Corals Mediterranean Eng

The corals of the
Mediterranean
The corals of the

Mediterranean
Index
01. Introduction
Brief explanation of the ta xonomy of anthozoans and the terminology used in this document
08
Configuration
Exoskeletons for all tastes
In touch
12
The evolution of corals in the Mediterranean

Exclusive corals and imported corals
20
Coral reefs
Coralline algae
Large concentrations of anemones
On rocks, walls and hard substrates
In caves and fissures
On muddy and sandy floors
In shallow waters and at great depths
Dirty water
Living on the backs of others
- On algae and seagrasses
- On living creatures
In company: corals and symbiotic/commensal animals
32
02. Physical characteristics of corals
03. Coral species of the Mediterranean
04. Coral habitats and corals as habitats
05. Coral reproduction
Sexual and asexual

Synchronized reproduction
Oviparous, viviparous
[]
04
The corals of the Mediterranean
Berried anemone (Alicia mirabilis) OCEANA/ Juan Cuetos
Budding, fission and laceration

Egg, larva/planula and polyp phases
Separate sexes and hermaphrodites
38
Trouble with the neighbours: space

Grow th
Size mat ters
A seat at the table: coral nutrition
Guess whos coming to dinner: natural predators of corals
Corals with light
Corals that move
46
Coral diseases
Climate change
Other anthropogenic ef fects on corals
- Ripping out colonies
- Chemical pollution
- Burying and colmatation
Fishing and corals
52
Commercial exploitation of corals

Corals and medicine
09. Protected corals
56
10. Oceana and corals
60
06. The fight for survival
07. Threats to corals
08. Coral uses
- Prohibiting the use of destructive fishing gear over coral seabeds

- Regulating the capture of corals
[]
01.
Leptogorgia sarmentosa OCEANA/ Juan Cuetos
[]
Introduction
There are corals which live as solitary animals

or in colonies, composed of rigid, semi-rigid
or sof t structures, and which protect themselves under rock formations or grow erect
like trees; there are those which prefer exposure to the sun or which live in zones of darkness; those which generate light or which
have medicinal properties, existing in shallow waters or at depths of more than 5,000
meters; those with polyps that grow anew
each year, or in colonies as old as 50 to 1,000
years on reefs which have taken more than
8,000 years to develop. Although the number
of coral species found in the Mediterranean represents less than 5% of those ex tant
throughout the world today, the diversity of
the types and forms of life serve as an example to us, demonstrating the full importance
of these animals in relation to the global marine ecosystem.
Corals are simple animals and as such, are
capable of forming very complex and diverse
communities. Contrary to popular belief, simple organisms show the highest capacity for
adaptation and mutation, since complex organisms are more specialized and therefore
less likely to undergo genetic and physical
modifications over a short period of time.
False coral (Myriapora truncata) OCEANA/ Juan Cuetos
Fire coral (Millepora planata) OCEANA/ Houssine Kaddachi
Many people believe that corals are plants.

This is because the great majority of these
creatures are species which live fixed to the
substrate, and with a quick glimpse they do
not seem to be very active. Because we are
terrestrial animals, we are used to drawing a
distinction between plants and animals according to their respective ability to move.
Yet, what is an obvious observation as far as
land creatures are concerned, does not apply to the sea. Appearances to the contrary,
there are other animals which also spend all
or part of their life anchored to rocks or other
substrates, or even to other organisms. This
is seen in porifers (sponges), bryozoans, hydrozoans and a great number of worms, mollusks and crustaceans. The fact that some
species look like tree branches only serves
to increase the confusion.
Corals are animals whose cells are organized
in tissues. They have a nervous system, grow
and reproduce, form colonies and can feed
directly from the organisms which surround
them in the water.
All of them are uniquely marine species which
exist in all of the ocean habitats known to us,
from shallow water and tide pools to the greatest depths known to support marine life.
[]
01. Introduction
Corals belong to one of the oldest ex tant

