 m depth by scuba diving. 

The observations indicate that the two variables 

are inversely related, as shown in Fig. 1. Density 

is very high at 30 m depth (about 300 colonies 

m-2), but the average height of the colonies is low 

(about 5 cm). At this depth, the colonies mainly 

growth hanged to the roof of the crevices of the 

coralligenous conglomerate. The biocoenosis 

includes several sponge species insinuating, 

encrusting and massive or erected. The stony 

coral Leptopsammia pruvoti is often present. 

Among bryozoans, Smittina, Rynchozoon, and 

Reteporella are among the commonest genera.

At 50 m depth, the size of the colonies drastically increases but, contemporaneously, the density 

is reduced. Here red coral settles perpendicularly to the cliff together with several sponges and 

bryozoans. L. pruvoti is very abundant.

At 70 m depth, the colonies show a commercial size: 1-2 cm in the basal diameter and about 15 

cm in height, but their densities drop down to only 2 colonies m-2. Recruits and juvenile colonies 

are very rare or completely lacking. At this depth, the colonies live on sparse rocks arising from 

the detritical bottom. The sedimentation is very high and a sheet of thin sediment covers the rocks. 

Leptopsammia and bryozoans are rare, while only erected sponges are common. Haliclona mucosa

that in shallow waters produces thin crusts inside crevices, here has a thick ear-like shape. 

Growth Rate Variation 

Several information regarding the community structure of the Portofino red coral in the 50’ years 

of the last century suggest it was not so different from today. A group of colonies collected during 

the 50’ years and now conserved at the Museo di Storia Naturale of Genoa has an average height of 

20 cm. In a popular book about describing a long series of diving along the Portofino Promontory, 

Roghi (1966) stated that in this area the coral was always been very abundant but of small size. 

All the interviews with old divers that collected coral in this area agree that on the cliff, until 50 

m depth, the colonies were abundant but very small; from 50 to 60 m depth, on rocks sparse in 

the detritical bottoms some branches 15 cm height with a weight of about 100 g were present. An 

unusual situation was observed on the tip of the promontory (Punta del Faro) were, 80 m depth, 

colonies until 250 g of weight were collected.

0

5

10

15

20

25

0 100 200 300 400

density (N m-2)

colony height (mm)

30 m

50 m

70 m

Fig. 1 – Density/size relationships along a depth 

transect on the Portofino Promontory

Fig. 1. Density/size relationships 

along a depth transect on the Portofino 

Promontory.

153
The first quantitative information about the 

colony size of the Portofino coral are due to 

Marchetti (1965) who recorded, at 30 m depth, 

a DW biomass m-2 lower than 400 g. This datum 

was determined by a low number of colonies 

(< 100) with an average weight of about 3.5 g. 

About 30 years after, Cattaneo-Vietti et al. (1993) 

came back in the Marchetti stations, recording 

an increasing in biomass, reaching about 800 

DW g m-2. The colony density was increased of 

more than three times, but the average weight of a 

single colony decreased until 2.5 g.

During 1999 and 2003 and, with less intensity in 

2005 and 2006, the populations of red coral of the 

Ligurian Sea were strongly affected by episodes 

of mass mortality, sometimes killing more than 

90% of the colonies. Together with red coral also 

several sea fans, sponges and bryozoans (almost 

23 species in 1999 and 14 species in 2003) were 

affected drastically, changing the structure of the 

whole biocoenosis (Cerrano et al. 2000, 2008; 

Perez et al. 2000; Garrabou et al. 2001). 

Recent surveys, conduced in the summer 2008, 

on the same stations, showed a further increasing 

of the biomass, until 1000 DW g m-2. This value 

is due to an increasing of the average size of the 

colonies that sometimes reach 15 cm in height 

and 20 g in DW (Fig. 3). Contemporaneously, the 

density of population again decreases (Fig. 2).

Discussion

Among benthic cnidarians, the Mediterranean red coral shows a peculiar life strategy, not 

completely clarified: its population density is inversely related to colonies size, along both spatial 

and temporal trends. These structural characteristics, observed studying the red coral populations 

living along the Portofino cliffs (Ligurian Sea), agree with ROV studies conducted on the Spanish 

coasts (Rossi et al. 2008). 

In shallow populations, density and size of the Mediterranean red coral can be affected by 

harvesting: recently Tsounis et al. (2006) have demonstrated that the fishing in Costa Brava has 

deeply influenced the population structures, transforming them from a “forest-like” structure to a 

“grass plain-like” one. This has been evidenced comparing harvested populations with “natural” 

ones, living in the Medas Islands MPA.

0

200

400

600

800

1000

1200

1400

1965 1993 2008

total weight/m2 (g)

0

100

200

300

400

500

1965 1993 2008

Colonies/m2 0

1

2

3

4

5

1965 1993 2008

average weight of colonies (g)

Fig. 2 – Historical and present data of biomass 

(A); density (B) and average colony weight (C) of 

the coral population of the Portofino Promontory 

Fig. 2. Historical and present data of biomass (A); density (B) and average colony 

weight (C) of the coral population of the 

Portofino Promontory.

154
The harvesting not only affects the size structure of a shallow population for the obvious reason 

that fishermen collect preferably colonies of large size, but also enhancing the larval settling. 

In 1990, we have experimentally scraped all the colonies present in a 30 x 30 cm surface inside a 

dense Portofino Promontory population. All the colonies were counted and measured. Four years 

after, all the colonies present in the same surface were again 

collected: their number was increased, but the average size 

decreased (Fig. 4). This result suggested a strong intraspecific 

competition as the available space induced a high larval 

settling (Cerrano et al., 1999). Under this condition, the coral 

populations appear like a “copse wood” where, after the cut, 

the new young trees grow densely and quickly, forming a 

bushy structure that remains in this situation for a long span 

of time due to intraspecific competition. Tsounis et al. (2006) 

stated that 14 years of protection at the Medas Island were 

not sufficient to reach a “forest-like” structure.

At the same, the Portofino coral populations were heavily 

exploited during the ‘50-60 years, but in the last 30 years no 

more fishing activities have been conduced. During this long 

span of time, these shallow water populations remained in an 

immature stage, confirming the idea that the “young forest 

effect” could stop for long time the further size increase of 

the population.

An open question is why the recruitment is so high only in shallow waters, while in the deeper 

ones no juveniles were observed although it is well known that the large deeper colonies are able 

to release thousands of planulae (Santangelo et al. 2003). 

It is possible that the high sedimentation recorded 70 m depth, on the rocks emerging from the soft 

bottom, could prevent the recruits as avoids the settling of other benthic organisms, like encrusting 

sponges and bryozoans. It is possible to hypothesise that the planulae released by deep colonies, 

in the absence of a suitable substratum, move to shallow habitats. By this way, the superficial 

populations are continuously supplied by the larvae produced by themselves and by those released 

by deeper colonies, while in deep waters the larval settling remains rare, linked to the stochastic 

presence of favourable micro-environmental situations. Following this hypothesis, the high density 

and small average colony size of the superficial populations could be a natural phenomenon, locally 

enhanced by harvesting and due to the high recruitment typical of these habitats.

On the other hand, free from competition, the deep colonies are able to grow more quickly. 

Moreover, living in more constantly colder water, their final size could be higher as recorded for 

the greatest part of ectothermic organisms (Atkinson and Sibly 1997).

As indicated by our data, the situation of shallow water population can be changed by stochastic 

event like the mass mortality episodes occurred in the Ligurian Sea during 1999 (Cerrano et al. 

2000) and 2003 and, with low intensity, in 2005 and 2006 (Cerrano et al. 2008). Since the first mass 

mortality episode, the Portofino populations have shown an unusually fast growth rate, reaching 

Fig. 3 – A large colony of 30 g 

collected on the Portofino cliff.

Fig. 3. A large colony of 30 g 

collected on the Portofino cliff. 

155
maximal colony sizes (until 15 cm in height and 

1 cm of basal diameter, 20 g DW) values never 

recorded in the previous 43 years.

The obvious explication of this phenomenon 

is that for the few, survived colonies, the 

intraspecific competition decreased and the 

greater availability of several resources gave 

them the opportunity of a faster growth rate. It 

is intriguing that, although the average size of 

colonies is now the largest of the past 43 years, 

the density is greater that in 1965. This fact 

suggests that intraspecific competition is not 

the only constrain able to affect coral growth: 

very likely also inter-specific competition plays 

a significant role. This is particularly evident in the relationships between red coral and the stony 

coral Leptopsammia pruvoti: on the cliff of the Portofino Promontory although the two species 

are able to coexist, they tend to colonise alternative areas, along a same horizontal transect, due to 

imperceptible micro-environmental conditions, by favouring alternately one species or the other 

(Cattaneo-Vietti et al. 1993).

All these evidences suggest that the growth of the Mediterranean red coral is strongly affected by 

intra- and inter-specific interactions linked not only to local variables by also to stochastic events.

0

1200

1990 1994

density (col. m-2)

0

100

1990 1994

height (mm)

0

10

1990 1994

basal diameter (mm)

Fig. 4 – Comparison of density and size of the 

original and the reconstituted population after 4 

years 

Fig. 4. Comparison of density and size of the 

original and the reconstituted population after 

4 years.

156
References

Atkinson, D., Sibly, R.M. 1997. Why are organisms usually bigger in colder environments? Making 

sense of a life history puzzle. Trends in Ecology and Evolution 12: 235-239.

Bramanti, L., Magagnini, G., De Maio, L., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean red coral Corallium rubrum (L. 1758), a 4-year study. Journal 

of experimental Marine Biology and Ecology 314: 69-78.

Caddy, J.F. 1993. Considerazioni sulla strategia di gestione a rotazione della risorsa corallo rosso. In: 

Cicogna, F., Cattaneo-Vietti, R. (Eds.) Red coral in the Mediterranean Sea: Art, History and 

Science. Ministero delle Risorse Agricole, Alimentari e Forestali, Roma: 201-227. 

Cattaneo-Vietti, R., Bavestrello, G., Senes, L. 1993. Il popolamento a corallo rosso del Promontorio 

di Portofino (Mar Ligure). In: Cicogna, F., Cattaneo-Vietti, R. (Eds.) Red coral in the 

Mediterranean Sea: Art, History and Science. Ministero delle Risorse Agricole, Alimentari 

e Forestali, Roma: 181-200.

Cattaneo-Vietti, R., Bavestrello, G. 1994. Four years rearing experiments on the Mediterranean red 

coral. Biologia Marina Mediterranea 1: 413-420.

Cerrano, C., Bavestrello, G., Cattaneo-Vietti, R. 1999. Nuove esperienze sul trapianto e sugli effetti 

del prelievo del corallo rosso in Mar Ligure. In: Cicogna, F., Bavestrello, G., CattaneoVietti, R. (Eds.) Red Coral and other Octocorals Biology and Protection. Ministero per le 

Politiche Agricole, Roma: 57-73. 

Cerrano, C., Bavestrello, G., Bianchi, C.N., Cattaneo-Vietti, R., Bava, S., Morganti, C., Morri, C., 

Picco, P., Sara, G., Schiaparelli, S. 2000. A catastrophic mass-mortality episode of gorgonians 

and other organisms in the Ligurian Sea (NW Mediterranean). Ecological Letters 3: 284-

293.

Cerrano, C., Bavestrello, G. 2008. Medium-term effects of die-off of rocky benthos in the Ligurian 

Sea. What can we learn from gorgonians? Chemistry and Ecology 24: 1-10.

Cicogna, F., Cattaneo-Vietti, R. 1993. Red coral in the Mediterranean Sea: Arts, History and Science. 

Ministero delle Risorse Agricole, Alimentari e Forestali, Roma, 1-263.

Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. 1999. Red Coral and other Octocorals Biology and 

Protection. Ministero per le Politiche Agricole, Roma, 1-338.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: insights into conservation and management 

needs. Journal of Animal Ecology 71: 966-978.

Garrabou, J., Perez, T., Sartoretto, S., Harmelin, J.G. 2001. Mass mortality event in red coral Corallium 

rubrum populations in the Provence region (France, NW Mediterranean). Marine Ecology 

Progress Series 217: 263-272. 

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in precious red coral Corallium rubrum (L.). Coral Reefs 23: 423-432.

Marchetti, R. 1965. Ricerche sul Corallo Rosso della Costa Ligure e Toscana. Il Promontorio di 

Portofino. Rendiconti dell’Istituto Lombardo di Scienze e Lettere B 99: 279-316.

157
Perez, T., Garrabou, J., Sartoretto, S., Harmelin, J.G., Francour, P., Vacelet, J. 2000. Mortalité massive 

d’invertébrés marins: un événement sans précédent en Méditerranée nord-occidentale. 

Comptes rendus de l’Académie des sciences Ser III Life Sci 323:853-865.

Roghi, G. 1966. La febbre rossa Mondo sommerso, 6, anno VIII.

Rossi, S., Tsouni,s G., Orejas, C., Padrón, T., Gili, J.M., Bramanti, L., Teixidó, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Marine Biology 54: 533-545.

Russo, G., Ulianich, L., Cicogna, F. 1999. Autotomia e frammentazione, una nuova strategia 

riproduttiva per il corallo rosso. In: Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. (Eds.) 

Red Coral and other Octocorals Biology and Protection. Ministero per le Politiche Agricole, 

Roma: 45-56. 

Santangelo, G., Carlietti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Marine Ecology 

Progress Series 248: 99-108.

Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean). Biological Conservation 121:391-397.

Tsounis, G., Rossi, G., Gili, J.M., Arntz, W. 2006. Population structure of an exploited benthic 

cnidarian: the case study of red coral (Corallium rubrum L.). Marine Biology 149: 1059-

1070.

ZIibrowius, H., Marques, V., Grasshoff, N. 1984. La ripartition du corail rouge dans l’Atlantique. 

Tethys 11: 163-170.

* This is an improved version of a paper presented to the 1st Mediterranean symposium on the conservation 

of the coralligenous and other calcareous bio-concretions, 15-16 January 2009 Tabarka, Tunis, UNEPMAP-RAC/SPA

158
LONG-TERM SETTLEMENT PLATES: 

SOME CLUES FOR DEMOGRAPHY AND RESTORATION 

OF MEDITERRANEAN RED CORAL (CORALLIUM RUBRUM) 

SHALLOW WATER POPULATIONS

1,2Bramanti L., 1

Vielmini I., 1Santangelo G.

1

 Department of Biology, University of Pisa, via Volta 6, 56126 Pisa, Italy

2

 ICM - CSIC, Passeig Maritim de la Barceloneta 37-49, 08003 Barcelona, Spain

Abstract

Recovery of coral populations is a slow process, especially in Mediterranean Sea where octocorals, 

characterized by slow growth rates, are dominant. Until now, restoration techniques are based on 

transplantation, a technique requiring high degree of manipulation leading to high colony mortality. 

Mediterranean red coral (Corallium rubrum, L 1758) is a long-lived, slow-growing gorgonian, 

endemic to Mediterranean rocky shores. Its high economic value determined over-harvesting 

which brought many coastal, shallow-water populations to depletion. Moreover, shallow-water 

populations could be extremely vulnerable to mass mortality events putatively linked to global 

warming. Therefore, these populations need conservation and restoration actions. Due to their 

fragility, transplantation of adult colonies does not give any effective result. Within this framework 

we set out a method suitable to foster red coral recruitment on artificial, carbonatic substrates 

(marble tiles), following a patented protocol. This protocol allowed red coral larvae settlement and 

growth on marble tiles. In the framework of a study carried out in Italy with 54 100 cm2

 marble 

tiles, overall, 388 settlers colonized the tiles and their densities varied between 12.37 ± 6.1 and 

2.75 ± 2.4 dm-2. Even if mortality affected these colonies (24.35 ± 9.12 colonies % y-1), after 4 

years the tiles still harbored a persistent population (19 ± 4.97 dm-2) with positive net-recruitment 

rates (recruitment-mortality). As permanently colonized tiles may easily be removed and fixed 

again in new areas by a central screw, red coral re-colonization may be fostered by transferring 

and re-fixing tiles in areas depleted of red coral colonies. Moreover, none of the colonies settled on 

such tiles was affected by boring sponges, which are one of the main sources of red coral mortality 

and also reduce their economic value. These findings suggest marble tiles could be a useful tool 

both to study recruitment and to foster restoration of red coral shallow-water populations.

Mediterranean red coral (Corallium rubrum, L 1758) is a 

colonial Anthozoan endemic to the Mediterranean Sea (Fig. 

1). Its bathymetric range is comprised between 10 and 800 

m depth (Zibrowius et al. 1984; Costantini et. al. 2009). Due 

to the high economic value of its axial, carbonate skeleton, 

this coral has been harvested and traded since ancient times 

(Fig. 2). Nowadays a reduction of overall fishing yield was 

recorded and in the last two decades the Mediterranean 

yield reduced by 2/3 (Santangelo and Abbiati 2001; Tsounis 

et al. 2007). Red coral is a long-lived, gonochoric, brooder 

species, whose larvae (planulae) do not travel very far 

from the parental colonies (Vighi 1972; Weinberg 1979). 

This species is structured in genetically differentiated, 

Fig. 1. Corallium rubrum colony 

with open polyps.

159
self-seeding populations (Abbiati et al. 1993). 

Reproduction is discrete and occurs within a limited 

time interval in early summer (Santangelo et al. 

2003). On a bathymetric distribution point of view, 

two kind of populations can be distinguished:

• Coastal shallower populations (up to 70 

meters depth), characterized by colonies with 

small size, few branches, a small average basal 

diameter, a high density (Tsounis et al. 2007), 

and a limited economic value due to colony 

small size and to parasitic boring sponges 

infection (more than 50% of the colonies are 

affected by boring sponges; Corriero et al. 

1997).

• Deep populations (below 70 m depth) are 

characterized by large, sparse colonies with a 

high economic value. 

Studies on different shallow-water populations 

indicate their vulnerability to mortality events due 

to global warming. In late summer 1999 and 2003, a 

mass mortality affected several benthic invertebrates 

(mainly sponges and gorgonians) in the western Mediterranean Sea (Cerrano et al. 2000). Adult 

colonies of a red coral population dwelling at Marseille were heavily affected by this event 

(Garrabou et al. 2001) while in the population of Calafuria (Livorno, Italy) recruitment, more than 

adult colonies, suffered mortality (Bramanti et al. 2005; Santangelo et al. 2007). At the same time, 

an anomalous temperature increase has been recorded in the same geographic area (Cerrano et al. 

2000; Romano et al. 2000). Such anomaly was considered to be (the putative) cause causing the 

mass mortality (Coma and Ribes 2003).

Data on shallow populations suggest moreover that colony size reduced (and probably density 

increased) due to the over-harvesting these populations faced (Garrabou and Harmelin 2002; 

Tsounis et al. 2007; Bramanti et al. 2009). Consequently, several coastal shallow-water populations 

have been brought to depletion. Therefore these populations need conservation and restoration 

actions. 