classes of animals in the world. Their fossil
remains can be traced back to the pre-Cambrian era,1 when there was a great surge in
oceanic life over 500 million years ago.
Occupying 1.1% of the surface of the worlds

oceans and 0.3% of all salt water, the Mediterranean no longer shelters the great coral
reefs that thrived 60 million years ago. This
is due to millennia of climactic and oceanographic changes. However, even today this
sea harbors a spectacular array of corals, including some which are not found anywhere
else.
Brief explanation of the

taxonomy of anthozoans
and the terminology
used in this document
The very term coral is ambiguous in itself,

since it can be a common term for a few species with rigid skeletons or, specific anthozoan groups. Sometimes, it is used to describe species which belong to other classes
of marine life, such as hydrozoans and bryozoans. This occurs with fire, or stinging, coral
(Millepora sp.) and with false coral (Myriapora
truncata).
We have selected to use corals as the name
to group together all anthozoans species, including true corals, black coral, sea fans (or,
gorgonians), sea feathers, and anemones.
We do not include here the other species
commonly referred to as corals, such as fire
coral. These are hydrozoans, a class of animals with distinct dif ferences.
The anthozoans, or flower animals from the

original Greek translation, are a class within
the Cnidaria phylum. They are a group of of
animals which spend their whole life in the
polyp phase.
[]
The cnidarians, or coelenterates, derive their

name from their cnidocy tes or stinging cells
- cnidocy te means stinging or itching needle,
derived from the Greek word knid (net tle).
The other older name they are known by,
coelenterates means vacuous intestine - from
the Greek koilos (vacuous) and enteron (intestine).
The Cnidarian phylum is divided into four
classes of species: hydrozoans, cubozoans,
scyphozoans and anthozoans.
The hydrozoans and cubozoans spend part

of their life as medusas and part as polyps.
The scyphozoans only exist as medusas and
never enter the polyp stage. Anthozoans
only exist as polyps and do not go through
a medusoid phase. The cnidarians once included a fif th class, conulata, which went extinct in the Triassic period.
As for the anthozoans, which we have decided to treat as corals and to which this writing is dedicated, they have traditionally been
subdivided into two subclasses: Octocorallia
and Hexacorallia.
The Octocorallia (or Alcyonacea) derive their

name from the 8-fold tentacular symmetry of
their polyps, and an equal number of septa
or complete, but unpaired, mesentery (the
layers of the gastrovascular cavity which
reaches from the mouth to the anus). They divide into five orders: Stolonifera (organ-pipe
coral; tree fern coral), Alcyonacea (sof t corals), Gorgonacea (gorgonians, or sea fans;
sea feathers), Helioporacea or Coenothecalia
(Indo-Pacific blue coral) and Pennatulacea
(sea pens).
Hexacorallia (or Zoantharia) is not only the

name for the species with six tentacles, but
also signifies the species with more than
eight tentacles (many times they are found in
multiples of six), with six complete and paired
mesenteries. They divide into seven orders:

Actinaria (sea anemones), Scleractinia or Madreporaria (stony corals), Ceriantharia (tubedwelling anemones), Antipatharia (black
corals), Corallimorpharia (coral anemones),
Zoanthidea or Zoantharia (colonial anemone)
and Ptychodactiaria. Other orders that used
to belong to this class are now ex tinct, such
as the Rugosa, Tabulata, Heterocorallia, etc.
Two-thirds of all anthozoans in the world belong to this subclass.
Various colourations of the Leptogorgia sarmentosa
OCEANA/ Juan Cuetos
In the second half of the 20th century, some

authors proposed a third subclass to unite
Ceriantharia and Antipatharia under a common order, Ceriantipatharis, given their similarities during the larval stage as well as their
unpaired mesenteries, among other common
traits. This in turn was a subclass that could
be divided into two very distinct orders: the
Ceriantharia, or tube-dwelling anemones,
which can retract entirely into their tubes,
and the Antipatharians, or black corals, which
have retractable tentacles. In contrast to all
other anthozoans, the lat ter do not form a
ring around their mouths.
There is no scientific consensus as to the