To conservation purposes we perform a demographic approach to population through life 

history tables and population demographic models (Santangelo et al. 2004, 2007; Bramanti et 

al. 2009). Life history tables, which merge mortality and reproduction of individuals within a 

population, are suitable for demographic studies of long-lived species and have been proven to 

be (useful) appropriate conservation tools approaching overexploited species (Gotelli 1991; Ebert 

1999; Fujiwara and Caswell 2001; Santangelo et al. 2004, 2007). Life tables moreover represent 

indispensable tools for compiling transition matrices (non-linear Leslie matrices of population 

growth), which in turn provide projections of population trends over time in stage-subdivided 

populations (Tuljapurkar and Caswell 1997).

Fig. 2. Precious handcraft objects 

obtained from Corallium rubrum axial 

skeleton.

160
To restoration purposes we patented a method to avoid manipulation of colonies. Mediterranean 

red coral is characterized by a slow growth rate (Marschal et al. 2004; Torrents et al. 2005) and 

therefore the recovery of depleted populations is a 

slow process. Transplantation of adult colonies is 

a restoration technique normally used in tropical 

corals (Oren and Benayahu 1997). This technique is 

based on the manipulation of the colonies harvested 

in a place and transplanted to a different one. This 

manipulation represents a stress that result in a high 

mortality of transplanted colonies. In the case of 

Corallium rubrum, moreover, fragility of colonies 

makes transplantation techniques not effective. 

To obtain reliable data on early life history of 

Corallium rubrum and to develop a restoration 

technique avoiding manipulation, we set out a 

method suitable to foster red coral recruitment 

on artificial, carbonatic substrates according to a 

patented protocol (marble tiles, Fig. 3). 

This protocol allowed red coral larvae settlement and growth. Marble was chosen because its 

chemical composition is similar to that of the biogenic substrate (coralligenous) on which red coral 

settles, which is composed mainly of calcareous algae and carbonate skeletons of invertebrates 

(Bramanti et al. 2003). White tiles, moreover, made for easy identification of red coral settlers and, 

being permanent, enabled us to follow the life history of both individual settlers and each cohort 

for an extended period of time. Tiles are photographically sampled for the period they remain 

submerged so that, when tiles are removed, all the settled colonies can be divided in different 

cohorts (one cohort for each year of submersion) basing on the identification provided by the 

photographic sampling. Diameter, height and number of polyps of each colony in each cohort are 

measured, and the growth rates determined. This procedure allows us to determine colony yearly 

growth rates (the ratio between the average size of the colonies in two successive cohorts) and the 

average annual growth rates permitting to group the population colonies into size/age classes. 

In the framework of a 4 years study carried out in Calafuria (Italy), 54 marble tiles (100 cm2 each) 

have been placed on the vault of crevices where adult colonies were present following the patented 

method. Tiles were put in place at the end of June, some weeks before red coral larval release, to 

allow a bioorganic film to cover their surfaces (Keough and Raimondi 1996) and photographically 

sampled monthly. This procedure provided us with a settlement and survival data series. Four 

settlement cycles and the subsequent survival of newly settled colonies were thus followed monthly 

for a 4-year period.

Overall, 388 Corallium rubrum larvae settled on the tiles. Density of settled colonies varied 

between 12.37 ± 6.1 and 2.75 ± 2.4 dm2. Mortality affected 66.4% of the Year 1 cohort. The yearly 

survival of cohorts (the survival rate between two consecutive years) was similar throughout the 

different years 76.4 ± 7.1 (SD) %. The highest mortality affected all cohorts during their first 

year of life. Despite the mortality, after 4 years the tiles still harbored a persistent population 

Fig. 3. White marble tiles are fixed to the 

rocky wall by a central steel screw onto the 

vault of crevices, in which red coral adult 

colonies are present.

161
(19 ± 4.97 dm-2) with positive net-recruitment rates 

(recruitment-mortality, Fig. 3).

Colonized tiles may easily be removed from the 

substrate and fixed again in new areas (by a central 

screw). This technique does not imply colony 

manipulation as colonies settled on tiles are not even 

touched. Re-colonization of red coral populations 

may be fostered by removing colonized tiles from a 

population in good conditions and re-fixing them in 

depleted areas. 

Another source of mortality for red coral colonies 

is represented by the infection of boring sponges 

(Corriero et al. 1997) producing holes in the axial 

skeleton resulting in a reduction of economic value 

of the colonies and in a higher mortality rate of 

infected colonies. After the 4 years study, colonies settled on tiles were examined for the presence 

of boring sponges. Results showed that none of the colonies was affected while more than 50% of 

colonies of the same age in natural substrate resulted to be affected (Corriero et al. 1997). These 

findings suggest that marble tiles could be a useful tool both to conservation purposes, giving 

important demographic data on the first years of life of the colonies, and to restoration purposes of 

red coral shallow-water populations by means of a transplantation technique that does not imply 

colony manipulation. 

References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean alcyonarian Corallium rubrum. Mar. Ecol. Prog. Ser. 95: 

245-250.

Bramanti, L., Magagnini, G., Santangelo, G. 2003. Settlement and recruitment: the first stages in the 

life cycle of two epibenthic suspension feeders. Ital. J. Zool. 70: 175-178. 

Bramanti, L., Magagnini, G., DeMaio, L., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean Red Coral Corallium rubrum (L 1758), a four-year study. J. 

Exp. Mar. Biol. Ecol. 314: 69-78.

Bramanti, L., Rossi, S., Tsounis, G., Gili, J.M., Santangelo, G. 2007. Settlement and early survival 

of red coral on settlement plates: some clues for demography and restoration. Hydrobiol. 

580: 219-224.

Bramanti, L., Iannelli, M., Santangelo, G. 2009. Mathematical modelling for conservation and 

management of gorgonians corals: young and olds, could they coexist? Ecological Modelling 

220 (2009): 2851-2856.

Cerrano, C., Bavestrello, G., Nike-Bianchi, C., Cattaneo-Vietti, R., Bava, S., Morganti, C., Morri, 

C., Picco, P., Sara, G., Schiaparelli, S., Siccardi, A., Spogna, F. 2000. A catastrophic massmortality episode of gorgonians and other organisms in the Ligurian Sea (North-western 

Mediterranean), summer 1999. Ecol. Lett. 3: 284-293.

0

5

10

15

20

25

30

1998 1999 2000 2001

recruits/dm2

Fig. 3. Net recruitment rate (recruitment 

minus mortality). Error bars represent 

SD. Drawn from Bramanti et al. 2005.

162
Coma, R., Ribes, M. 2003. Seasonal energetic constrains in Mediterranean benthic suspension 

feeders: effects at different levels of ecological organization. Oikos 101: 205-215.

Corriero, G., Abbiati, M., Santangelo, G. 1997. The sponge complex inhabiting a Mediterranean red 

coral population. PSZN Mar. Ecol. 18: 147-155. 

Costantini, F., Taviani, M., Remia, A., Pintus, E., Schembri, P.J., Abbiati, M. 2009. Corallium rubrum

in the deep sea: genetic characterization and comparison with shallow water colonies. Mar. 

Ecol. (in press).

Ebert, T.A. 1999. Plant and animal populations. Methods in demography. Ac. Press, S. Diego CA.

Fujiwara, M., Caswell, H. 2001. Demography of endangered North Atlantic right whale. Nature 414: 

537-541.

Garrabou, J., Perez, T., Sartoretto, S., Harmelin, J.G. 2001. Mass mortality event in red coral 

(Corallium rubrum) populations in the Provence region (France, NW Mediterranean). Mar. 

Ecol. Prog. Ser. 217: 263-272.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in NW Mediterranean: insights into conservation and management needs. 

J. An. Ecol. 71: 966-978.

Gotelli, N.G. 1991. Demographic models for Leptogorgia virgulata, a shallow water gorgonian. 

Ecology 72: 457-461. 

Marschall, C., Garrabou, J., Harmelin, J.G. 2001. A new method for measuring growth and age of the 

precious red coral Corallium rubrum (L). Coral Reefs 23: 423-432. 

Romano, J.C., Bensoussan, N., Walid, A.N.Y., Arlhac, D. 2000. Anomalie thermique dans les eaux du 

golfe de Marseille durant l’été 1999. Une explication partielle de la mortalité d’invertebrés 

fixés? C R Acad Sci Paris, Sciences de la vie/Life sciences 323: 415-427.

Oren, U., Benayahu, Y. 1997. Transplantation of juveniles corals: a new approach for enhancing 

colonization of artificial reefs. Mar. Biol. 127: 499-505.

Santangelo, G., Abbiati, M. 2001. Red coral: conservation and management of an overexploited 

Mediterranean species. Aquat. Cons. Mar. Freshw. Ecosys. 11: 253-259.

Santangelo, G., Carletti, E., Maggi, E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Mar. Ecol. Prog. 

Ser. 248: 99-108.

Santangelo, G., Maggi, E., Bramanti, L., Bongiorni, L. 2004. Demography of the overexploited 

Mediterranean red coral Corallium rubrum. Sci. Mar. 68: 199-204.

Santangelo, G., Bramanti, L., Iannelli, M. 2007. Population dynamics and conservation biology of 

the over-exploited Mediterranean Red coral. J. Theor. Biol. 244: 416-423. 

Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean). Biol. Cons. 121: 391-397.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W.E. 2007. Red coral fishery in the Costa Brava (NW 

Mediterranean): case study of the overharvested precious coral. Ecosystems 10: 975-986.

163
Tuljapurkar, S., Caswell, H. 1997. Structured-population models in marine, terrestrial, and freshwater 

systems. Chapman and Hall, N.Y.

Vighi, M. 1972. Etude sur la reprodution du Corallium rubrum. Vie et Milieu 23: 21-32.

Weinberg, S. 1979. The light dependent behaviour of planulae larvae of Eunicella singularis and 

Corallium rubrum and its implications for octocorallian ecology. Bijdr. Dierkd. 49: 16-30.

Zibrowius, H., Montero, M., Grashoff, M. 1984. La ripartition du Corallium rubrum dans l’Atlantique. 

Thetis 11: 163-170. 

164
BIODIVERSITY AND ECOLOGY OF SPONGE 

BORING INTO PRECIOUS CORALS:

A WORLDWIDE OVERVIEW

Barbara Calcinai1, Carlo Cerrano2, Nozomu Iwasaki3, Giorgio Bavestrello1

1

DiSMar, Università Politecnica delle Marche, Via Brecce Bianche, 60131 Ancona, Italy

2

DipTeRis, Università di Genova, Corso Europa 26, 16132 Genova, Italy

3

Usa Marine Biological Institute Kochi University, Kochi, Japan

Abstract

Boring sponges live inside calcareous substrata where they produce holes and galleries filled 

by sponge choanosome. A group of these species is associated to precious corals both in the 

Mediterranean and Pacific Ocean. The sponges produce a spongiosus pattern in the coral scleraxis 

resulting in a considerable decreasing in its economic value. Stoeba, Holoxea, Alectona, Delectona, 

Thoosa, Spiroxya, Cliona, Dotona, and Aka are the most involved genera. Alectona and Spiroxya

(family Alectonidae), both with 6 species, are the most common, while Cliona (family Clionaidae), 

widely diffused also in other organic substrata such as coral reefs, is poorly represented. The 

total number of boring species recorded in the different species of precious coral is 27 with only 

two present both in the Mediterranean and Pacific Ocean. Alectonidae family (Alectona, Dotona, 

Delectona, and Spiroxya) accounts for 14 species, suggesting a strong affinity of this family for the 

compact calcitic scleraxis of the precious coral. In particular the genus Spiroxya has the strongest 

affinity for precious coral (with 6 species out of 9, recorded in coral colonies). This affinity is also 

indicated by the geographic distribution of this genus that has exactly the same Tethyan distribution 

as the genus Corallium.

The way of penetration of boring sponges into precious corals is not completely clarified but it is 

stated that sponges are not able to penetrate through the living coenosarc. Therefore the penetration 

of sponges in the coral scleraxis starts from the dead portions of the colonies or from the substratum. 

The petrographic structure of the substratum (calcareous vs. not calcareous) can therefore affect 

the sponge infection of the coral population and, on the base of substrate composition, the level of 

infestation of a bank can be predicted.

The sponge infestation may compromise the survivor-ship of smallest size classes. Although 

generally the boring activity decreases the steadiness of the colonies, it is difficult that large 

specimens are detached from the substrate. On the contrary, when sponges attack newly settled 

specimens (1 mm in diameter), the erosion is able to weak the base of the colonies. According to 

different kinds of substrata the sponge activity may therefore result in a high level of post-larval 

mortality. 

Introduction

The species belonging to Corallium and Paracorallium (Octocorallia, Coralliidae) are known all 

over the world due to their heavy exploitation by the jewel industry. Considering the general slow 

growth rate of these taxa (Bramanti et al. 2005; Roark et al. 2006), there are several international 

projects devoted to their protection and management. The main areas where precious coral fishery 

165
is strongly active are the Mediterranean Sea for the red coral Corallium rubrum (Cicogna et al. 

1999) and the Pacific Ocean for several other species (UICN 2007; Parrish and Baco 2007).

Precious coral colonies are characterized by a scleraxis composed of magnesiferous calcite 

organized in an extremely compact structure of acicular to lamellar calcite crystals that grow 

centripetally, forming fan-like structures (Cortesogno et al. 1999). Often the collected colonies 

cannot be carved owing to the presence of boring sponges, strongly decreasing the commercial 

value of the coral (Liverino 1984). 

When occupied by boring sponges the scleraxis of the precious coral shows a series of chambers 

interconnected by channels giving rise to a spongiosus pattern in the carbonate. The knowledge of 

the involved sponge species, their relative destructive power, their distribution and their relationship 

with the affected coral colonies is important to help both coral industry and coral scientists to draw 

together adequate measures for management and exploitation of this resource.

Morphology of Perforation

Until now no comparative studies were conduced to detect differences in the morphology of 

the perforations produced by different taxonomic groups of boring sponges. The genus Thoosa

produces in the coral scleraxis irregular, ovoid, and elongated chambers (2-5 mm in diameter) 

connected by canals (Azzini et al. 2007). A similar pattern is shown by Holoxea excavans but the 

chambers are smaller (1-2 mm) (Calcinai et al. 2008). The genus Aka, belonging to the family 

Phlaeodictyidae, produces in the coral scleraxis a completely different boring pattern composed 

by minute oval canals and galleries giving rise to a characteristic spongious aspect of the eroded 

substratum (Calcinai et al. 2003). A different pattern is shown by the species of Alectona and 

Delectona that grow continuously enlarging a single wide boring chamber (Bavestrello et al. 

1996a, 1998).

More detailed morphological data on the erosion micro-patterns recognizable on the wall of 

boring chambers were obtained by SEM observations. The pitting pattern is the result of a two 

step – chemical and mechanical – activity of peculiar etching cells of boring sponges (Rützler and 

Rieger 1973). Previous studies (Calcinai et al. 2003, 2004a) have demonstrated that the recorded 

differences between different species are not related with substrate features. Among the species 

boring in precious coral the pitting patterns were clearly recognizable at least at generic level. 

Cliona produces smooth pits while those of Spiroxya and Holoxea are finger-print like. On the 

contrary, the scars produced by Alectona and Delectona create pits with deep concentric, irregular 

erosion lines. Particularly, Alectona shows a radial pattern that overlaps on the concentric bands, 

making the pits of this genus very distinctive. Finally, in the genus Aka, pits are generally regular, 

with the surface subject to deep grooves that extend to the pit center. There is often an outgrowth 

in the center of the pit. These data are interesting allowing the identification of borers also in dead 

or subfossil coral branches deprived of their living parasites.

Biodiversity and Biogeography

In the Mediterranean Sea, during a series of surveys conducted on different red coral populations 

16 boring species were identified (Melone 1965; Barletta and Vighi 1968; Corriero et al. 1988; 

Calcinai et al. 2002; Calcinai et al. 2004a; Calcinai et al. 2007). 13 species were recorded during 

166
our studies (Bavestrello et al. 1995, 1996, 1998; Calcinai et al. 2001, 2002, 2004b) on the Pacific 

species of precious coral. The total number of species involved in the boring activity in the precious 

coral is 27 with only two species, Dotona pulchella and Alectona verticillata, being common to 

both geographic areas and the genus Delectona is recorded only in the Mediterranean coral (Table 

1). 

The genera more involved are 8: Holoxea, Alectona, Delectona, Thoosa, Spiroxya, Cliona, Dotona, 

Aka but sometimes Jaspis were also recorded. The most represented genera are Alectona and 

Spiroxya (family Alectonidae), both with 6 species, while Cliona (family Clionaidae), which is 

very common in calcareous, organic substrata such as coral reefs, is poorly represented. While 

Alectona has 2 species in the Mediterranean and 5 in the Pacific, Spiroxya has 3 species both in 

the Mediterranean and in the Pacific Ocean. The family Alectonidae (Alectona, Dotona, Delectona 

and Spiroxya) accounts for 14 species, suggesting strong affinity for the compact calcite of the 

precious coral scleraxis. The affinity of the species of this family for precious coral could be 

related to their typical bathophily. 

Nevertheless, while Spiroxya is a genus with a strong affinity for precious coral (with 6 species 

out of 9, recorded in coral colonies), Alectona is common in other organic carbonates. This is 

also indicated by the geographic distribution of these genera: while Alectona has circumtropical 

distribution, the genus Spiroxya has exactly the same Tethyan relict distribution as the genus 

Corallium (Calcinai et al. 2004b).

Way of Penetration

The way of penetration of boring sponges into the precious coral is not completely clarified but it is 

generally accepted that sponges are not able to penetrate through the living coenosarc. Following 

this idea it is possible to hypothesize three different ways of penetration: (i) the substratum, (ii) 

dead portions of the colonies, (iii) the borderline of the base where, probably, the coenenchime is 

thinner. 

In all the examined specimens affected by the sponge erosion the boring chambers are more 

abundant in the base while they strongly reduce and quickly stop in the distal portion of the colony. 

If the colonies show some dead apical branches, they are never affected by the erosion. 

If the penetration arises from the substratum its petrographic structure is able to affect the sponge 

infection. Coral banks living on calcareous substrata, where sponges are able to penetrate would 

be more prone to boring activity. On the contrary, corals living on granitic substrates, which cannot 

be bioeroded, would be free from erosion. Unfortunately, we have not a screening of the percent 

of affected coral colonies in banks settled on different petrographic substrata but, for example, 

unpublished data obtained from a Japanese population of Paracorallium japonicum settled on 

basaltic rocks indicate a very scarce incidence of boring.

On the other hand, studies conduced on the carbonatic substrata of the Portofino Promontory have 

clearly shown that different species of boring sponges have a different level of selectivity toward 

the coralligenous conglomerate substrate or towards the colonies or C. rubrum. For example, Jaspis 

jonstoni and Spiroxya heteroclita were recorded commonly in the conglomerate and, sometimes, 

in the coral colonies. Other species, like Aka insidiosa and Cliona janitrix, were mainly observed 

167
in coral colonies but are rarely able to extend in the surrounding substratum. Spiroxya corallophila 

and Stoeba sp. were only recorded in the corals, while Stoeba plicatus was only present in the 

conglomerate. These data suggest that probably the way of penetration of boring sponges in the 

coral skeleton is different in different species. In more generalistic species, the penetration arises 

from the substratum, while in species specialized in the coral erosion, the penetration probably 

start from the borderline of the coral base. 

Effect on the Demography of Coral Populations

It is evident that the activity of boring sponges in the coral scleraxis decreases its stiffness 

enhancing the possibility of break and detach of the colonies from the substratum. Nevertheless, 

the influence of boring sponges is variable according to the size of the affected colonies. While 

adult colonies cannot become unstable by erosion, the sponge infestation may compromise the 

stability of smallest size classes. In fact, when sponges attack newly settled specimens (1 mm in 

diameter), the erosion is able to weak adhesion of the base of the colonies. According to different 

kinds of substrata the sponge activity may therefore result in a high level of post-larval mortality. 

Detailed studies conduced on the rocky cliff of the Portofino promontory clearly indicate that in 

September, immediately after the summer reproduction of red coral, on the substratum are present 

thousands of just settled juvenile colonies composed by few polyps and with a basal diameter of 

about 1 mm. On the contrary, six month later a large majority of these small sponges are not more 

present on the substratum. A survey conduced on these juveniles specimens indicate that they are 

affected by sponge erosion, probably arising from the substratum. This fact is strongly supported 

by the evidence that small colonies, settled on artificial plastic panels, are not affected by the 

erosion. It is probable that the activity of boring sponges represents the main constrain, at least on 

carbonatic substrata, in the regulation of the population of red coral (Cerrano et al. 2001).

References

Azzini, F., Calcinai, B., Iwasaki, N., Bavestrello, G. 2007. A new species of Thoosa (Demospongiae, 

Hadromerida) excavating precious coral Corallium sp. from Midway. Italian Journal of 

Zoology 74 (4): 405-408.

Barletta, G., Vighi, M. 1968. Ricerche sul corallo rosso: 5-poriferi perforanti. Rendiconti Istituto 

Lombardo Accademia di Scienze e Lettere, Milano. Scienze Biologiche e Mediche 102: 

145-159. 

Bavestrello, G., Calcinai, B., Sarà, M. 1995. Two new species of Cliona (Porifera, Demospongiae) 

boring the scleraxis of Corallium elatius from the western Pacific. Bollettino di Zoologia, 

62: 375-381.

Bavestrello, G., Calcinai, B., Cerrano, C., Pansini, M. Sarà, M. 1996. The taxonomic status of some 

Mediterranean clionids (Porifera, Demospongiae) according to morphological and genetic 

characters. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique (Ser. Biologie), 

Bruxelles 66: 185-195.

Bavestrello, G., Calcinai, B., Sarà, M. 1996a. Delectona ciconiae n. sp. (Porifera, Demospongiae) 

boring into the scleraxis of Corallium rubrum. J. Mar. Biol. Ass. U.K. 76: 867-873.

168
Bavestrello, G., Calcinai, B., Cerrano, C., Sarà, M. 1998. Alectona species from Western Pacific 

(Demospongiae: Clionidae). Journal of the Marine Biological Association of the United 

Kingdom 78: 59-73.

Bramanti, L., Magagnini, G., De Maio, L., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean red coral Corallium rubrum (L 1758), a 4-year study. Journal 

of Experimental marine Biology and Ecology 314: 69-78.

Calcinai, B., Bavestrello, G., Cerrano, C., Sarà, M. 2001. Boring sponges living into precious corals 

from the Pacific Ocean. Italian Journal of Zoology 68: 153-160.

Calcinai, B., Cerrano, C., Bavestrello, G. 2002. A new species of Scantiletta (Demospongiae, 

Clionaidae) from the Mediterranean precious red coral with some remarks on the genus. 

Bulletin of Marine Science 70: 919-926.

Calcinai, B., Arillo, A., Cerrano, C., Bavestrello, G. 2003. Taxonomy-related differences in the 

excavating micro-patterns of boring sponges. J. Mar. Biol. Ass. UK 83: 37-39. 

Calcinai, B., Bavestrello, G., Cerrano, C. 2004a. Bioerosion micro-patterns as diagnostic 

characteristics in boring sponges. In: Pansini, M., Pronzato, R., Bavestrello, G., Manconi, 

R. (Eds.) Sponge Science in the New Millennium, Bollettino dei Musei e degli Istituti 

Biologici dell’Università di Genova 68: 229-238.

Calcinai, B., Azzini, F., Bavestrello, G., Iwasaki, N., Cerrano, C. 2004b. Redescription of Alectona 

verticillata (Johnson) (Porifera, Alectonidae) boring into Japanese precious coral. Italian 

Journal of Zoology 71: 337-339.

Calcinai, B., Cerrano, C., Bavestrello, G. 2007. Three new species and one re-description of Aka. 

Journal of the Marine Biological Association of the United Kingdom 87: 1355-1365.

Cerrano, C., Bavestrello, G., Bianchi, C.N., Calcinai, B., Cattaneo-Vietti, R., Morri, C., Sarà, M. 

2001. The role of sponge bioerosion in the Mediterranean coralligenous accretion. In: 

Faranda, F.M., Guglielmo, L., Spezie, G. (Eds.) Mediterranean Ecosystems: Structures and 

Processes, Springer-Verlag, Milano: 235-240.

Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. 1999. Red coral and other Mediterranean octocorals, 

biology and protection. In: Cicogna, F., Bavestrello, G., Cattaneo-Vietti, R. (Eds.) Red 

coral in the Mediterranean Sea: art, history and science. Ministero delle Risorse Agricole, 

Alimentari e Forestali, Roma.

Corriero, G., Pansini, M., Sarà, M. 1988. Boring sponges (Demospongiae, Clionidae) perforating 

Corallium rubrum in the Mediterranean sea. FAO Fishery Reports 413: 73-78.

Cortesogno, L., Gaggero, L., Bavestrello, G., Cerrano, C., Cattaneo-Vietti, R. 1999. Struttura, 

mineralogia, minerochimica e chimismo del corallo rosso. In: Cicogna, F., Bavestrello, 

G., Cattaneo-Vietti, R. (Eds.) Biologia e tutela del corallo rosso e di altri ottocoralli del 

Mediterraneo. Ministero per le Politiche Roma, 83-97.

Liverino, B. 1983. Il Corallo. Ed. Analisi. Bologna: 229 p.

Melone, N. 1965. I poriferi associati a Corallium rubrum (L.) della Sardegna. Annali del Museo 

Civico di Storia Naturale Giacomo Doria 75: 344-358. 

169
Parrish, F.A., Baco, A.R. 2007. State of deep coral ecosystems in the U.S. Pacific Islands Region: 

Hawaii and the U.S. Pacific Territories. In: Lumsden, S.E., Hourigan, T.F., Bruckner, A.W., 

Dorr, G. (Eds.) The state of deep coral ecosystems of the United States: 2007. 159-194 

(Chapter 4). U.S. Dep. Commer., NOAA Tech. Memo CRCP-3. Silver Spring, MD 365 p.

Roark, E.B., Guilderson, T.P., Dunbar, R.B., Ingram, L. 2006. Radiocarbon-based ages and growth 

rates of Hawaiian deep-sea corals. Marine Ecology Progress Series 327: 1-14.

Rützler, K., Rieger, G. 1973. Sponge burrowing: fine structure of Cliona lampa penetrating calcareous 

substrata. Mar. Biol. 21: 144-162.

UICN. 2007. Les Analyses UICN/TRAFFIC des propositions d’amendement aux Annexes de La 

CITES pour la Quatorzième session de la Conférence des Parties à la CITES www.iucn.org/

themes/ssc/our_work/wildlife_trade/citescop14/cop14analyses.htm.

170
Table 1. Sponge species boring precious corals (Corallium and Paracorallium) in Mediterranean 

Sea and Pacific Ocean.

Species Locality

Mediterranean Pacific

Holoxea furtive Topsent, 1892 X

H. excavans Calcinai et al., 2001 X

Alectona millari Carter, 1879 X

A. wallichi Carter, 1874 X

A. triradiata Levi & Levi, 1983 X

A. verticillata (Johnson, 1899) X X

A. sorrentini Bavestrello et al., 

1998 X

A. sarai Calcinai et al., 2008 X

Delectona ciconiae Bavestrello et al., 

1996 X

Cliona janitrix Topsent, 1932 X

C. lobata Hancock, 1849 X

C. desimoni Bavestrello et al., 

2005 X

Spiroxya acus (Bavestrello et al., 

1995) X

S. corallophila (Calcinai et al., 2002) X

S. heteroclita (Topsent, 1988) X

S. levispira (Topsent, 1898) X

S. macroxeata (Calcinai et al., 2001)

S. sarai (Melone, 1965) X

Thoosa armata Topsent, 1888 X

T. bulbosa Hancock, 1849 X

T. midway Azzini et al., 2007 X

T. mollis Volz, 1939 X

Dotona pulchella Carter, 1880 X X

Aka labyrinthica (Hancock, 1849) X

A. corallirubri Calcinai et al., 2007 X

A. insidiosa (Johnson, 1899) X

Stoeba spp. X X

171
 BIOMETRICAL ANALYSIS AND OBSERVATIONS ON DEEP RED CORAL

 CORALLIUM RUBRUM (L.) IN N.W. SARDINIA SHOALS (ITALY)

L. A. Chessa1

, M. Scardi2

1

 Dipartimento di Protezione delle Piante – Ecologia,

Università di Sassari, via E. De Nicola, 9 - 07100 Sassari, Italy,

2

 Dipartimento di Biologia, Università di Roma ‘‘Tor Vergata,’’ Roma, Italy

Monitoring deep red coral shoals and studying size structure of this species are both very important 

tasks in areas of intense harvesting. Since fishing dramatically affects the size of the largest 

colonies, the colony size is a good indicator of harvesting impact. A study was performed on 

coralligen shoals, emerging from a bioclastic sandy bottom, 7 nm SW of Alghero (Sardinia, Italy). 

Registered divers practice coral fishing in this area, although only threatening the largest colonies. 

The study analyzed some morphometrical parameters of nearly 200 colonies, which were collected 

at 130 m depth. A linear model to estimate wet weight from the number of apices in the colonies 

was defined, while observations on deep coral settlement and on the state of the ramifications 

were also performed. We concluded that the red coral of these fishing grounds seem not to be 

overexploited and that most colonies are in good health. It seems quite clear that the overall weight 

of a colony can be effectively inferred on the basis of the number of apices.

172
EFFECTIVE LARVAL DISPERSAL AND SCALES OF 

CONNECTIVITY IN CORALLIUM RUBRUM

F. Costantini, C. Fauvelot, E. Pintus, M. Abbiati

Centro Interdipartimentale di Ricerca per le Scienze Ambientali

in Ravenna and Dipartimento di

Biologia Evoluzionistica Sperimentale, 

University of Bologna, Via S. Alberto 163, I – 48100 Ravenna, Italy.

E-mail: federica.costantini@unibo.it

Dispersal patterns play a fundamental role in determining the dynamics of marine populations 

defining the extent of breeding units as well as the level of connectivity among them. Persistence, 

resilience, and recovery of marine populations affected by anthropogenic and natural threats 

depend on the supply and survival of recruits. Understanding spatial scales of genetic structuring 

provides indirect estimates of species dispersal ability. Limited dispersal reduces chances for 

re-colonization and could promote genetic drift, reduction of the evolutionary potential of the 

species, and increase risk of population extinction. Corallium rubrum is a long-lived, slowgrowing gorgonian. Red coral is one of the most exploited subtidal sessile invertebrates in the 

Mediterranean Sea, mostly owing to human harvesting. The main goal of this study is to assess the 

level of connectivity and effective larval dispersal capability of the species using microsatellites. 

Scales of genetic structuring were analyzed in C. rubrum samples collected in the Ligurian Sea in a 

range from meters to 100s of meters. At spatial scales of 10s of meters, a surprisingly strong pattern 

of genetic structuring has been found, supporting the hypothesis of limited dispersal ability of the 

planulae and suggesting a high inbreeding rate. Significant heterozygote deficiencies and chaotic 

genetic structuring at spatial scales of 1 meter have been also detected. These findings suggest 

that biological processes (e.g. pre- and post-settlement mortality and stochasticity of reproductive 

success) may favor the occurrence of genetically distinct pools of recruits at meter distances. The 

outcomes of these studies are of great importance to infer about red coral populations resilience 

and to identify the appropriate scales for the conservation of this species.

173
DEEP-WATER CORALLIUM RUBRUM (L. 1758) FROM THE MEDITERRANEAN SEA: 

PRELIMINARY GENETIC CHARACTERISATION

F. Costantini, M. Taviani, A. Remia, E. Pintus, P. J. Schembri, M. Abbiati

Centro Interdipartimentale di Ricerca per le Scienze 

Ambientali in Ravenna and Dipartimento di Biologia 

Evoluzionistica Sperimentale, University of Bologna, 

Via S. Alberto 163, I – 48100 Ravenna, Italy.

E-mail: federica.costantini@unibo.it

The precious red coral Corallium rubrum lives in the Mediterranean Sea and adjacent eastern 

Atlantic Ocean on subtidal hard substrates. Corallium rubrum is a long-lived gorgonian coral that 

has been commercially harvested since ancient times for its red axial calcitic skeleton and which, at 

present, is thought to be in decline because of overexploitation. The depth distribution of C. rubrum 

is known to range from ca. 15 – 300 m. Recently, live red coral colonies have been observed in the 

Strait of Sicily at depths of ca. 600 – 800 m. This record sheds new light on the ecology, biology, 

biogeography, and dispersal mechanism of this species and calls for an evaluation of the genetic 

divergence occurring among highly fragmented populations. A genetic characterization of the deep 

sea red coral colonies has been done to investigate biological processes affecting dispersal and 

population resilience, as well as to define the level of isolation ⁄ differentiation between shallow 

and deep-water populations of the Mediterranean Sea. Deep-water C. rubrum colonies were 

collected at two sites (south of Malta and off Linosa Island) during the cruise MARCOS of the 

R⁄V Urania. Collected colonies were genotyped using a set of molecular markers differing in their 

level of polymorphism. Microsatellites have been confirmed to be useful markers for individual 

genotyping of C. rubrum colonies. ITS-1 and mtMSH sequences of deep-water red coral colonies 

were found to be different from those found in shallow water, suggesting the possible occurrence of 

genetic isolation among shallow- and deep-water populations. These findings suggest that genetic 

diversity of red coral over its actual range of depth distribution is shaped by complex interactions 

among geological, historical, biological, and ecological processes. 

174
FEEDING CHARACTERISTICS OF THE RED CORAL 

CORALLIUM RUBRUM

C. Ferrier-Pagès

Centre Scientifique de Monaco

Corallium rubrum is a characteristic gorgonian of the Mediterranean Sea. Quantitative data on 

prey ingestion and selection by this species are important issues for defining its ecological role, 

and also explaining its distribution and abundance. Whereas several studies have examined feeding 

habits of gorgonians in general, only two have been performed on C. rubrum specifically (Tsounis 

et al. 2006; Picciano and Ferrier-Pagès 2007). Tsounis et al. (2006) found high feeding rates of C. 

rubrum on detrital particulate organic matter and copepods, which each represented from 15 to 

50% of the diet of the red coral. In terms of carbon, this food can bring up to 3192 ng C polyp-1 

d-1 to C. rubrum. There was, however, a high annual variability in the rates of prey capture, 

compared to other gorgonians. Picciano et al. (2007) investigated the feeding rates on smaller prey, 

in the pico-nanoplankton size. The effects of flow, prey concentration and seawater temperature 

were tested. It was shown that C. rubrum was able to prey on both pico- and nanoplankton cells, 

with a preference for flagellates, which represented from 43 to 70% of the C and N ingested from 

small prey. Flow speed (2, 6, and 11 cm s-1) had no effect on grazing rates. Conversely, these 

rates increased with the increase in pico-and nanoplankton concentration and with temperature. 

In the range of concentrations typically found in the Ligurian Sea, the ingestion of pico- and 

nanoplankton can bring 148 ng C (polyp)-1 d-1 and 28 ng N (polyp)-1 d-1 to the red coral. This 

type of food represents only ca. 4.5% of the total carbon gained by C. rubrum from the different 

sources, but might be important in terms of nitrogen, phosphorus, and other essential elements. 

175
SMALL-SCALE DISTRIBUTION OF JAPANESE RED CORAL

OFF AMAMI ISLAND, SOUTHERN JAPAN 

N. Iwasaki1, T. Fujita2, G. Bavestrello3, R. Cattaneo-Vietti4

1 Usa Marine Biological Institute, Kochi University, Japan

2 National Museum of Nature and Science,Tokyo, Japan

3Università Politecnica delle Marche, Italy

4

Università di Genova, Italy

Using a ROV, small-scale distribution patterns of Japanese red coral (Paracorallium japonicum) 

populations were investigated at the depths of 196 – 211 m off Amami Island, southern Japan in 

March 2009. The ROV navigated 1158 and 1475 m covering the areas of 23,160 and 29,490 m2 at 

each site. The average density was 0.47 colonies/100 m2

. Variance/average ratio and Morisita’s Iδ 

indicated that the distribution pattern of Japanese red corals was aggregated.

An additional remark on economic aspects of Japanese precious corals will be reported.

176
RADIOCARBON-BASED AGES AND GROWTH RATES 

OF HAWAIIAN CORALLIUM SP. 

E. Brendan Roark1

, T. P. Guilderson2,3, R. B. Dunbar4

1

Department of Geography, Texas AandM University, College Station, TX 77843, USA

2

Center for Accelerator Mass Spectrometry, Lawrence Livermore National Laboratory, 

Livermore, CA, USA

3

Department of Ocean Sciences and Institute of Marine Sciences, 

University of California, Santa Cruz, Santa Cruz, CA, USA

4

Department of Environmental Earth Systems Science, Stanford University, Stanford, CA, USA

E-mail: broark@geog.tamu.edu

The radial growth rates and ages of the Hawaiian deep-sea coral Corallium sp. were determined 

using radiocarbon measurements. Specimens of Corallium secundum were collected from 450 ± 

40 m depth at the Makapuu deep-sea coral bed off the southeast coast of Oahu, Hawaii, USA, using 

a submersible (PISCES V) from the Hawaii Undersea Research Laboratory. The primary source 

of carbon to the calcitic C. secundum was determined to be in situ dissolved inorganic carbon 

(DIC). Using ‘bomb 14C’ time markers we calculated radial growth rates of 170 μm yr-1 and ages 

of 67 ± 9 to 71 ± 9 years for two specimens up to 28 cm tall. These radial growth rates are similar 

to the radial growth rates (110 ± 20 μm yr-1) of an Atlantic specimen determined by excess lead210 (Pbex) dating (Druffel et al. 1990). The calculated age (71 ± 9 years) for the one specimen 

collected intact with a height of 28 cm is two times larger than the age calculated (29 years) based 

on equations correlating colony height (length) and age based on growth ring counts (Grigg 1976, 

2002). Examination of thin cross sections (petrographic method) in these specimens yielded a 

maximum of 7 density growth rings. These results suggest that further research is required to refine 

the radiocarbon results and validate periodicity of any growth rings. Finally, we will present some 

preliminary results that Sr/Ca measurements in C. secundum specimens may be a viable proxy of 

sub-surface temperature variability.