classification of these animals. Anthozoa is
an animal phylum which has yet to be adequately defined. This can be expected, as
the ta xonomic classifications and the subsequent identification of their related species
can be very divergent.
The ta xonomic classification which we have
decided to use in this report is the one used
in Sistema Naturae 20002 (excluding the
Scleractinia, which follow the definition of
Vaughan T.W. & J.W. Wells, 19433, updating
certain types and species based on the MARBEF,4 ITIS5 and Hexacorallia of the World6 databases). On the other hand, we have kept
the names Octocorallia and Hexacorallia instead of Alcyonacea and Zoantharia to avoid
confusing, respectively, the two subclasses
and two orders.
[]
02.
Physical characteristics
of corals
White gorgonia (Eunicella singularis) OCEANA/ Juan Cuetos
[]
A coral is a polyp which can live alone or in

colonies and cover itself with a hard or sof t
exoskeleton, but its overall constitution is
rather simple.
The polyps are sac-like with radial symmetry and two layers of tissue: an ex ternal one
called the ectoderm or epidermis, and an internal one known as the grastrodermis. There
is also a third gelatinous layer in between
called the mesoglea.
The polyp produces calcium carbonate from
within the ectoderm for the purpose of building the firm skeleton of many corals.
On their bot tom side they have an anchoring or basal disk which is used to anchor the
polyp to the substrate or other structure securing a large colony, while the disk-shaped
tentacular mouth is found on the upper side,
facing away from the substrate.
Corals only have one orifice serving as both

mouth and anus, leading to the pharynx, a
short tube connecting the mouth with the
gastrovascular cavity. This is separated by
the wall of the septa or mesentery, forming
the chambers in which the digestive cells are
located.
One particularly characteristic trait of anthozoan and other cnidarians are the cnidos
(greek), stinging needles known as cnidocy tes. There are three dif ferent types:
1) nematocysts, shaped like small harpoons,
which can shoot out and penetrate the tissue
of prey, inoculating it with the toxin they contain. They are found in the tentacles and gastrovascular cavity of all anthozoans;
2) spirocysts, only found among hexacorallias, which use adhesion with their victims instead of needles; and
3) ptychocysts, also adhesive, and used by
ceriantharias (tube-dwelling anemones) to
construct their tubes.
Configuration
Corals can be found living in isolation or in
immense colonies; they may display only
their sof t bodies live inside a tube or create
erect structures that are rigid, semi-rigid or
sof t and on which they anchor their polyps.
They may rise into branch formations, with
many or few dendrites, or carpet vertical
walls on the sea-floor, making it look like a
carpet of grass; they may take on the aspect
of cushions, balls, feathers, whips, globules,
etc. There are also those which have taken
over the ex ternal housing of sponges or other
corals.
The great variety these colonies present has
created an endless naming process and attempt to categorize them. Some have lit tle
branching and are whip-like (such as Elissela
paraplexauroides or Viminella flagellum), cable-like (such as Eunicella filiformis), or feather-like (such as Virgularia mirabilis or those of
the Pennatula type).
Of ten, these forms reflect a specialized adaptation for survival in dif ferent marine environments, conditioned by hydrodynamics,
temperature, etc.
In some cases, the coral colonies form treelike structures, like most gorgonians or some
scleractinias such as Dendrophyllia ramea or
Pour talosmilia anthophyllites. For these types
of colonies, the structure is usually rigid or
White gorgonia (Eunicella singularis) with protruding calyces
OCEANA/ Thierry Lanoy
[]
semi-rigid, but there are also sof t body colonies, such as dead mans fingers (Alcyonum
spp.) or the sea feathers (Pennatulacea),
among others.
One of the structures most characteristic of

coral formation is the reef. In the Mediterranean, coral reefs are scarce and there are
only a few species of colonial anthozoans
that have the capacity to create them.
However, there are other cnidarians which

form complex and sizeable colonies, even
though these may not always be considered
reefs. Some examples of these are the anemone colonies of the type Epizoanthus, Parazoanthus and Cor ynactis, or some corals living
in colonies such as the Madracis pharensis,
Astroides calycularis, Polycyathus muellerae,
Phyllangia mouchezii and Hoplangia durotrix.
Clavularia sp. OCEANA/ Juan Cuetos
One must not forget that although the small

size of some of the stoloniferas and alcyonaceas colonies makes them less visible, the
abundance of individual specimens may be
considerable, enough to create micro habitats. This is the case of the Clavularia sp., Cornularia sp., or Maasella edwardsi.
Although the general perception of corals
is usually one of large colonies, there are a
number of species which either live in isolation or in small groups.
[10]
White gorgonia (Eunicella singularis) with bulging calyces

OCEANA/ Juan Cuetos
Solitary anthozoans may be found living in the

habitats created by other corals. Some deepwater species are of ten found existing in cold
water coral reefs. In the Mediterranean, this
is seen with species such as Desmophyllum
cristagalli or Stenocyathus vermiformis.7
All anemones live in isolated units or in small

colonies, although they can sometimes form
habitats when there are enough of them in a
given area. This also occurs with many true
corals of the Balanophyllidae and Caryophyllidae families.
An alternative favored by some corals is to occupy structures made by other anthozoans.
The sof t coral Parer y thropodium coralloides
and the false black coral (Gerardia savaglia)
form such habitats. While they are capable of
creating branch-like colonies on their own,
they sometimes invade gorgonian colonies
and partly or entirely cover them over.
Some species of corals may also appear different according to the environmental conditions in which they have grown.
Although Cladocora caespitosa takes its

name from its typical presentation in sweeping grass carpets or as small cushions, they
may rise into shrub-like formations when the
hydrodynamics are low and permit it. The
white (Eunicella singularis) or yellow (Eunicella cavolini) gorgonia, may also look dif fer-
Maasella edwardsii OCEANA/ Juan Cuetos
ent depending on the ambient hydrodynamics;8 while some hardly branch at all, and the
branching appears flat, they may also have a
fuller, bushy appearance. Even their exoskeletons can me modified.9
Exoskeletons for all tastes
The exoskeleton of hard corals (scleractinias)

is composed of a crystallized form of calcium
carbonate (CaCO3), known as aragonite,10
which also composes the shells of many mollusks. In rare cases or with varying marine
chemicals, aragonite has been found to be a
substitute of calcite.11
The high mineralization rate of the soluble calcium carbonate found in corals is ma ximized
in certain symbiotic algae (zooxanthelae) living in their tissue.12
Tube anemone (Cerianthus membranaceus) OCEANA/ Juan Cuetos
There are other corals which use composites of proteins, carbohydrates and allogeneic composites such as gorgonian to form
a horn-like skeleton13, sometimes dot ted with
calcareous spicules, common among octocorallias. It is sof ter than scleractinias, making
for greater flexibility. Some octocorallias have
opted for an intermediate system, substituting
gorgonian for calcium carbonate. This is the
case of scleractinias. Thanks to these alterations, their structure is more rigid. This can be
seen with red coral (Corallium rubrum).
There are also examples of sof t skeletons,

as with anemones and zoantharias. Or those
that have opted for a tube instead of a skeleton, using stinging cells (ptychocysts) and
mucus to insulate the polyp, and to fix sand
and other particles14 in the manner of some
polychaete worms.
In touch
When polyps live in colonies, they must find