177
BIOMINERALIZATION IN CORALLIUM RUBRUM: 

ORGANIC MATRIX COMPARATIVE STUDY OF SCLERITES AND SKELETON 

S. Tambutté 

Centre scientifique de Monaco

Biomineralization is the dynamical and physiological process by which organisms transform ions 

in solution into a solid structure: the biomineral. Biominerals are composed of two fractions: a 

mineral fraction and an organic fraction named the organic matrix (OM). This organic matrix 

is mainly composed of proteins, lipids, and sugars. It is responsible for the nucleation, growth, 

orientation of the crystals, as well as the micro- and macro-architecture of the final skeletal 

product. The comparative study of various animal models is necessary for a better understanding 

of biomineralization. The genus Corallium, including the red coral Corallium rubrum, is a very 

interesting model to study biomineralization since it possesses two different skeletal structures 

with very different shapes: an axial skeleton and sclerites. The comparative study of proteins 

present in the OM of axial skeleton and sclerites has allowed us to treat a mechanistic aspect of 

biomineralization within a same species. Moreover we have determined that part of the color of 

the skeleton and sclerites are due to pigments present in the OM.

178
DETERMINATION OF CORALLIUM RUBRUM POPULATION AGE STRUCTURE 

Vielmini I.1°, Bramanti L. 1,2, Tsounis G.2, Rossi S.3, Gili J.M.2, Cattaneo-Vietti R. 4, Santangelo G.1

1

 Department of Biology, University of Pisa, via Volta 6, 56126 Pisa, Italy

2

 ICM-CSIC, Passeig Maritim de la Barceloneta 37-49, 08003 Barcelona, Spain

3

 ICTA, Autonomous University of Barcelona, Cerdanyola del Vallés 08193 Barcelona, Spain

4

 Dip.Te.Ris., University of Genova, C. so Europa 26, 16132 Genova, Italy

°E-mail: ivielmini@biologia.unipi.it

Introduction 

Management approaches based on a detailed knowledge of specific population demographic 

features are fundamental to ensure long-term exploitation sustainability (Knittweis et al. 2009). 

Demography focuses on the specific dynamics of each population, allowing to project population 

structure over time. This approach allows setting out local conservation strategies for wildlife and 

for exploited species structured into discrete local populations, such as the Mediterranean red coral 

(Abbiati et al. 1993). Regarding corals, long-term studies are scarce due to their slow growth and 

longevity (Connell 1997; Garrabou and Harmelin 2002).

Because of its high economic value, the precious Mediterranean red coral Corallium rubrum 

(Cnidaria, Octocorallia, Gorgonacea) has been harvested for more than 2000 years and its shallowwater populations (living between 20 and 70 m depth), indicate a regression and even a partial 

economic extinction (Tsounis et al. 2007). Demographic studies based on sound age structure and 

growth rate assessment in different populations are nevertheless scarce (Santangelo et al. 2009). 

In this study we compared the age structure and the basal growth rate of two shallow Corallium 

rubrum populations dwelling in two marine protected areas.

Colony and Population Age Assessment

In this study, a sclerochronological approach has been applied to supply basic knowledge on colony 

growth rate and population age structure of two shallow Corallium rubrum populations.

These dwell within two distinct Marine Protected Areas in Spain and Italy: Cap de Creus (Costa 

Brava, North-Western Mediterranean; 42°29’,21 N; 3°30’,18 E, Spain) and Portofino (Eastern 

Ligurian Sea 44°18’,18 N 09°12’,83 E, Italy). These populations are differently managed: in 

Cap de Creus, C. rubrum professional harvesting is regulated, while in Portofino harvesting is 

banned since 1999. Despite this, illegal harvesting by poacher occurs in the Spanish area, seriously 

affecting the local Mediterranean red coral population.

The age of the Corallium rubrum colonies has been determined according to an indirect method by 

counting growth rings in thin sections (50 μm thick) of the colony axial skeleton (Marschal et al. 

2004). This sclerochronological technique labels the circular layer of the organic matrix deposed 

each year (Fig. 1).

179
According to Marschal et al. (2004), each 

growth ring corresponds to a year of colony life 

except for the first 3 or 4 years when the calcium 

carbonate deposition of the core of the colony is 

still in progress, resulting as an irregular shape in 

the centre of the thin section.

Colony age corresponds to the number of rings 

counted in the axial skeleton to which 4 further 

years have be added, i.e. the first years of colony 

life. The annual diameter growth rate is the 

quotient of the colony basal diameter and the 

estimated age.

Populations Age Structure

The average population age is 25.3 ± 6.5 SD year and 18.7 ± 3.2 SD year, respectively, at Portofino 

and Cap de Creus. The oldest colony detected in Portofino is aged 39, while the oldest one in Cap 

de Creus is of age 27.

The estimated annual diameter growth rate (average basal diameter/colony age) is 0.22 ± 0.04 SD 

mm/yr for the population of Portofino and 0.24 ± 0.06 SD mm/yr for that of Cap de Creus.

Our results indicate, for both populations, a significant negative linear correlation between the 

annual diameter growth rate and colony age (Pearson’s correlation coefficient: P = 0.001). For 

both populations, the relationship between colony age and colony basal diameter is well fitted by 

a power curve (AIC test; Fig. 2). Such findings highlight a decrease of the diameter growth rate 

during the life-span of the colony.

Implications of this Analysis

The difference between the two age structures indicates that the population of Portofino is recovering 

from previous exploitation occurred until the 1970s (Bavestrello et al. 2009). The population of 

Cap de Creus, still object of both legal and illegal harvesting (Tsounis et al. 2007), shows a shorter 

life span by about 25%, and a population structure dominated by young colonies as confirmed by 

recent data (Tsounis et al. 2006; Vielmini et al., in preparation).

The colony growth rates found are similar to those measured in other populations (Garrabou 

and Harmelin 2002; Torrents 2007), suggesting a stable growth pattern within and between the 

investigated areas. It is worth noting that colony diameter growth could be strongly affected by 

habitat features (Torrents 2007).

Fig. 1. Thin section (50 μm thick) of the axial 

skeleton of a colony of Corallium rubrum.

180
The decrease in diameter growth rate with age suggests that diameter may not be the most reliable 

descriptor of colony growth. Other descriptors, such as weight or branching pattern, should be 

taken into account to assess Mediterranean red coral.

Nowadays, the colony minimum size limit for commercial harvest of C. rubrum in Cap de Creus 

is fixed above 7 mm; in the light of our findings, such limit could represent a conservative measure 

to allow adult colonies to reproduce about 25 times during their life-span. Despite this, it is worth 

to note that colonies bigger than 7 mm in diameter are very scarce in Cap the Creus (only 4 on 

163 colonies collected on whole, and not considered in population age determination because of 

morphological limitation). 

Further, a sustainable fishery management plan should maximize the yield by harvesting colonies 

at their most productive age (Grigg 1976). Management should ensure an adequate number of 

sexually mature individuals (Grigg 1976): While 7 mm base diameter is above the age of sexual 

maturity, it does not allow for an efficient fishery with maximum sustainable yield, and according 

to recent studies it may not be adequate to ensure the survival of the populations (Santangelo et al. 

2007; Tsounis et al. 2010).

Data on age structure, merged in static life-history tables, will be the base to set out demographic 

models that will allow to project population trends over time. This approach, applied to populations 

with different life-histories, will allow to compare their demographic structures and trends and to 

set out conservation and management strategies suitable for specific local populations.

Fig. 2. Relationship between colony basal diameter (mm) and age in the two Mediterranean 

red coral populations (Portofino: blue spots, N = 75; Cap de Creus: red spot N = 36).

181
Acknowledgements

This study was funded by the Common research project between the University of Pisa, Department 

of Biology (Italy) and CSIC-ICM (Spain): “Population dynamic of Corallium rubrum.” The authors 

are grateful to Valentina Cappanera and Toni Garcia for their invaluable help during field work.

References

Abbiati, M., Santangelo, G., Novelli, S. 1993. Genetic variation within and between two Tyrrhenian 

populations of the Mediterranean Alcyonarian Corallium rubrum. Mar. Ecol. Prog. Ser. 95: 

245-250.

Bavestrello, G., Cerrano, C., Cattaneo-Vietti, R. 2009. Biological interactions affecting the growth 

rates of red coral (Corallium rubrum) colonies. Proceedings of the 1st Symposium on the 

Coralligenous and other calcareous bio-concretions of the Mediterranean. Tabarka, 15-16 

January 2009.

Connell, J.H. 1997. Disturbance and recovery of coral assemblages. Coral Reefs 16(5): 101-113.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: insights into conservation and management 

needs. Journal of Animal Ecology 71: 966-978.

Grigg, R.W. 1976. Fisheries management of precious and stony corals in Hawaii. UNIHISEAGRANT-TR-77-03.

Knittweis, L., Jompa, J., Richter, C., Wolff, M. 2009. Population dynamics of the mushroom coral 

Heliofungia actiniformis in the Spermonde Archipelago, South Sulawesi, Indonesia. Coral 

Reefs 28: 793-804.

Marschal, C., Garrabou, J., Harmelin, J.G., Pichon, M. 2004. A new method for measuring growth 

and age in the precious red coral Corallium rubrum (L.). Coral Reefs 23: 423-432.

Santangelo, G., Bramanti, L., Vielmini, I. 2009. What we have learned about red coral and what 

we have to learn for its rational management. In: Bruckner, A.W., Roberts, G.G. (Eds.) 

Proceedings of the first international workshop on Corallium Science, Management and 

Trade. NOAA Technical Memorandum NMFS-OPR-43 and CRCP-8 Silver Spring, MD.

Torrents, O. 2007. Biologie des populations du corail rouge Corallium rubrum (L. 1758) de 

Méditerranée nord-occidentale. PhD Thesis. Université de la Méditerranée Aix Marseille II.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2006. Population structure of an exploited benthic 

cnidarian: the case study of red coral (Corallium rubrum L.). Marine Biology 149: 1059-

1070.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W.E. 2007. Red Coral Fishery at the Costa Brava (NW 

Mediterranean): Case Study of an Overharvested Precious Coral Ecosystems DOI: 10.1007/

s10021-007-9072-5.

Tsounis, G., Rossi, S., Grigg, R.W., Santangelo, G., Bramanti, L., Gili, J.M. 2010. The Exploitation 

and Conservation of Precious Corals. Oceanography and Marine Biology: An Annual 

Review, Volume 48 (in press).

182
Section IV

Management

183
POPULATION DYNAMICS AND LIFE HISTORY TRAITS 

OF CORALLIUM RUBRUM POPULATIONS: 

IMPLICATIONS FOR SUSTAINABLE MANAGEMENT AND TRADE

Andrew W. Bruckner1,2

1

NOAA Fisheries, Office of Habitat Conservation, Silver Spring, MD USA

2

Khaled bin Sultan Living Oceans Foundation. Landover, MD USA

Abstract

Population demographics, especially density, abundance, size structure, and morphology (branching 

pattern), as well as certain life history traits (e.g. growth rates, reproductive strategy, and longevity), 

must be considered when developing fisheries management strategies for precious corals in the 

family Coralliidae, as these provide indicators of the status of these species and their vulnerability 

to harvest. All species of Coralliidae are sessile, modular (colonial) gorgonians characterized 

by slow-growth, low rates of natural mortality, and a long lifespan. They typically occur at low 

densities, but are capable of forming large, highly branched, tree-like structures in absence of 

disturbance. Most representatives in this family broadcast gametes into the water column on an 

annual cycle, relying on external fertilization and an extended planktonic development phase, 

while the endemic Mediterranean species, C. rubrum, is a brooder that releases well-developed, 

competent planula. This trait has allowed C. rubum to form dense assemblages in shallow water, 

where populations are dominated by small (< 5 cm tall), unbranched colonies. These populations 

exhibit high rates of early mortality, especially in environments frequently disturbed by natural 

stressors (e.g. temperature anomalies, bioerosion) and heavy collection pressure. However, this is 

not the normal situation: C. rubrum colonies are capable of achieving a much larger size (30-50 

cm height) and more extensive ramifications, as reported historically, in deep water populations, 

and in shallow water when protected from fishing. The brooding life history strategy may have 

allowed the persistence of C. rubrum in marginal environments and when subjected to heavy 

fishing pressure, because planula display lower rates of mortality and higher settlement success 

than related broadcast spawning species. Nevertheless, these populations are highly susceptible 

to extirpation due to the selective removal of the largest colonies by SCUBA divers, followed by 

collection of progressively smaller colonies; these smaller colonies produce exponentially fewer 

larvae per reproductive event than large colonies and they reproduce only a limited number of 

times before being removed from the population. Furthermore, C. rubrum exhibits a single annual 

discrete reproductive event unlike other corals that are brooders, and local sources of planula 

are needed to maintain populations due to limited potential for long-distance dispersal. Deep 

populations may be even more vulnerable to extirpation, since they occur at much lower densities; 

they may experience allee effect (depensation) if harvest further reduces colony density below 

levels needed for successful fertilization. An increase in the minimum size of legal harvest (to at 

least 10 mm diameter and 10 cm height) is needed to rebuild overexploited stocks of C. rubrum and 

provide insurance against catastrophic disturbances. This measure would maximize reproductive 

output by increasing colony size, allowing colonies to develop third order branching patterns, and 

increasing the number of times a colony reproduces before removal by collectors. Harvest should 

be limited in deeper areas (> 50 m) where densities are lower until population dynamics and life 

history traits from the deep populations are better understood. Other management approaches, such 

as an increase in the number and size of no-take marine protected areas, are needed to maintain 

184
high reproductive output and compensate for losses due to fishing pressure and other stressors 

outside of protected areas.

Introduction

Habitat forming corals are modular organisms made up of multiple, identical interconnected 

units (polyps) that form massive, plating, encrusting, and branching colonies. These corals are 

permanently attached to the bottom with exception of a short dispersal period during their larval 

phase. Like unitary animals, corals can be classified as r- or k-selected depending on their life 

history strategy, although some species exhibit a continuum across these traits. Most k-selected 

corals form large, long-lived colonies that expend considerable energy in growth and maintenance, 

and less in reproduction. These include some of the dominant framework corals found on shallow 

reefs (e.g. Montastraea annularis), many of which are hermaphroditic (Szmant 1991). K-selected 

corals broadcast millions of gametes into the water column during a single discrete event, once per 

year, which are fertilized externally. These larvae spend an extended period in the water column 

during their development phase, and have a high potential for long-distance dispersal, but they 

tend to exhibit very high rates of larval mortality and low settlement success. K-selected corals 

tend to live for decades to centuries, often sustaining partial mortality that increases over time, but 

colonies continue to grow indefinitely, and the likelihood of total colony mortality declines with 

size (and age) (Bak and Engel 1979; Bak and Meesters 1998). These corals must also achieve a 

certain minimum size before first reproduction, but individual colonies can reproduce indefinitely, 

as long as they remain above that minimum size. 

In contrast, r-selected species are often considered “weedy species” that begin reproducing at a 

smaller size. These corals invest more energy in reproduction and less in growth, but they tend 

to have relatively short life spans. They are often the first species to colonize an area after a 

disturbance because they have naturally high production rates and great inter-annual variation in 

recruitment driven by stochastic environmental factors (Musik 1999). R-selected species include 

most brooders; these species have separate male and female colonies, where fertilization is internal, 

and females release well-developed planula. They produce fewer gametes per reproductive event 

as compared to broadcast spawners, but larvae have higher rates of survival and colonies often 

reproduce monthly on a lunar cycle to overcome shortfalls in larval release. Larvae are competent 

shortly after release and settle within close proximity to the parent, which may also enhance their 

success by colonizing habitats where their parents were most successful. 

Modular Organisms

In addition to selected advantages associated with the different modes of reproduction, there 

are numerous advantages to a modular life history, such as the ability to survive with partial 

mortality. However, these organisms require very specific conditions for successful settlement 

(e.g hard substrate, low rates of sedimentation and algal biomass, and a narrow temperature 

range); once they settle and begin growing, they are generally unable to relocate if the conditions 

become unfavorable. They do exhibit the potential to adapt to their environment through various 

mechanisms such as modifications in growth form. For instance, some species of corals that 

contain zooxanthellae (symbiotic algae) form hemispherical colonies in shallow water, becoming 

more plate-like in deeper environments to maximize their surface area for optimal light capture. 

Branching corals may form short, thickened, robust branches in shallow areas with high wave 

185
exposure and longer, slender branches in calm protected environments. Branching species also 

often dominate large areas and individual colonies attain large dimensions (Gilmore and Hall 

1976). Large sizes increases resistance to all but the most devastating physical disturbances and 

implies greater longevity, while small size implies either a cessation in growth, slow growth, or 

high colony mortality (Szmant 1986).

Modular organisms also exhibit highly complex growth patterns that can mask the age–size 

relationships that are characteristic of unitary organisms such as fishes (Hughes and Jackson 1980; 

Hughes 1984, 1990; Hughes and Tanner 2000). For instance, predation, disease, and other chronic 

disturbances that cause partial mortality can result in colonies of an identical age with dramatically 

different sizes or branching patterns. The combination of such variability in partial mortality and 

annual variation in recruitment makes it unlikely that a coral population ever achieves a stable 

age distribution, unless they are driven to an altered population structure dominated by small 

individuals that never reach their full potential for growth such as that observed in certain shallow 

C. rubrum populations (e.g. Calafuria, Italy; Santangelo et al. 2004). Because they are permanently 

attached to the substrate, they are also more susceptible to total colony mortality associated with 

human activities, such as collection, than motile fisheries species. 

Reproduction and Recruitment

While short-term population dynamics of large, massive, or fast-growing branching species 

is influenced mostly by rates of recruitment, their ability to colonize new areas or areas that 

were catastrophically disturbed, and their ability to persist depends on the degree of success of 

reproduction (Szmant 1986). Reproductive strategies vary across genera and families of corals, with 

some species reproducing sexually through brooding and others via broadcast spawning, and some 

also capable of asexual propagation through fragmentation, polyp bailout, and other mechanisms. 

A coral’s reproductive strategy is not necessarily correlated with polyp size or morphology, but 

reproductive differences may imply important differences in evolutionary selective pressures 

(Szmant 1986). 

The family Coralliidae is one example of gonochoric corals that have a well defined seasonal 

pattern of sexual reproduction, which occurs only once per year (Santangelo et al. 2003; Torrents 

et al. 2005; Tsounis 2005; Tsounis et al. 2006b). Within the family, most species are thought to be 

broadcast spawners, while C. rubrum is the only representative known to be a brooder (Grigg 1993). 

This unique trait has major implications for the survival of C. rubrum when exposed to catastrophic 

acute disturbances (e.g. unusual temperature anomalies) and pressure associated with harvesting. 

First, the planula released by C. rubrum have a much shorter pelagic stage, which allows them to 

bypass high rates of mortality associated with planktonic development. They also exhibit higher 

recruitment rates, at least in shallow water, because they tend to settle in locations where their 

parents were successful. Nevertheless, there are also several factors that limit reproductive output in 

C. rubrum. Most corals that brood larvae are smaller than broadcast spawners; they reproduce on a 

lunar cycle, with many reproductive cycles per year, to compensate for the lower larval production 

per gametogenic cycle as compared to broadcast spawners that release millions of gametes (von 

Moorsel 1983; Szmant 1986; Richmond and Hunter 1990). C. rubrum only reproduces once per 

year, which results in much lower annual levels of larval production. Populations of brooders also 

tend to have shorter dispersal phases, which results in a high degree of genetic separation on a 

186
short spatial scale, and the maintenance and survival of these populations requires local sources of 

healthy reproductive populations to replace colonies that die or are collected. 