ways to coordinate themselves. For gorgonians and other octocorallias living in branching
colonies, the polyps are in contact internally
via living tissues such as the coenenchyme. In
addition, the gastrovascular cavities are also
intercommunicated via canals and tubes.
But if there can be only one particularly striking technique to stay in touch with one another, it is that exhibited by the stoloniferas
corals. These are simple polyps that are
united by means of of fshoots containing internal canals which maintain continuous contact. When a single polyp or its of fshoots is
touched, all of the animals retract simultaneously into their calyx.
[11]
03.
Coral species
of the Mediterranean
Mediterranean madreporaria (Cladocora caespitosa) OCEANA/ Juan Cuetos
[12]
More than 200 species of coral (from a total of 5,600 species which have been described worldwide, 500 of which are in Europe) live in the Mediterranean. Some of the species living there are
endemic, while others have a subtropical origin from the warmer waters of the Atlantic. Still others
are more common in arctic zones, while some are found everywhere.
Anthozoan species found in the Mediterranean:
OCTOCORALLIA
Alcyionacea
Pennattulacea
Alcyionidae
Funiculinidae
Dead mans fingers Alcyonium acaule

Dead mans fingers Alcyonium palmatum
False red coral Parerythropodium coralloides
Paralcyionidae
Maasella edwardsi
Paralcyonium spinulosum
Gorgonacea
Acanthogorgiidae
Acanthogorgia armata
Acanthogorgia hirsuta
Elliselliidae
Elisella paraplexauroides
Viminella flagellum
Gorgoniidae
Yellow sea fan/gorgonia Eunicella cavolini

Cable gorgonia Eunicella filiformis
Eunicella gazella
Senegalese gorgonia Eunicella labiata
White gorgonia Eunicella singularis
Pink gorgonia Eunicella verrucosa
Guinea gorgonia Leptogorgia guineensis
Portuguese gorgonia Leptogorgia lusitanica
Leptogorgia sarmentosa
Leptogorgia viminalis
Isididae
Isidella elongata
Plexauridae
Bebryce mollis
Echinomuricea klavereni
Muriceides lepida
Red gorgonia Paramuricea clavata
Spiny gorgonia Paramuricea macrospina
Crown gorgonia Placogorgia coronata
Placogorgia massiliensis
Spinimuricea atlantica
Spinimuricea klavereni
North Sea gorgonia Swiftia pallida
Villogorgia bebrycoides
Tall sea pen Funiculina quadrangularis
Kophobelemnidae
Koster sea pen Kophobelemnon stelliferum

Leuckart sea pen Kophobelemnon leucharti
Veretillidae
Cavernularia pusilla
Pluma de mar redonda o Veretilo Veretillum cynomorium
Pennatulidae
Sea Feather Pennatula aculeata

Phosphorescent Sea Feather Pennatula phosphorea
Red Sea Feather Pennatula rubra
Grey Sea Feather Pteroeides griseum
Virgularidae
Sea Pen VIrgularia mirabilis
Stolonifera
Cornularidae
Cornucopia Cornularia cornucopiae