Brooders are also more sensitive to changes in density, because the chance for successful 

fertilization is much lower. Colonies that are broadcast spawners synchronously release a large 

a pool of gametes that have a high likelihood of direct contact and fertilization and an obligatory 

development period before they become competent to settle (Willis 1990). C. rubrum is more 

similar to a pine tree that releases pollen. The pollen is ready immediately and becomes less viable 

with time. It requires a chance encounter with a female colony for successful fertilization, which 

will occur only if colonies are located within close proximity or water currents are suitable to 

transport sperm to female colonies. The magnitude of the dispersal for a brooder like C. rubrum, in 

terms of number of larvae dispersing and distance travelled, is much less than larvae of broadcast 

spawning corals (Willis 1990) 

Size-dependent Reproduction 

One of the most critical biological aspects for a sessile (attached) organisms that form complex, 

branching colonies and have long life-spans is colony size, as 1) reproductive output continues to 

increase with size and age; 2) growth to a larger size conveys a greater resistance to disturbance; and 

3) it implies longevity. This is partially due to the modular nature of C. rubrum, where each polyp 

is interconnected with other polyps by a series of canals that are responsible for the translocation of 

water and nutrients, allowing all polyps to function together as a single, larger unit. These colonies 

must reach a certain size before they become sexually mature, but once that size is achieved they 

can reproduce indefinitely. In branching gorgonians, reproductive output increases exponentially 

with size and the largest colonies in the population are thought to produce 90% or more of the 

offspring (Babcock 1991; Coma and Gili 1995; Beiring and Lasker 2000; Santangelo et al. 2003; 

Torrents et al. 2005; Bruckner 2009). 

Although C. rubrum is a brooder, it exhibits many traits in deep water that are similar to other 

gorgonians that are broadcast spawners (including other species of Corallium), presumably because 

fishing pressure and large scale environmental disturbances are less common which allows these 

corals to achieve much larger sizes. Most species in the family Coralliidae are in the low-productivity 

range as colonies are relatively slow-growing and long-lived, with population turnover estimated 

to occur once every 15-25 years (Grigg 1976) or longer, based on more recent estimates of age 

(Marschal et al. 2004). Brooding and early reproduction may be an adaptation that allows the coral 

to occupy marginal habitats in shallow water, because the colonies can never achieve a large size 

before dying (Santangelo 2003), which results in a much more rapid population turnover. These 

populations usually display high rates of recruitment (0-32 recruits/m2

 per yr from 1995 to 1999 in 

Spain (Linares et al. 2000), and 0-12.5 recruits/m2 per yr from 1979 to 2000 in France (Garrabou et 

al. 2001)), although recruitment occurs in pulses with no successful settlement in some years, up 

to 95% of the larvae die before settling (Tsounis 2005), and 66-70% of the new recruits die over 

the first four years of life (Bramanti et al. 2005). Small colonies that dominate shallow areas of 

the Mediterranean today are also more vulnerable to total colony mortality, and their contribution 

to future generations is much less than larger colonies because they contain substantially fewer 

reproductive modules (polyps). 

187
Significant differences in the size of Corallium rubrum colonies have been reported between 

locations and depths, and most areas affected by coral fishing and recreational diving have shown a 

general trend of declining size structure over the last four decades. For instance, colonies collected 

in the 1950s and 1960s were frequently up to 50 cm height and 2 cm diameter. A 1962 collection 

off Costa Brava, Spain consisted of colonies with basal diameter of 10-45 mm (mean = 16 mm) 

and height of 10-50 cm (mean = 11.5 cm) with the largest corals estimated at 50-80 years in age 

(Garrabou and Harmelin 2002). Colonies sampled off Costa Brava were still substantially larger in 

1986 than that observed today with 85% of the colonies having a basal diameter > 5 mm (GarcíaRodríguez and Massò 1986). Shallow populations from Costa Brava (10-25 m depth) are now 

skewed towards small corals (mean height = 3.1 cm, max = 20 cm; mean diameter = 4-5 mm), 

most of which are young (< 8 years old) and only 35% have basal diameters > 5 mm. Colony 

height increases with depth to 4 cm height at 25-50 m and 6 cm at 50-90 m; the largest colonies 

(13-16 cm height) are found in non-harvested areas below 50 m depth (Rossi et al. 2008). Colonies 

in an adjacent protected area (Medas Island, Spain) closed to fishing for 15 years were 20-50% 

larger (mean = 4 cm height, max = 20 cm) than colonies at similar depths in fished areas off Costa 

Brava. Populations examined in 1998 in fished areas off France (Riou archipelago) were larger 

(mean height = 6 cm; range = 4-13 cm) than that reported for Spain (although small colonies were 

not examined in this study), but these colonies were still only about half the size of a similar habitat 

in France protected from fishing since 1983 (mean = 11.8 cm, range = 8-17 cm). 

Differences in total reproductive output are directly related to colony size structure and the degree 

of branching. Corallium rubrum are gonochoric brooders that were reported to first produce 

gonads at 2-10 years age (minimum 2-3 cm height, 1.8-3.6 mm diameter), with colonies achieving 

100% fertility at 4-6 cm height. The total number of polyps increased exponentially with colony 

size from about 25-90 polyps in a small (2-3 cm height) single branched colony to over 4000 

in a 12 cm tall colony with over 35 branches, and 15,000 for colonies 30 cm in height. Even in 

the Ligurian Sea, where colonies never exceed 10 cm in height and most had few branches, the 

number of planula increased 7 fold from the smallest reproductively mature colonies (1.4-2.3 mm 

diameter, 24 planula) to the largest (> 4.6 mm diameter, 158 planula) (Santangelo et al. 1993). 

While small colonies may be the most important component in terms of reproductive contribution 

in some shallow populations today (Santangelo et al. 2007), this is only because the small colonies 

are numerically dominant, versus very few larger colonies. Nevertheless, when compared to their 

potential to achieve larger sizes the contribution of these small colonies to future generations 

would be much less if a population exhibits a more normal size frequency that is not skewed 

towards juvenile colonies. They are also less likely to sustain the synergistic impacts of fishing, 

natural stressors and climate change, and degraded areas have a delayed potential for recovery due 

to limited potential for long distance dispersal.

Relationships between Size and Density

Abundance and biomass are usually important factors in assessing population status and trends 

of fishery species. However, for sessile colonial species, size and relationships with density are 

more important measures, as size is a major determinant of first reproduction, reproductive output, 

and colony survival, and density is related to the likelihood of fertilization. In the case of C. 

rubrum, smaller colonies tend to occur in large aggregates because they settle relatively close to 

their parent; these smaller-sized colonies lack a branching morphology and exhibit higher rates of 

natural mortality. As they increase in size, they require more space, but survival also increases with 

188
size. If they are able to reach a large size, populations will ultimately become less dense, with a 

lower overall abundance throughout a bed, but populations will potentially have a higher number 

of polyps overall. For instance, a single large colony 300-500 mm in height can contain more 

polyps (up to 10,000 or more) than 100 or more small colonies that are 30-50 mm height (approx. 

100 polyps), like those found today in the Mediterranean. The increase in polyps is partially related 

to size (larger colonies have more polyps overall), but large colonies lack polyps at the basal 

portions and polyps may be more dense but smaller at the branch tips. Most importantly, polyp 

number also increases in proportion to the degree of branching, which is much more extensive in 

large colonies. For these reasons, size/age structures give a better indication of population decline, 

as the fishery is size-selective (largest colonies are removed first) and a decline in the number of 

large colonies may result in an increase in small colonies as more space becomes available to 

support settlement of larvae. 

Data on the population structure are also more useful in identifying a changing proportion of 

mature/immature colonies, which is more functional as a basis for management decisions that 

need to ensure minimum recruitment, especially for sessile animals that require a certain density 

to ensure fertilization success (Bruckner 2009; Tsounis et al. in press). Abundance data cannot 

be directly compared among populations or within individual beds over time, due to their patchy 

occurrence within these beds, differences in the available habitat within an area occupied by this 

coral, and highly variable features such as crevices and outcrops that are not uniformly distributed 

throughout the habitat. Also, numbers of colonies per unit area are unlikely to provide an indication 

of the population status or trends due to their life history traits, which include: 1) a sessile (attached) 

existence; 2) density-dependent recruitment within localized areas that result in dense populations 

of small, unbranched colonies; 3) the ability to form of large, highly branched, three-dimensional 

colonies that are long-lived and will reproduce annually for the duration of their lifespan, in the 

absence of physical disturbance (collection).

In Spain, depth was found to be only weakly correlated to population abundance while there was 

a stronger correlation with size, with larger colonies found in deeper water where fishing pressure 

was lower (Tsounis et al. 2006b). The weak relationship between density and depth may be due 

to small scale variation in environmental and habitat attributes. In very shallow areas (18-25 m) 

colonies occur in dense aggregations in cracks and crevices, under overhangs and on the walls 

and ceiling of small caves, where they form very small patches. In contrast at depths below 35 m, 

colonies occur in the open at lower densities but they cover a larger area and are larger in size. This 

trend was apparent several decades ago, where densities in Palma de Mallorca were reported at 55 

colonies per square meter at 40 m depth and 20 colonies/m2 at 60 m depth along the Costa Brava, 

while 90-100 colonies/m2

 were observed in Corsica (FAO 1984). These abundances are much 

lower than reported in the Ligurian Sea (Santangelo et al. 1993), but those populations may consist 

of 2/3 or more juvenile colonies. One of the limitations of these data is related to differences in 

sampling regimes; the scraping technique detects all corals, small and large, while other methods 

under count recruits. Furthermore, recruitment in the densest populations (Ligurian Sea) appears 

to be an order of magnitude higher than other areas and juvenile mortality is lower (Bramanti et al. 

2006), but some of these dense shallow populations are much more susceptible to mass mortality 

events and partial mortality from sponges and other factors (Garrabou et al. 2001) and population 

turnover is very rapid, with few colonies ever achieving a minimum diameter of 7 mm. 

189
Recommendations for Improved Management

• Increase minimum size of harvest to 10 mm basal diameter and 10 cm height: With 

exception of new measures adopted in Sardinia, most other locations in the Mediterranean that 

use size limits in their national coral management plans establish a minimum size of harvest for 

C. rubrum at 7 mm basal diameter. In one of the earliest demographic study of C. rubrum, GarcíaRodríguez and Massò (1986) documented the harvest of colonies that were 5-14 years in age 

which they concluded were well below maximum sustainable yield (MSY). Based on recent age 

studies that indicate Corallium colonies are 2-4 times older and growth rates are 2.6-4.5 times 

lower than previously thought (Marschal et al. 2004; Roark et al. 2006). Tsounis et al. (2007) 

estimated MSY of 98 years, whereas the current practice of harvesting colonies once they achieve 

a 7 mm basal diameter (11 year old colonies) results in only 6% of the potential yield. This increase 

in size would allow colonies time to develop branches, reach 100% sexual maturity, and reproduce 

multiple times before collected.

• Reduce collection at depths above 50 m depth: These populations are reported to be 

overexploited throughout the Mediterranean. Collection needs to be limited until colonies 

reestablish a larger size structure. Once these areas recover, collection should be limited to small 

portions of the total habitat and large-size colonies only (e.g. above 10 cm height).

• Create networks of protected areas (MPAs): MPAs should encompass all depths/habitats 

occupied by Corallium and portions of each bank where Corallium is collected. Existing no take 

areas (protected areas) for C. rubrum are relatively small and it is unclear whether they are sited 

such that they would replenish fished areas, given the high degree of genetic structure and evidence 

that populations are self seeding (Costantini et al. 2007). 

• Rotational harvest: It is unclear how effective this measure would be unless rotational 

periods exceed 30-40 years to allow full recovery of populations as colonies in areas protected 

from fishing for 15-30 years are still less than half the reported historic size (Caddy 1993; Francour 

et al. 2001; Tsounis et al. 2006a). 

• Limit harvesting in deep water: Proposals to limit collection to 80-130 m (mixed gas diving) 

appear to be short sighted at this time because we know too little about these populations. This 

move to deeper water may be a reflection of the depleted nature of the resource – all acceptable 

(large) colonies have been removed from shallow water and collectors are now targeting areas that 

have not been fished for 20 years or more (e.g. since elimination of the dredge). Harvest in these 

areas should be restricted until population structure is quantified and more data as both growth 

rates and recruitment may be much different in deep water due to environmental variations.

• Establish monitoring plans: Baseline studies on population dynamics must be completed 

before a new area is open to fishing. Conservative estimates of landings from these areas should 

be adopted based on the size of the bank that contains Corallium, its abundance and size structure. 

Collection areas should then be monitored on an annual basis to determine collection impacts and 

to modify measures as needed to prevent a shift in size structure as that observed in shallow water. 

• Enforcement: Better monitoring of landings that include data on numbers of colonies, sizes 

(height and diameter), and numbers of branches, along with total weight, should be undertaken 

190
at ports and marinas, instead of voluntary reporting. Divers should be required to submit daily 

log books, including site locations, and strict penalties should be adopted for offenses. Once a 

minimum size is established, there should not be a variance allowed (e.g. 20% as included in 

the Sardinia plan), as this diminishes the value of a minimum size and it is an indication that 

inadequate numbers of larger colonies exist in an area if collectors are removing smaller corals.

Conclusions

For a modular organism that characteristically forms highly complex, branched colonies, and can 

reach sizes of 30-50 cm, a mean height of 3-5 cm, as reported from most shallow areas (< 50 m) 

in the Mediterranean, is equivalent to a loss of 80-90% of the reproductive modules (polyps) of 

individual colonies, due to the absence of 2nd, 3rd and 4th order branches. These small colonies can 

become sexually mature at a young age (2-3 cm height), although they don’t achieve 100% fertility 

until about 6 cm height and 10-20 years in age or older. The spawning potential in C. rubrum (and 

other gorgonians) increases exponentially with size, with larger arborescent colonies producing up 

to 90% of the recruits. Given settlement rates of no more than 5% of the total larval production, 

and continued removal of colonies by fisheries after they have reproduced no more than one or two 

times, typical shallow population today may produce 80-90% fewer recruits than in the 1960s, and 

about half of that produced by populations that have been protected from fishing for 15-20 years 

and contain colonies twice as large.

Using only colony abundance and density as an indication of population size and viability can be 

misleading, as dense populations are limited to only C. rubrum and these tend to be dominated by 

small, reproductively immature colonies. For colonial organisms, change in population structure 

(size frequency distribution) is a more suitable measure of decline than changes in the absolute 

numbers of colonies. Commercial extraction primarily eliminates the largest corals, followed 

by smaller colonies over time, but it is the largest, oldest colonies that contribute most to the 

replenishment of the population. Furthermore, shifts in the size structure of populations due to 

fishing pressure can be directly compared, while density and abundance cannot. This is because 

these measures differ depending on how they are assessed (colony density measured over the entire 

suitable habitat is much less than the density of small patches occupied by the coral within this 

habitat), and the life stage of the population. In fact, a less-dense population is likely to represent 

an older, more stable and viable population as open substrates of suitable habitat can support 

high numbers of recruits, but these exhibit size-related survival that increases as the colonies get 

larger. Thus, populations with a high abundance and density, such as those seen in Mediterranean 

shallow water, are an indication of frequent continuing perturbations responsible for rapid turnover 

of populations and a persistent state of early-stage recovery. This is similar to observations of 

other corals that brood their larvae; however, most other corals that are brooders are considered 

early colonizing, “weedy” species, while C. rubrum is a long-lived species that may be attempting 

to adapt to increasing localized (direct human impacts) and global stressors (climate change). 

These types of populations are much less resilient to other stressors and are more likely to exhibit 

localized extirpations when compounded by fishing pressure than populations that contain a mix of 

small (10-50 mm tall), medium (60-140 mm), and large (150-500 mm) colonies, like that formerly 

observed in the Mediterranean and still present in some deep-water areas that have not been 

targeted by fisheries for several decades.

191
Throughout most of the Mediterranean, C. rubrum colonies in shallow water now form small, 

high density aggregations within much larger areas of suitable habitat, while the corals become 

less dense and larger in size in deeper areas with lower fishing pressure. These dense shallow 

aggregations are dominated by recruits and small adult colonies, generally no more than 3-5 cm 

in height with few colonies over 10 cm in height, whereas historically colonies were 20-50 cm in 

height. Because of the sessile nature of the species, further reductions in density associated with 

the selective removal of the largest colonies in deeper coral beds may alter reproductive potential 

due to allele effects. Also, the smaller organisms in shallow water may experience catastrophic 

mortalities when fishing pressure is compounded by other stressors such as climate change. 

References

Babcock, R.C. 1991. Comparative demography of three species of scleractinian corals using age and 

size-dependent classification. Ecol 61: 225-244.

Bak, R.P.M., Engel, M.S. 1979. Distribution, abundance and survival of juvenile hermatypic corals 

(Scleractinia) and the importance of life history strategies in the parent coral community. 

Mar Biol 54: 341-352.

Bak, R.P.M., Meesters, E.H. 1998. Coral population structure: the hidden information of colony sizefrequency distributions. Mar Ecol Prog Ser 162: 301-306.

Beiring, E.A.,Lasker, H.R. 2000. Egg production by colonies of a gorgonian coral. Mar Ecol Prog 

Ser 196: 169-177.

Bramanti, L., Magagnini, G., Maio, L.D., Santangelo, G. 2005. Recruitment, early survival and 

growth of the Mediterranean red coral Corallium rubrum (L 1758), a 4-year study. J Exp 

Mar Biol Ecol 314: 69-78.

Coma, R., Gili, J.M. 1995. Reproduction and cycle of gonadal development in the Mediterranean 

gorgonian Paramuricea clavata. Mar. Ecol. Prog. Ser. Vol 117: 173-183.

FAO. 1984. Technical consultation on red coral resources in the Western Mediterranean. FAO 

Fisheries Report 306, Roma, 142 p.

García-Rodríguez, M., Massó, C. 1986a. Estudio biométrico de poblaciones de coral rojo (Corallium 

rubrum L.) del litoral de Gerona (NE de España). Bol Inst Esp Oceanogr 3: 61-64.

Garrabou, J., Perez, T., Sartoretto, S., Harmelin, G. 2001. Mass Mortality Event in Red Coral 

Corallium rubrum Populations in the Provence Region (France, NW Mediterranean). Mar 

Ecol Prog Ser 17: 263-272.

Garrabou, J., Harmelin, J.G. 2002. A 20-year study on life-history traits of a harvested long-lived 

temperate coral in the NW Mediterranean: Insights into conservation and management 

needs. Journal of Animal Ecology 71: 966-978.

Gilmore, M.D., Hall, B.R. 1976. Life history, growth, habitats, and constructional roles of Acropora 

cervicornis in the patch reef environment. J Sediment Petrol 46: 519-522.

Grigg, R.W. 1993. Precious coral fisheries of Hawaii and the U.S. Pacific Islands - Fisheries of 

Hawaii and U.S. - Associated Pacific Islands. Marine Fisheries Review 55: 50-60.

Hughes, T.P. 1984. Population dynamics based on individual size rather than age: a general model 

with a reef coral example. Amer Naturalist 123: 778-795.

192
Hughes, T.P. 1990. Recruitment limitation, mortality, and population regulation in open systems: a 

case study. Ecol 71: 12-20.

Hughes, T.P. 1994. Catastrophes, phase shifts, and large scale degradation of a Caribbean reef. 

Science 265: 1547-1551.

Hughes, T.P., Tanner, J.E. 2000. Recruitment failure, life histories, and long term decline of Caribbean. 

Musick, J.A. 1999. Ecology and conservation of long-lived marine animals. In: Musick, J.A. (Ed.) 

Life in the slow lane: ecology and conservation of long-lived marine animals. Am. Fish Soc. 

Symp. 23 Bethesda MD USA, 1-10.

Richmond, R.H., Hunter, C.L. 1990. Reproduction and recruitment of corals: comparisons across the 

Caribbean, the Tropical Pacific, and the Red Sea. 60: 185-203.

Rossi, S., Tsounis, G., Padrón, T., Orejas, C., Gili. J.M., Bramanti, L., Teixidor, N., Gutt, J. 2008. 

Survey of deep-dwelling red coral (Corallium rubrum) populations at Cap de Creus (NW 

Mediterranean). Mar Biol 154: 533-545.

Santangelo, G., Abbiati, M., Giannini, F., Cicogna, F. 1993. Red coral fishing trends in the western 

Mediterranean Sea during the period 1981-1991. Sci Mar 57: 139-143.

Santangelo, G., Carlietti, E., Maggi E., Bramanti, L. 2003. Reproduction and population sexual 

structure of the overexploited Mediterranean red coral Corallium rubrum. Marine Ecology 

Progress Series 248: 99-108.

Santangelo, G., Maggi, E., Bramanti, L., Bongiorni, L. 2004. Demography of the over-exploited 

Mediterranean red coral (Corallium rubrum L. 1758). Scientia Marina 68: 199-204.

Santangelo, G., Bramanti, L., Lannelli, M. 2007. Population dynamics and conservation biology of 

the overexploited Mediterranean red coral. J. Theor. Biol. 244: 416-423.

Szmant, A.M. 1986. Reproductive ecology of Caribbean reef corals. Coral reefs 5: 43-54.

Szmant, A.M. 1991. Sexual reproduction by the Caribbean reef corals Montastraea annularis and M. 

cavernosa. Mar Ecol Prog Ser 74: 13-25.

Torrents, O., Garrabou, J., Marschal, C., Harmelin, J.G. 2005. Age and size at first reproduction in 

the commercially exploited red coral Corallium rubrum (L.) in the Marseilles area (France, 

NW Mediterranean. Biological Conservation 121: 391-397.

Tsounis, G. 2005. Demography, reproductive biology and trophic ecology of red coral (Corallium 

rubrum L.) at the Costa Brava (NW Mediterranean): ecological data as a tool for management. 

Reports of Polar and Marine Science. 512. Alfred Wegener Institute for Polar and Marine 

Research, Bremerhaven.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2006a. Population structure of an exploited benthic 

cnidarian: the case study of red coral (Corallium rubrum L.). Mar. Biol. 149: 1059-1070.

Tsounis, G., Rossi, S., Gili, J.M., Arntz, W. 2006b. Effects of spatial variability and colony size 

on the reproductive output and gonadal development cycle of the Mediterranean red coral 

(Corallium rubrum L.). Mar. Biol. 148: 513-527.

Van Moorsel, G.W.M.N. 1983. Reproductive strategies in two closely related stony corals (Agaricia, 

Scleractinia). Mar Ecol. Prog Ser. 13: 273-283. 

193
Willis, B.L. 1990. Species concepts in extant scleractinian corals: considerations based on reproductive 

biology and genotypic population structures. Systematic Botany 15: 136-149.

194
LESSONS LEARNED FROM NATIONAL TRADE REGULATIONS ESTABLISHED 

FOR THE RED CORAL, CORALLIUM RUBRUM

J. Dietrich

Head CITES Management Authority

The timely restricted (January 1987 – June 1997) strict protection of Corallium rubrum in 

Germany through the listing of the species in Annex 1 of Germany’s Federal Ordinance on Species 

Conservation resulted in a total prohibition of any commercial trade into Germany, both from EU 

and non-EU Member States.

Due to this situation German, customs was confronted with commercial imports of premanufactured and manufactured products made of other Corallium species. The listing revealed 

significant (irresolvable) identification problems on the species level for enforcement officials in 

particular for pre-manufactured products, jewelry, or products made of coral dust.

Wrong identifications lead to wrong declarations of seizures of red coral shipments from countries 

where the species doesn’t occur in the wild.

In the light of a current pending consultation process initiated by the U.S. government for a 

possible CITES listing of the entire family Coralliidae, and based on Germany’s experience with 

its past national protection of Corallium rubrum, the U.S. proposal is not only evaluated but also 

recommendations are made how to enhance conservation and management of red corals on our 

planet by taking other suitable approaches under CITES.

195
CITES NON-DETRIMENT FINDINGS FOR PRECIOUS CORALS

V. Fleming

UK CITES Scientific Authority (Fauna), JNCC, 

Monkstone House, City Road, Peterborough PE1 1JY, UK

The implications of a potential CITES listing for Coralliidae are discussed, especially with respect 

to the requirement under CITES to make a ‘non-detriment finding’ before any export is permitted. 

In this respect, the outcome of the workshop on non-detriment findings, held in Mexico (November 

2008), and the general guidelines and principles recommended by that meeting are presented, 

especially with respect to aquatic invertebrates. This guidance on aquatic invertebrates suggested 

an ongoing cyclical process of: a) assessing risks, b) regulating harvests, c) recording harvests, 

trade, and population responses and, finally, d) reviewing, revising, and refining, if necessary, the 

management measures. Case studies from the workshop, especially those for black and stony corals, 

provide examples of approaches to non-detriment findings for taxa, already listed on CITES, which 

have similar life histories to red corals. Some of the perceived difficulties of any CITES-listing 

for Coralliidae, especially with respect to nomenclature, identification of worked specimens, and 

administrative burdens, are also addressed with respect to experience from CITES-listed taxa such 

as black and stony corals and recent discussion in the Hong Kong workshop (March 2009).

196
REGULATION OF PRECIOUS CORAL FISHERIES IN THE WESTERN PACIFIC AREA

Sadao KOSUGE*, Hideo KIUCHI**,Yasuo TANABE**, Katsuo KAMEI**

*) Laboratory for Research of Organic Jewelry of Institute of Mala-cology of Tokyo, Japan.

 **) Japan Cooperative Society of Commerce and Industry of Precious Coral, Kochi, Japan.

Precious coral of the Mediterranean Sea (Mediterranean red coral) introduced to Japan as one of 

the Buddhist Jewelry about more than 1200 years ago, and this is the first record that the precious 

coral appear in Japan. Buddhist scriptures described as the heaven (paradise of Buddha) decorated 

with the seven treasures, namely Gold, Silver, Precious Coral, Crystal, Lapis Lazuli, Agate, and 

Giant Clam Shell.

In Buddhism, precious coral is referred as the one of the treasures of paradise and is used to 

decorate Statue of Buddha along with other gem stones until present time. This is most evident in 

Lamaism (Tibetan Buddhism), where precious coral used not only to decorate religious statues, but 

also as religious artifacts, such as personal talismans.

According to the stabilization and extension of Japanese Society many objects imported from 

China in Eighth Century, of which generation represented by Emperor Shoumu (701-756). His 

treasures preserved in the National Treasure House named “Hyo-So-In” of To-daiji Temple. These 

treasures were donated to the To-daiji Temple by the Empress Koumyou on June 21st of 756 AD 

which was 49 days after the death of her husband, Emperor Shoumu, as 49 days are an important 

date for the deceased person in Buddhism.

This collection features many relics of ancient Persia, China, and other ancient countries, and these 

examples are no longer exist in their mother (native) countries. The number of collection of this 

treasure house is 9020 items in total. 

Beads and small branches of Mediterranean red coral are included in this collection. The beads 

were used in the decoration of a crown which the Emperor Shoumu wore at the ceremony for 

newly built statue of Buddha in 752 AD. 

On precious coral, these situation continued long time in Japan. For instance, in the TokugawaShogunate (ruled by Samurai, from 1607 to 1867 AD), foreign trade was strictly restricted and 

only permitted with Holland (Netherlands) and China at the Port of Nagasaki (Deshima), Kyushu, 

Japan. 

Import of precious coral from the Mediterranean Sea during these days can traced from the records 

of trade with Holland which were preserved in the national Archives of both countries. Volumes of 

trade of precious coral were not great, and most of the precious coral brought to Japan were used 

as specific gifts to the Shogun and officers of Tokugawa Shogunate. Journals of the successive 

Heads of the Dutch Trade Delegations to Japan, preserved in the Central National Archives of the 

Netherlands in Hague, reveal that the director of each delegation was obligated to travel to Edo 

(now, Tokyo) for an audience with the Shogun.

197
At that time, the trade delegation heads brought many gifts for Shogun, his Prince and other top 

level officials of the Tokugawa Shogunate, and the Mediterranean red coral always included in 

these gifts.

Similar gifts were also presented to the officials at the port of Nagasaki, in which Mediterranean 

red coral were presented to the important Offices in the port. It is said that Mediterranean red 

coral was regarded as one of the most valuable gifts during the Tokugawa Shogunate Period. 

During Tokugawa Shogunate, precious coral was also imported via China through the trade routes 

between the Mediterranean area and China. Although the amount of Mediterranean red coral from 

China was not so big as that from Holland, trade records indicate that precious coral was always 

included among the cargoes of ships from China.

So we, the Japanese, have a special emotion to the Mediterranean red coral, and wish to say our 

special thanks for the people of the Mediterranean area for their long time supply and support.

Despite the long history of the using of precious coral in Japan, precious coral fisheries in Japan 

have a rather short history to compare with the Mediterranean red coral fisheries. Because precious 

coral off the coast of Japan inhabits depths of more than 70 to 100 meters, it was hard to harvest 

them using the primitive fishing methods at that time.

Coral fisheries in Japan did not start until the early 18th century. Also unlike Mediterranean coral, 

coral in Japan did not wash up on beaches after storms or heavy surf, because they lived on 

the deep seabed. On the other hand, the Tosa Clan in Tokugawa Shogunate prohibition against 

gathering coral was not strictly enforced. While the Clan did not wish to see common people 

develop unregulated fisheries and free trade in its Domain, neither did it intend to levy severe 

penalties for contingent coral harvests that accompanied other types of fishing. This is supported 

by the document issued 1838 AD that stated anyone harvesting precious coral was prohibited from 

selling it freely and should offer to the Tosa Clan in stead of selling to the outside. A warning was 

added stating that anyone violating this regulation would be punished.

During the Meiji Restoration the feudal system was abolished along with all Tosa Clan regulations 

and laws. Soon afterward precious coral fisheries began operating along the coast of the Muroto 

region (Tosa Bay) in 1871.

The equipment used at this time was the coral gathering net that Mr. Konojoi Ebisuya had invented. 

Modern equipment still uses the same basic principles Ebisuya used in designing his net. Konojoi 

Ebisuya’s work in opening and developing coral fisheries was recognized when he was given 

an award at the 1897 Fisheries Exhibition. Also white precious coral (shiro) was dedicated in 

his honor as Corallium konojoi by Kishinoue in 1903. At the present, coral fisheries in Japan 

operate in Kochi area (Tosa Bay) by net and southern part of Kyushu by submarine. Both fisheries 

operations are regulated by the provincial government of each area. The details of the regulations 

are presented by the previous report by Hideki Moronuki.

Explanation 1.

Detail of harvest amounts of the past 20 years from Tosa Bay, Kochi, Japan is explained with the 

data in each year including harvest amount, traded price, and number of boats. From these data, 

198
there are no signs of over-harvest or depletion of precious coral resources along the Japanese coast. 

Further, harvested coral by boat are almost all dead ones and its percentage is 85 to 90% of total 

harvest amount. 

Explanation 2.

Displayed actual material of three species of precious coral (C. japonicum, elatius, and konojoi) in 

various phases (alive, 3 phases of dead material, to worm eaten phase), and natural to manufactured 

ones. People can understand that dead material is still useful for manufacturing and rather hard to 

discriminate the differences among these final products of these samples. So they can understand 

that dead coral are still useful and no problem for manufacturing.

Explanation 3.

Recently, the Taiwan Government established very severe regulations of the coral fisheries around 

the Taiwan area, which started from 2008. Issue of fisheries licenses is strictly investigated. Annual 

harvest amounts of each boat are strictly restricted. Each boat must be set up with GPS equipment, 

so each boat can be traced by the government. Fisherman must be present daily reports on his 

fisheries results within 3 days after the fisheries operation. Permit areas for coral fisheries around 

the Taiwan territory are strictly regulated.

Report of harvest amount in Taiwan (2009)(unit is kg):

 January 0.111 February 140.147 March 154.964

 April 346.447 May 348.615 June 478.286

 July 263.352 August 325.004 September 355.692

These reports will be issued every month in the future.

References

Jeng, M.S. 2009. Present condition of the regulation of precious coral. Fisheries in Taiwan. 

International Forum on precious coral, Rept. of first meeting in Hong Kong. 33-35.

Kosuge, S. 1993. History of the precious coral fisheries in Japan. Precious corals and Octocoral 

Research 1: 30-38. 

Kosuge, S. 2009. Buddhist Scripture described on the Heaven. International Forum on Precious 

Coral, Rept. of first Meeting in Hong Kong back cover.

Kosuge, S. 2009. Analytic data on the harvest amount of precious coral in Tosa Bay. Precious Corals 

and Octocoral Research 6: 1-5.

199
THE INCLUSION OF CORALLIUM JAPONICUM, CORALLIUM KONOJOI, CORALLIUM 

ELATIUS, AND CORALLIUM SECUNDUM IN APPENDIX-III OF CITES. 

THE ITALIAN EXPERIENCE OF CORPO FORESTALE DELLO STATO. 

M. Mei, C. Lungo, L. Corbetta, S. Di Domenico

Corpo Forestale dello Stato – Central CITES Unit

Abstract

Until about two decades ago, much of the red coral worked in Italy came from Corallium 

rubrum, characteristic of the entire range of the Mediterranean Sea. Currently, about half of 

the manufactured product comes from the Chinese area. In 2008, four species of red coral from 

People Rep. of China (Corallium japonicum, Corallium konojoi, Corallium elatius, and Corallium 

secundum) were included into Appendix-III of CITES. This was applied by the European Union 

with Reg (EC) 407/09 (entry into Annex C). This inclusion requires specific procedures for the 

import and re-export of raw coral and finish products derived from it, about the enforcement of 

which is responsible in Italy the State Forestry Corps. In fact, for specimens importation of the four 

species considered, either need a Certificate of Origin, or re-export certificate or an export license. 

Sales in Italy and then to the EU does not require specific permissions, but in case of re-export to 

countries outside the EU, it is required that the specimens are accompanied by an export certificate, 

issued by the CITES Service of State Forestry Corps. At this moment, about 50% of raw red coral 

is purchased in Italy from Pacific area/P. Rep. of China/Taiwan and it is often very difficult to 

distinguish a finish product made with Chinese Coral in comparison to another from C. rubrum, as 

in the raw product you can recognize morphological and microscopical differences. 

The species of Corallium and Paracorallium (Bayer and Cairns 2003) genus are commercially 

exploited by man for over 5000 years. Manufacturing activity in red coral in Italy is an important 

source of income for many people, with small firms located in Naples hinterland and, in particular, 

in Torre del Greco. Until about two decades ago, much of the raw material processed in Italy came 

from Corallium rubrum, characteristic of the entire range of the Mediterranean Sea. Currently, 

about 40% of rough/semi-rough product comes from Pacific/Japan/Philippines/ PDR of China/P 

of Taiwan area.

On April, 2, 2008, by Notification n° 27, the CITES Secretariat in Geneva informed Parties that, 

at the request the People’s Democratic Republic of China, four species of red coral – Corallium 

japonicum, Corallium konojoi (synonym of C. konjoi), Corallium elatius, and Corallium secundum 

– have been added in Appendix-III of CITES. This inclusion in the European Union only became 

operational on 22 May 2009 with the publication of Reg (EC) of Commission n° 407, May 14, 

2009, replacing the EC Reg 318/2008, and where these four species of red coral have been included 

in Annex C of modified Reg (EC) Council Regulation n° 338/97. This inclusion required the 

adoption of specific procedures for the import and re-export of raw coral and by-products.

The inclusion in Annex C of Reg (EC) 338/97 requires exact obligations for the State Forestry 

Corps, that is in charge for certification activities and control of specimens of species protected 

by CITES. State Forestry Corps discharge these duties across more than 50 offices spread all 

over the country: CITES Territorial Services and CITES Operations Units for customs controls, 

200
both coordinated by a CITES Central Service established in Rome (Fig. I). For the importation 

of live coral or dead, rough, parts and derivatives “specimens” belonging to one of four species 

considered, if they do not come from People’s Republic of China, they must always be accompanied 

by a “Origin certificate” or re-export certificate issued by the Official CITES Authority of the 

exporting country. When instead such specimens come from PRC, they must be accompanied 

by an export permit issued by the Official CITES Authority of that country. In either cases, when 

the specimens entry into Italy, CITES Operations Units of the Forestry Corps, fill a specific form 

named “Import Notification” on the basis of the documentation accompanying goods at customs 

point. The subsequent sale in the country and EU, based on Article 8 of Reg. (EC) 338/97, does not 

require specific authorization under the CITES legislation. In any case, the seller will provide care 

to the client, about the purpose of resale outside the EU, the information on the CITES document 

of importation.

In case of re-export to countries outside the European Community of specimens with coral 

belonging to four species in question, under provisions of article 5.4 of Reg. (EC) 338/97, the same 

specimens need of a re-export certificate, which is issued by the CITES Service of Forestry Corps. 

This kind of certification is issued, as provided in Article 5 of Reg. (EC) 338/97, only on the basis 

of documentary evidence attesting the correct importation of source material. 

As instead regards the other species of coral belonging to the genus Corallium sp, in particular 

C. rubrum, which at present are not subject to the provisions of CITES and Reg. (EC) 338/97, 

CITES Service of the State Forestry Corps provides for the issuance, at the request of the subject, 

of a so-called “declaration of non-inclusion,” which is placed on the bills of sale on the basis of 

documentary evidence of the non-inclusion in the lists of CITES. For personal and households 

effects, under the CITES legislation is not required any certificate, preserving the possibility of 

stricter domestic measures by the Parties. It was launched a consultation on this matter with some 

states outside the EU, trough Italian Main CITES Management Authority which is constituted 

by the Ministry of Environment and the protection of Territory and Sea: the United States, in 

particular, have confirmed that any kind of document is not required for this type of objects as 

species included in Appendix-III. 

And now, with the next figures (1, 2) are showed provided data on notifications for import 

compiled in 2009 (May – September 2009) by CITES Operations Units (NOC) and re-export 

certificates issued by CITES Territorial Services (SCT) of State Forestry Corps. Against n° 31 

import notifications related to three of the four species considered, compiled by NOC operating in 

the regions of Veneto, Lombardia, Lazio, and Campania, it was issued a total (for the same period) 

of n° 31 re-export certificates, by SCT operating in Vicenza, Rome, and Naples in particular. 

Further, during the period 1/1/2009 – 10/09/2009, on a total amount of 149 issued Import 

Notifications, about 25% regarded the 4 species of Corallium included in Appendix-III. In that 

period, CITES Service of Forestry Corps in Naples has issued 26 re-export certificates, that are 

about 6.5% of the total re-export documents. In these few months of application, more than an 

increasing of the issued certificates (+ 6.5%), Naples Forestry Corps have pointed out an increasing 

of the necessary time to deliver permits, due to the considerable checks of the big amount of preconvention rough, semi-rough, manufactured material, and relative documentation. As already 

mentioned, at this time approximately 40% of the rough/semi-rough coral used for manufactured 

product crafted by Italian companies is purchased in Pacific/Japan/Philippines/PDRC/P of Taiwan 

201
Fig. 1. Import notification delivered from 1/1/2009 to 10/09/2009.

Fig. 2. Re-export certificates delivered from 1/1/2009 to 10/09/2009.

202
area. This is possible because red coral belonging to the species C. japonicum in particular and 

C. elatius and C. secundum presents chemical and physical characteristics that make it very 

similar to C. rubrum, so that the worked products, especially in the smaller items such as small 

beads, cabochons, or other shapes used in jewelry, looks alike very similar from each other and 

consequently it is very difficult to recognize the single species. 

This fact can creates difficulties for those who, as officers of the State Forestry Corps but generally 

CITES Enforcement workers, have to do routine controls required by rules and subsequently issue 

specific certificates for specimens in entry/exit from Italy. 

On the contrary, in rough or semi-rough products it is possible identify peculiar differences in the 

distribution and orientation of color and canals, as you can see in every piece of coral like these 

showed in the next picture (Fig. 3).

As results of such identification problems, there were some cases of shipment seizures about 

manufactured products, even if they were escorted by official declaration of Italian competent 

authority, with labels declaring that they have been obtained by the species “Corallium rubrum,” 

not subjected to CITES provisions. 

Conclusions 

Therefore, on the basis of the considerations set out above, the short lack of time from CITES 

inclusion and the findings after the first few months of application of Reg. (CE) 407/09, there are 

real difficulties associated with: 

• Identification of specimens.

• Increase of administrative activity and time taken for checking of invoices, bills, accounts, 

and other administrative documents, also to cover stocks of coral, everything necessary for 

the issuance of certificates.

Fig. 3.

203
Priority should therefore look for measures to ensure at the same time:

• Conservation of species.

• Avoid of contentious in their application.

• Bearable burden for the certification and control authorities.

• Reliability and efficiency of technical–administrative action to reduce at minimum level the 

possibility of contentious, related in particular to identification of species and big amount 

of pre-convention stocks.

204
IMPLICATIONS OF AN APPENDIX-II LISTING OF THE GENUS

CORALLIUM FOR A SUSTAINABLE INDUSTRY 

Stephen Nash

CITES Secretariat

The presentation examines the implications of a possible listing of the genera Corallium and 

Paracorallium in Appendix-II of the Convention on International Trade in Endangered Species of 

Wild Fauna and Flora (CITES) on the red coral industry. As with any species listed in AppendixII, international trade is allowed under specific circumstances, upon advice from a Scientific 

Authority. The presentation will review the conditions and State obligations under the Convention 

that a listing of Corallium/Paracorallium would entail, and consider practical issues relating to 

law, science, management, and administration. A further consideration is whether such a listing is 

enforceable, taking into account matters of identification, treatment of personal effects, and preConvention specimens. Included in the discussion is consideration of counterfeit red coral and red 

coral substitutes, the possible impacts of a split-listing by species and/or geographical populations, 

the impact on the reporting burden, and taxonomic uncertainty. After a careful consideration of 

the implications of a listing, Parties will be able to decide whether CITES offers the best means 

to ensure protection from over-exploitation and support for a long-term future for the red coral 

industry. 

205
LACK OF COMPREHENSIVE CENSUS DATA HAMPERS CONSERVATION 

AND MANAGEMENT OF CORALLIUM RUBRUM COLONIES 

A. Deidun

Physical Oceanography Unit, University of Malta, Malta 

E-mail: alan.deidun@um.edu.mt

Besides being listed in a suite of European and regional environmental legislation to which Malta 

is party (e.g. Habitats Directive, SPA Protocol, Bern Convention) and in Maltese environmental 

legislation (e.g. LN 311 of 2006, as well as in the local Red Data Book), no systematic census of 

Corallium rubrum (Qroll ahmar in Maltese) has been conducted to date within Maltese coastal 

and offshore waters. Anecdotal evidence exists of the presence of red coral in relatively deep 

waters around the Maltese Islands, but no definite distribution map for the species within Maltese 

waters exists, rendering the adoption of a coherent conservation and management plan for the 

species difficult. Such a situation has arisen from the verbatim transposition of Habitats Directive 

species lists into local environmental legislation. A similar scenario exists for large swathes of 

the Eastern Mediterranean, where records of the red coral species have transpired from sporadic 

sampling events. The situation is compounded by sweeping statements such as the one which 

claims that the only commercially valuable red coral beds are now located along North African 

coasts from Morocco to Tunisia, off the western coast of Sardinia (Bonifacio Straits), and along 

the Spanish coasts. In addition to the need for a comprehensive distribution map of red coral 

in the Mediterranean, the need for a harmonized pan-Mediterranean regulation and management 

framework, possibly under the aegis of FAO, is also felt. At present, whilst Corallium rubrum 

is strictly protected in Malta and Gibraltar, where no exploitation is contemplated, regulated 

exploitation is permissible in a number of Mediterranean countries, including Spain, France, and 

Tunisia.

206
EUROPEAN UNION LEGISLATION ON WILDLIFE TRADE (CITES) 

M. Valentini 

 European Commission DG. ENV – E2 – CITES Team

The presentation aims to give an overview on the European Union legislation implementing the 

CITES provisions, fixing stricter rules, fostering common interpretation, scientific and enforcement 

coordination.

Tasks and responsibilities of Scientific Authorities, as well the rules of procedure of the Scientific 

Review Group, that coordinates the Scientific Authorities of the 27 Member States; Annexes A, 

B, C, and D to the EU Regulation Import, export procedures, and the consequences of the listing;

Stricter rules in the EU legislation; Gaborone amendment; Annotations to the listing; Consultation 

with third countries; How the non detrimental findings are carried out; Delivery of the SRG 

opinion; SRG import restrictions and mechanisms for trade suspension.

207
Working Group Reports
CONCLUSIONS FROM SCIENCE AND MANAGEMENT WORKING GROUP

The working group expressed support for cross-Mediterranean approach to red coral management 

– for example through the GFCM. Desirable to have a common assessment of current management, 

monitoring and research measures across the region.

Need local management plans under this regional ‘umbrella’ – using case studies (e.g. Sardinia/

Hawaii) to improve management.

A working group needs to be formed to work post-conference on developing these plans and to 

seek funding – there is scope to strengthen lobbying for such funding from sources such as EC, 

national governments and private sector.

Management should be based on key scientific parameters such as population structure, density 

genetics, colony and population growth rates (reproduction and recruitment) and connectivity; 

management should be adaptive based on feedback from fishery dependent and independent data 

and input from relevant stakeholders.

The group agreed that shallow water populations are well studied but deeper water populations are 

less well studied and extrapolating from shallow water studies (in selected locations) may not be 

appropriate. 

There is a general consensus that shallow water populations are, or have been, over-exploited but 

the overall extent of decline in the Mediterranean is not known. Therefore, there is a clear need to 

determine the extent of decline and/or recovery. 

Shallow water populations need to be fully protected from harvesting. The vertical distribution 

of these populations depends on local environmental characteristics. However, as a guideline for 

enforcement and management, these populations might be defined by the limits to which airbreathing divers can descend (typically 70 m).

Deep water populations (typically 70-150 m) are a priority for research and fishery management 

because these are the most heavily exploited populations and knowledge of population parameters 

are poorly known. These populations should be managed by selective harvesting using appropriate 

size limits. The deepest populations (> 150 m) are not harvested, some may never have been 

harvested, and should not be subject to harvest and could provide refugia for the species.

The decision to ban non-selective gears was a major step forward in the regulation of the fishery. 

However, there is consensus that management measures and enforcement in the Mediterranean 

overall should be improved further taking into account the case studies referred to above. Such 

management measures should be population specific and include, but not be limited to, minimum 

basal diameter and height of colony and branching pattern, quotas, regulation of collection 

methods, harvest reporting and monitoring, harvest rotation periods and the number and extent of 

no-take zones/protected areas. These management measures should be adaptive and be informed 

by proper and comprehensive scientific research on, inter alia, stock status and management 

209
models. Absence of scientific information is not sufficient excuse to not manage populations on a 

precautionary basis.

Experience has shown that management regimes that are not adequately enforced fail to yield 

the anticipated conservation benefits. There is a strong need for existing and future management 

measures to be properly enforced with inter alia sufficient resources and penalties – including 

confiscation – that are commensurate with the offence and represent a meaningful deterrent.

The working group emphasised the need for further research to underpin the management and 

conservation of the species; research should be collaborative and the results shared between 

Mediterranean countries and internationally. Regular stock assessments and national reporting, in 

the context of regional management, is desirable.

210
CONCLUSIONS FROM THE TRADE WORKING GROUP

It was not possible for the workshop, in plenary, when considering the report of the trade 

working group, to reach any appropriate consensus, on trade related issues, because some 

participants objected to any reference to any CITES listings while others felt that discussion 

of the implications of a hypothetical CITES Appendix-II listing were essential. This report 

represents general agreement of the working group but consensus was not reached on all of 

the text.

Identification

Corallium rubrum is the focus of the Mediterranean Workshop. However, since there are other 

Corallium in trade in the Mediterranean, four of which are already subject to international 

controls, identification difficulties already arise even in the absence of any CITES Appendix-II 

listing. Different species of Coralliidae are utilized in manufacturing of jewelry and semi-finished 

products, and discriminating between species is a challenge. Many of the issues discussed here, 

including identification and pre-Convention materials, are not unique to coral. Many of these 

issues apply to all countries, including Italy and other Mediterranean countries already trading in 

CITES-listed products and lessons may be learned from the implementation of already existing 

CITES listings. It was also noted that industry expertise and opinion should be included and taken 

into consideration for implementation. 

There are six species in the Coralliidae family that are commercially traded. It has been suggested 

that, in the event of an Appendix-II listing, amending CITES Resolution CONF. 12.3, to allow Parties 

to declare worked specimens at the family level would alleviate many identification challenges. 

However, reporting at the family level would limit the data available to asses the impact of trade 

on individual species, and this should be taken in to account in making non-detriment findings. 

Moreover, it was represented by some participants that significant administrative problems could 

arise related to the identification at the family level in deciding whether to issue an export or reexport permit. 

The following were recommended as tools to assist in identification. Industry expertise and 

opinion should be sought to ensure the identification tools are accurately represented. Moreover, 

it is recognized that these tools may not allow identification to the level of species in all cases of 

semi-finished and finished Coralliidae products:

1) ID guide to precious corals – tools need to be available online and in hard copy form; 

2) Reference collection;

3) Training of customs officials and training the ‘trainers;’

4) List of spp. and country of origin;

5) Access to government-approved experts for consultation (customs officials need access to 

scientists);

6) Capacity building (developing countries in particular);

7) Forensic tools (which may not be available for all countries, or feasible when it requires 

damage of the product);

211
8) Secretariat is creating a wiki identification website.

Additional needs: implementation plan useful before adoption; adopt taxonomic reference for corals. 

Administrative Burden

It was recognized that there are heavy administrative burdens associated with a CITES listing, 

including burdens to CITES management authorities for issuing permits and reporting trade, 

burdens to enforcement officials, and to industry. Recognition that national governments could 

develop procedures for dealing with pre-Convention stockpiles of Coralliidae specimens with 

inventory measures. It was also recognized that there are mechanisms under CITES that could 

be used to reduce these burdens (including use of personal effects exemption and streamlined 

permitting procedures). In many CITES countries, these mechanisms are not currently applied 

and it is up to national governments to determine their use. Ultimately, if there is a CITES listing 

proposal, the Parties will need to decide if the conservation benefits to the resources outweigh the 

administrative burdens. As for any CITES listing, it was agreed that there needs to be a real benefit 

and need for conservation of the species before proposing a listing. It is recognized that these will 

be burdens that apply to all countries, including developed countries.

1) There was the recognition that systems are in place to lessen administrative burden, and it 

is the choice of national governments to implement these systems. 

2) Personal effects. Not all countries recognize CITES personal effects exemption (EU, US do 

recognize personal effects exemption). Those that do recognize it, it’s often not recognized 

consistently.

3) Streamline permit procedure (e.g. using partially filled out forms).

4) Delayed implementation in order for countries to ready themselves (and industries) for a 

listing, in order to minimize the burden. 

5) Money and added capacity.

6) Direct animals committee to review/update existing notification on stony corals and 

taxonomic references/guide; do the same for precious corals. This issue is not unique to 

corals and could apply to other species listed under CITES.

Socioeconomics and Livelihoods

Socioeconomic and livelihood impacts that can accompany a CITES listing must be considered. 

For example, it was noted that only 31 shipments of Appendix-III listed species were re-exported 

from Italy, from beginning of 2009, representing an 80% reduction in the trade. A concern was 

also expressed about the perception of a CITES listing as being a ‘non-green.’ The Appendix-II 

listing of Strombus gigas was cited as an example that led to the collapse of the demand for objects 

made from this species. The consumer perception is that they’re buying something endangered. A 

marketing strategy may be beneficial in altering this perception. 

There are many different forms of trade in corals, not just jewelry, and they must also be considered. 

Res. 12.3 potential revision is working towards identification of products at the family level. This 

could be a potential solution to identifying different species in the same product or piece. 

212
It is recognized that there are challenges and enforcement issues relating to the international coral 

trade, such as poaching and illegal harvesting, the most environmentally – and cost-effective 

methods are to manage enforcement at the harvesting site with enforcement of national laws and 

adequate penalties.

It was noted that Taiwan adopted its own enforcement legislation in December 2008. It is critical 

to have cooperation between range states to effectively control the illegal harvest.

213
Appendix
1. Participants and Affiliations

215
Last Name First Name Affiliation Contact Information

Abbiati Marco Università degli Studi di 

Bologna - Italy marco.abbiati@unibo.it

Angeletti Lorenzo ISMAR-CNR - Italy lorenzo.angeletti@bo.ismar.cnr.it

Apa Maurizio ASSOCORAL - Italy info@assocoral.it

Ascione Caterina ASSOCORAL - Italy caterina@ascione.com

Ascione Mauro ASSOCORAL - Italy assocoral@libero.it

Avanzo Cristina Italian Forestry Corps - Italy c.avanzo@corpoforestale.it

Bailey Amy

US Fish and Wildlife Service, 

Office of Law Enforcement - 

U.S.A.

amy_bailey@fws.gov

Bavestrello Giorgio Università Politecnica delle 

Marche - Italy g.bavestrello@univpm.it

Beccaro Maria Antonino De Simone srl - Italy maribecc@tin.it

Bedin Luca

MIPAF Direzione Generale 

della Pesca Marittima e 

dell’Acquacoltura - Italy

l.bedin@politicheagricole.it

Benayahu Yehuda Tel Aviv University - Israel yehudab@tauex.tau.ac.il 

Berney Jaques IWMC - Switzerland iwmcch@bluewin.ch 

Bertelletti Mauro

MIPAF Direzione Generale 

della Pesca Marittima e 

dell’Acquacoltura - Italy

m.bertelletti@politicheagricole.it

216
Last Name First Name Affiliation Contact Information

Bianco Gennaro Università degli Studi di 

Napoli Parthenope - Italy gen-bia@libero.it

Borriello Antonella ASSOCORAL - Italy assocoral@libero.it

Bramanti Lorenzo Università degli Studi di Pisa 

- Italy lbramanti@biologia.unipi.it

Bruckner Andrew NOAA Fisheries - U.S.A. andy.bruckner@noaa.gov

Bunrei Umehara Ring An Jewelry Enterprise 

Co. Ltd. bunbun_umehara@hotmail.com 

Bussoletti Ezio

Università degli Studi di 

Napoli Parthenope, Ministero 

dell’Ambiente - Italy

bussoletti.ezio@minambiente.it

Calligaro Laura Ministero degli Affari Esteri - 

Italy laura.calligaro@esteri.it

Cannas Rita Università degli Studi di 

Cagliari - Italy rcannas@unica.it

Caocci Federica Università degli Studi di 

Cagliari - Italy f.caocci@virgilio.it

Carlson Robert U.S. Consulate General - 

U.S.A. carlsonrw@state.gov

Catalano Valerio Università degli Studi di 

Napoli Parthenope - Italy

valerio.catalano@uniparthenope.

it

Cataldo Gennaro SESTO CONTINENTE - Italy ASSOCORAL@LIBERO.IT

Cattaneo-Vietti Riccardo Università degli Studi di 

Genova - Italy catta@unige.it

217
Last Name First Name Affiliation Contact Information

Cau Angelo Università degli Studi di 

Cagliari - Italy cau@unica.it

Condito Ciro ASSOCORAL - Italy assocoral@libero.it

Cooper Ernie Director Traffic and wildlife/

WWF - Canada ecooper@wwfcanada.org

Corbetta Luisa Italian Forestry Corps - Italy l.corbetta@corpoforestale.it

Cosentino Aldo

Ministero dell’Ambiente e 

della Tutela del Territorio e del 

Mare - Italy

scn-dg@minambiente.it

Costantini Federica Università degli Studi di 

Bologna - Italy federica.costantini@unibo.it

Cottingham David

National Marine Fisheries 

Service/Office of Protected 

Resources - U.S.A.

david.cottingham@noaa.gov

De Simone Annunziata Elena 

Gioia Antonino De Simone srl - Italy info@antoninodesimone.it

De Simone Michele ASSOCORAL - Italy info@desimonefratelli.com

Debreuil Julien Centre Scientifique de Monaco 

- Principato di Monaco julien@centrescientifique.mc

Deidun Alan Physical Oceanography Unit, 

University of Malta - Malta alan.deidun@um.edu.mt

Del Gatto Ernesto Del Gatto srl - Italy info@delgattogioielli.it

Del Gatto Francesco Del Gatto srl - Italy info@delgattogioielli.it

218
Last Name First Name Affiliation Contact Information

Dietrich Jelden Head CITES Management 

Authority - Germany Dietrich.Jelden@BfN.de

Di domenico Sandra Italian Forestry Corps - Italy s.didomenico@corpoforestale.it

Di Stefano Floriana Università degli Studi di 

Napoli Parthenope - Italy

floriana.distefano@

uniparthenope.it

Doneddu Roberto Regione Autonoma Sardegna 

- Italy rdoneddu@regione.sardegna.it

Dounas Costas

Institute of Marine Biology 

and Genetics, Hellenic Centre 

of Marine Research - Greece

kdounas@her.hcmr.gr

Drougas Aimilia

MINISTRY OF 

AGRICULTUREHYDROGEOLOGY 

DIVISION - Greece

DELPHIS@HOL.GR

Ellalloussi Khalid Département de la pêche 

maritime; Rabat - Tunisia elalloussi@mpm.gov.ma

Fischer Johanne FAO johanne.fischer@fao.org

Fleming Vincent Joint Nature Conservation 

Committee - United Kingdom vin.fleming@ jncc.gov.uk

Follesa Maria Cristina University of Cagliari - DBAE follesac@unica.it

Gatta Rossana U.S. Consulate General gattar@state.gov

Grazioli Eleonora Università degli Studi di 

Cagliari - Italy eleonora.grazioli@alice.it

Grigg Richard University of Hawaii - U.S.A. rgrigg@soest.hawaii.edu

219
Last Name First Name Affiliation Contact Information

Harmelin Jean-Georges Centre d’Océanologie de 

Marseille - France

jean-georges.harmelin@univmed.

fr

Kiuchi Hideo

Japan Cooperative Society of 

Commerce and Industry of 

Precious Coral 

i.m.t.kosuge@excite.co.jp

Hsu Hsiao-Ling Japan International Precious 

Coral Research Organization sharon0606@hotmail.com

Iacobelli Luigia F.LLI IACOBELLI SRL - Italy info@iacobelli.it

Iannizzotto Antonio

Ministero dell’Ambiente e 

della Tutela del Territorio e del 

Mare - Italy

iannizzotto.antonio@

minambiente.it

Iazzetti Giovanni Università degli Studi di 

Napoli Federico II - Italy iazzetti@unina.it

Ibero Carlos ATECMA SL, Madrid - Spain cites@atecma.es

Iwasaki Nozomu

Usa Marine Biological 

Institute, Kochi University - 

Japan

iwasakin@cc.kochi-u.ac.jp

Kamei Katsuo

Cooperative Society of 

Commerce and Industry of 

Precious Coral

 i.m.t.kosuge@excite.co.jp

Kosuge Sadao Director Institute of 

Malacology of Tokyo - Japan i.m.t.kosuge@excite.co.jp

Koutsoubas Drosos Dept. Marine Sciences, 

University of Aegean - Greece drosos@aegean.gr

Linneman Jeremy Special Projects Director The 

ocean Foundation - U.S.A. jlinneman@oceanfdn.org

Liverino Vittorio vimach coral di v. liverino srl vimachliverino@alice.it

220
Last Name First Name Affiliation Contact Information

Lungo Ciro Italian Forestry Corps - Italy c.lungo@corpoforestale.it

Mastascusa Vincenza Università degli Studi di 

Napoli Parthenope

vincenza.mastascusa@

uniparthenope.it

Maturani Antonio

Ministero dell’Ambiente e 

della Tutela del Territorio e del 

Mare - Italy

dpn-cites@minambiente.it

Mazza Tommaso ASSOCORAL - Italy assocoral@libero.it

Mei Marco Italian State Forestry Corps - 

Italy m.mei@corpoforestale.it

Moronuki Hideki

Ecosystem Conservation 

Office Fisheries Agency - 

Japan 

hideki_moronuki@nm.maff.go.jp

Nash Stephen CITES Secretariat stephen.nash@cites.org

Noguchi Laura U.S. Fish and Wildlife Service 

- U.S.A. Laura_Noguchi@fws.gov

O’Criodan Colman WWF International, - 

Switzerland cocriodain@wwfint.org

Ozturk Bayram

İstanbul University ,Turkish 

Marine Research Foundation - 

Turkey

ozturkb@istanbul.edu.tr

Pani Marco ASSOCORAL - Italy pani.marco@gmail.com

Papa Marianna Università degli Studi di 

Napoli Parthenope - Italy marianna.papa@uniparthenope.it

Parente Albina U.S. Consulate General Parentea@state.gov

221
Last Name First Name Affiliation Contact Information

Pedoni Claudia Università degli Studi di 

Cagliari - Italy pedoni@unica.it

Pesci Paola Università degli Studi di 

Cagliari - Italy ppesci@unica.it

Roark Brendan

Department of Geography 

Texas A&M University - 

U.S.A.

broark@geog.tamu.edu

Roberts Glynnis NOAA Fisheries - U.S.A. Glynnis.Roberts@noaa.gov

Rodic 

Baranovic Petra State Institute for Nature Protection - Croatia petra.rodic@dzzp.hr

Rossi Sergio Institut de Ciencia i Tecnologia 

Ambientals-UAB - Spain sergio.rossi@uab.cat

Rossi Alessandra

Ministero dell’Ambiente e 

della Tutela del Territorio e del 

Mare - Italy

rossi.alessandra@minambiente.it

Russo Giovanni Università degli Studi di 

Napoli Parthenope - Italy giovanni.russo@uniparthenope.it

Sacco Flavio Università degli Studi di 

Cagliari - Italy flaviosacco@unica.it

Said Taleb Institut national de recherche 

halieutique - Morocco taleb@inrh.org.ma

Salomidi Maria Hellenic Centre for Marine 

Research - Greece msal@ath.hcmr.gr

Sandulli Roberto Università degli Studi di 

Napoli Parthenope - Italy roberto.sandulli@uniparthenope.it

Santangelo Giovanni Università degli Studi di Pisa 

- Italy gsantangelo@biologia.unipi.it

222
Last Name First Name Affiliation Contact Information

Sbarra Giuseppe ASSOCORAL - Italy giuseppesbarra@inwind.it

Scardi Michele Università degli Studi di Tor 

Vergata - Italy mscardi@mclink.it

Scarpati Massimo Coral Fisher oaks57@tiscali.it

Signoretti Federica U.S. Embassy signoretf@state.gov

Stampacchia Paolo Università degli Studi di 

Napoli Federico II - Italy paolo.stampacchia@unina.it

Taleb Said INRH - Morocco TALEB@INRH.ORG.MA

Tambutté Sylvie Centre Scientifique de Monaco 

- Principato di Monaco stambutte@centrescientifique.mc

Tanabe Yasuo

Japan Cooperative Society of 

Commerce and Industry of 

Precious Coral 

 i.m.t.kosuge@excite.co.jp

Taviani Marco ISMAR-CNR - Italy marco.taviani@bo.ismar.cnr.it

Torntore Susan University of Idaho torntore@uidaho.edu

Tsounis Georgios Instituto de Ciencies del Mar 

(CSIC) - Spain georgios@icm.csic.es

Valentini Marco Euripean Commisssion CITES 

Team - Belgium

Marco.VALENTINI@ec.europa.

eu

Vielmini Ilaria Università degli Studi di Pisa 

- Italy ivielmini@biologia.unpi.it

223
Last Name First Name Affiliation Contact Information

Villani Guido CNR - Italy gvillani56@libero.it

Vilne Ilona Nature Protection Board - 

Latvia ilona.vilne@dap.gov.lv

Benayahu Yehuda Dept. of Zoology, Tel Aviv 

University - Israel yehudab@tuaex.tau.ac.il

Weng Hsu-Shu RING AN ENTERPRISE CO., 

LTD - Japan ringan3535@xuite.net

Willens Todd ASSOCORAL - U.S.A. assocoral@libero.it

Xiaoping Lu

The Endangered Species of 

Wild Fauna and Flora Import 

and Export Management 

Office of P.R.C. 

lxpc263@yahoo.com

Zahzah Khaled Direction Générale des Forêts 

Morocco khaledzahzah2000@yahoo.fr

Zoubi Abdelaziz INRH - Morocco abdelaziz_zoubi@hotmail.com

224
2. Workshop Agenda

225
International Workshop on

Red Coral Science, Management, and Trade:

Lessons from the Mediterranean

University of Naples “Parthenope”

Villa Doria D’Angri

Via Petrarca, 80

September 23rd – 26th 2009

The aim of this Workshop is to define a management strategy for the Mediterranean red coral, 

emphasizing the various problems of conservation and the most suitable methods of sustainable 

harvest and trade.

Preliminary Program

September 22nd afternoon: 

Arrival of participants and registration at Villa Doria D’Angri, Via Petrarca, 80.

4.30pm - Meeting of the Steering Committee

September 23rd Morning: Plenary Session 

8.30am - Registration

9.30am - Welcome of Authorities (Chair: E. Bussoletti/D. Cottingham/A. Maturani)

Minister/Vice Minister MATTM

Dr. Aldo Cosentino, Director General DPN MATTM 

Prof. Gennaro Ferrara, Rector of University of Naples “Parthenope”

Prof. Raffaele Santamaria, Dean of Faculty of Science and Technology, University of Naples 

“Parthenope”

10.00am - Mediterranean red coral (Corallium rubrum): historical assessment. An overview 

of the role red coral played in the Mediterranean history, culture, traditions and economy.

•	 10.00-10.25 - C. Ascione - The precious Mediterranean red coral: an historical, cultural, 

economic heritage.

•	 10.25-10.50 – J. G. Harmelin Lacaze-Duthiers’ legacy: advances in scientific knowledge 

of Corallium rubrum from French works after 1864

226
•	 10.50- 11.15 G. Santangelo, L. Bramanti, I. Vielmini, M. Iannelli - The history of research 

and harvesting of the Mediterranean red coral: potential and limits.

11.15am - Coffee Break

11.30am-1.15pm - Introductory speeches 

•	 11.30-11.55 – S. Rossi, G. Tsounis, J.-M. Gili: The deep transformation of the Mediterranean 

red forest: the Catalonian coast case study 

•	 11.55-12.20 – R. Cannas, Caocci F., Follesa M.C., Grazioli E., Pedoni C., Pesci P., Sacco 

F., A. Cau: Multidisciplinary data on the status of red coral (Corallium rubrum) resource 

in Sardinian seas (Central Western Mediterranean).

•	 12.20-12.45 - A. W. Bruckner: Rate and extent of decline in Corallium (pink and red 

coral) populations: are existing data adequate to justify a CITES Appendix II listing?

•	 12.45-13.10 - R. W. Grigg: Management of precious corals: local science and enforcement 

versus a listing by CITES

1.15pm - Lunch time

2.45pm-6.30pm: 

•	 2.30–2.50 - M.Taviani, A.Freiwald, L. Beuck, L.Angeletti, A. Remia, A. Vertino, M. 

Dimech, P. Schembri: The deepest known occurrence of the precious red coral Corallium 

rubrum (L. 1758) in the Mediterranean Sea.

A. Vertino, H. Zibrowius, M. Taviani. Fossil ancestors of family Coralliidae in the 

Mediterranean basin

P. Montagna, D. Vieulzeuf, M. López Correa,.J. Garrabou, M. Taviani, C. Marschal, C. 

Linares, M. McCulloch, S. Silenzi, A. Freiwald: Geochemical investigation of Corallium 

rubrum from shallow and bathyal settings in the Mediterranean Sea

Harvest, Manufacturing and Trade – (Chair: R. Cattaneo-Vietti/N. Iwasaki/L. Noguchi)

•	 2.50-3.10 – A. Bailey: Overview of Law Enforcement Issues Regarding the Trade in 

Corallium.

•	 3.10-3.30 – A.E. De Simone: Diffusion of Coral in the world; symbol, use and trade.

•	 3.30-3.50 – C. Dounas, D. Koutsoubas and M. Salomidi: Red coral (Corallium rubrum, L. 

1758) fisheries in the Greek Aegean and Ionian Seas.

•	 3.50-4.10 – A. Dridi, A. Zoubi and S. Taleb: The exploitation of red coral in Morocco.

227
•	 4.10-4.30 – B. Öztürk: Red coral and ıts actual sıtuatıon ın Turkey.

4.30-4.45 – Coffee Break 

•	 4.45-5.05 – M. Pani: Analysis of FAO Statistics of harvest of Corallium rubrum in the 

Mediterranean 1978-2007.

•	 5.05-5.25 - P. Stampacchia: Il comparto del corallo di Torre del Greco.

•	 5.25-5.45 – S.J. Torntore: Coral as Product: Analyzing Production and Market Perspectives.

•	 5.45-6.05 – R. Doneddu: Red coral management strategies in Sardinian coastal waters: a 

successful experience.

•	 6.05-6.25 - E. Cooper: Development of a Guide to the Identification of Precious Corals

September 24th 9.00am-1.00pm: Plenary Session 

Science: Biology and ecology of red coral (Chair A. Zoubi/M. Abbiati/B. Roark)

The status of knowledge on biology and ecology of the Mediterranean red coral Corallium rubrum 

(L 1758) will be discussed with comparative information for Pacific species.

•	 9.00-9.20 – M. Abbiati, F. Costantini, C. Fauvelot, E. Pintus: Genetic structuring of 

Corallium rubrum in the Mediterranean Sea: implication for management and conservation.

•	 9.20-9.40 – G. Bavestrello, C. Cerrano, R. Cattaneo-Vietti: Biological interactions affecting 

the growth rates of red coral (Corallium rubrum) colonies.

•	 9.40-10.00 – L. Bramanti, Vielmini I., Santangelo G.: Long-term settlement plates: some 

clues for demography and restoration of Mediterranean red coral (Corallium rubrum) 

shallow water populations.

•	 10.00-10.20 – B. Calcinai, C. Cerrano, N. Iwasaki and G. Bavestrello: Biodiversity and 

ecology of the sponge boring into precious corals: a worldwide overview.

•	 10.20-10.40 – L.A. Chessa andM. Scardi: Biometrical analysis and observations on deep 

red coral Corallium rubrum (L.) in N.W.Sardinia shoals (Italy).

•	 10.40-11.00 – F. Costantini, C. Fauvelot, E. Pintus, M. Abbiati: Effective larval dispersal 

and scales of connectivity in Corallium rubrum.

11.00-11.15 – Coffee Break

228
•	 11.15-11.35 – F. Costantini, M. Taviani, A. Remia, E. Pintus, P.J. Schembri, M. Abbiati: 

Deep-water Corallium rubrum (l. 1758) from the mediterranean sea: preliminary genetic 

characterization.

•	 11.35-11.55 – C. Ferrier-Pagès: Feeding characteristics of the red coral Corallium rubrum.

•	 11.55-12.15 – N. Iwasaki, T. Fujita, G. Bavestrello, R. Cattaneo-Vietti: Small-scale 

distribution of Japanese red coral off Amami Island, Southern Japan.

•	 12.15-12.35 - E.B. Roark, T.P. Guilderson, R.B. Dunbar: Radiocarbon-based ages and 

growth rates of Hawaiian Corallium sp.

•	 12.35-12.55 – S. Tambutté: Biomineralization in Corallium rubrum: organic matrix 

comparative study of sclerites and skeleton.

•	 12.55-13.15 – I. Vielmini, Bramanti L., Tsounis G., Rossi S., Gili J.M., Cattaneo-Vietti R., 

Santangelo G.: Corallium rubrum age structure determination.

1.15pm - Lunch Time

2.45pm – 6.45pm: Plenary Session 

Management (Chair A. Maturani/V. Fleming)

The group will compile and assess information on the existing national management strategies, 

how the various jurisdictions are operating, and their adequacy in the different Mediterranean 

experiences. Comparisons with other World regions experiences are welcome (for example Hawaii 

black coral and Pacific red coral harvest).

•	 2.45-3.05 - A. W. Bruckner: Changes in size structure of fished Corallium rubrum

populations and implications for sustainable management and trade.

•	 3.05-3.25 – J. Dietrich: Lessons learned from national trade regulations established for the 

red coral, Corallium rubrum.

•	 3.25-3.45 – V. Fleming: CITES non-detriment findings for precious corals.

•	 3.45-4.05 - S. Kosuge, H. Kiuchi, Y. Tanabe and K. Kamei: Precious coral fisheries 

regulation in the western Pacific area in comparison with Sardinian regulation. 

•	 4.05-4.25- M. Mei, L. Corbetta, S. Di Domenico, P. Pandolfi: The inclusion of Corallium 

japonicum, Corallium konojoi , Corallium elatius and Corallium secundum in Appendix 

III of CITES. The Italian experience of Corpo Forestale dello Stato.

229
•	 4.25-4.45 - S. Nash: Implications of an Appendix II listing of the genus Corallium for a 

sustainable industry. 

•	 4.45-5.05 A. Deidun: Lack of comprehensive census data hampers conservation and 

management of Corallium rubrum colonies

•	 5.05-5.25 –. M. Valentini: European Union legislation on wildlife trade (CITES)

•	 5.25-5.45 M. Scarpati: Red Coral harvesting: past, present, future (Video).

5.45 – 6.00 pm Coffee Break

6.00 – 7.00 Discussions (potential contingency to absorb delays)

8.30pm - Gala Dinner at Villa Doria D’Angri Terrace

September 25th 9.30am-1.30pm: Meetings of three Working Groups

a) Harvesting , Manufacturing and Trade Working Group (P. Stampacchia, L. Noguchi)

b) Scientific Working Group (G. Bavestrello, B. Roark)

c) Management Working Group (G. Santangelo, V. Fleming)

1.30 pm – Lunch Time

2.00pm-3.30pm – continuation of Working Groups Meetings

3.30pm - Visit to Torre del Greco with the following program:

- visit to the “Red Coral Museum”;

- visit to a red coral manufacturing emplacement;

- dinner and historical Neapolitan music show.

September 26th morning/afternoon:

 Final Common Report of the three WGs (Chair: E. Bussoletti/D. Cottingham)

Outcomes to correctly sustain harvest and trade of Mediterranean red coral, potentially 

extended to Pacific red coral. 

• Sustaining research on specific scientific topics on which the lack of knowledge was 

assessed, which will lead to a rational harvesting (based on sound scientific data) of the 

resource and to rearing experiments.

230
• Evaluate the opportunity, as a follow-up of the event, to establish a Capacity Building 

Program specifically addressed to red coral in order to transfer know-how and positive 

experiences to other countries.

• Developing recommendations on changes to management approaches to enhance 

conservation.

• Identifying strong local management system and enforcement efforts to prevent 

unsustainable harvest of red coral, and analysing gaps where more steps are needed for 

the management

• Recommending additional measures that could be implemented on a local, regional, 

national, or international scale that are necessary to address possible threats affecting 

red coral and that could help to foster population recovery, and where these need to be 

applied.

OFFICIAL EVENTS:

∗	 Gala Dinner at Villa Doria D’Angri on September 24th, 20.30

∗	 On the evening of September 25th, a visit to Torre del Greco with the following program 

is planned:

- visit to the “Red Coral Museum”;

- visit to a red coral manufacturing emplacement;

- Dinner and historical Neapolitan music show.

231
3. Workshop Coordinators: 

Ezio Bussoletti 

David Cottingham

232
4. Steering Committee:

Marco Abbiati, Italy

Giorgio Bavestrello, Italy

Andy Bruckner, USA

Riccardo Cattaneo-Vietti, Italy

Angelo Cau, Italy

Jean-Georges Harmelin, France

Sadao Kosuge, Japan

Drosos Koutsoubas, Greece

Jeremy Linneman, USA

Laura Noguchi, USA

Glynnis Roberts, USA

Serge Rossi, Spain

Giovanni Russo, Italy

Roberto Sandulli, Italy

Giovanni Santangelo, Italy

Marco Taviani, Italy

Abdelaziz Zoubi, Morocco

233