Cervera Cervera atlantica
Clavulariidae
Common clavularia Clavularia crassa

Dwarf clavularia Clavularia carpediem
Clavularia marioni
Clavularia ochracea
Rolandia coralloides
Sarcodyction catenata
Scleranthelia microsclera
Scleranthelia rugos
Primnoidea
Callogorgia verticillata
Coralliidae
Red Coral Corallium rubrum
[13]
HEXACORALLIA
Actiniaria
Andresiidae
Andresia parthenopea
Edwardsiidae
Edwardsia beautempsi
Edwardsia claparedii
Edwardsia grubii
Edwardsia timida
Edwardsiella janthina
Edwardsiella carnea
Scolanthus callimorphus
Halcampoididae
Synhalcampella oustromovi
Halcampella endromitata
Halcampoides purpurea
Haloclavidae
Anemonactis mazeli
Mesacmaea mitchelli
Mesacmaea stellata
Peachia cylindrica
Peachia hastata
Boloceroididae
Bunodeopsis strumosa
Actiniidae
Actinia atrimaculata
Actinia cari
Actinia cleopatrae
Actinia crystallina
Actinia depressa
Beadlet anemone Actinia equina
Strawberry anemone Actinia fragacea
Actinia glandulosa
Actinia judaica
Actinia mesembryanthemum
Actinia phaeochira
Actinia rondeletti
Actinia rubra
Actinia rubripuncatata
Actinia striata
Actinia zebra
Anemonia cereus
Mediterranean snakelocks anemone Anemonia sulcata
Anthopleura ballii
Bunodactis rubripunctata
Gem Anemone Bunodactis verrucosa
Deep-water Anemone Condylactis aurantiaca
Thick tentacle anemone Cribrinopsis crassa
Paranemonia cinerea
Paranemonia vouliagmeniensis
Pseudactinia melanaster
Actinostolidae
Paranthus chromatoderus
Paranthus rugosus
[14]
Aiptasiidae
Aiptasia carnea
Aiptasia diaphana
Aiptasia saxicola
Trumpet anemone Aiptasia mutabilis
Aiptasiogeton pellucidus
Aliciidae
Alicia costae
Alica Alicia mirabilis
Aurelianiidae
Imperial anemone Aureliana heterocera
Condylanthidae
Segonzactis hartogi
Segonzactis platypus
Diadumenidae
Orange anemone Diadumene cincta

Orange striped anemone Haliplanella lineata
Hormathiidae
Actinauge richardi
Cloak anemone Adamsia carciniopados

Amphianthus crassus
Sea fan anemone Amphianthus dohrnii
Parasitic anemone Calliactis parasitica
Hormathia alba
Hormathia digitata
Hormathia coronata
Hormathia nodosa
Paracalliactis lacazei
Paracalliactis robusta
Isophellidae
Club tipped anemone Telmatactis cricoides

Swimming anemone Telmatactis forskalii
Telmatactis solidago
Metridiidae
Plumose or Frilled anemone Metridium senile
Phymanthidae
Phymanthus pulcher
Sagartiidae
Actinothoe clavata
Actinothoe sphyrodeta
Daisy anemone Cereus pedunculatus
Kadophellia bathyalis
Octophellia timida
Sagartia elegans
Sagartia troglodytes
Sagartiogeton entellae
Sagartiogeton undatus
Gonactiniidae
Gonactinia prolifera
Protanthea simplex
HEXACORALLIA (continued)
Corallimorpharia
Zoanthidea
- Corallimorphidae
Epizoanthidae
Corynactis mediterranea
Jewel anemone Corynactis viridis
- Sideractiidae
Sideractis glacialis
Scleractinia
Caryophyllidae
Cup coral Caryophyllia calveri

Cup coral Caryophyllia cyathus
Southerm cup coral Caryophyllia inornata
Devonshire cup coral Caryophyllia smithii
Ceratotrochus magnaghii
Coenocyathus anthophyllites
Coenocyathus cylindricus
Desmophyllum cristagalli
Hoplangia durotrix
Tuft coral Lophelia pertusa
Paracyathus pulchellus
Sphenotrochus andrewianus
Polycyathus muellerae
Pourtalosmilia anthophyllites
Phyllangia mouchezii
Thalamophyllia gasti
Faviidae
Thin tube coral Cladocora caespitosa

Cladocora debilis
Flabellidae
Javania cailleti
Monomyces pygmaea
Guyniidae
Guynia annulata
Stenocyathus vermiformis
Dendrophylliidae
Orange coral Astroides calycularis

Cup coral Balanophyllia cellulosa
Cup coral Balanophyllia europaea
Scarlet and gold star coral Balanophyllia regia
Cladopsammia rolandi
Yellow tree coral Dendrophyllia cornigera
Yellow coral Dendrophyllia ramea
Sunset cup coral Leptopsammia pruvoti
Brown epizoanthidae Epizoanthus arenaceus

Epizoanthis frenzeli
Epizoanthus incrustans
Epizoanthus mediterraneus
Epizoanthus paguricola
Epizoanthus paxi
Epizoanthus steueri
Epizoanthus tergestinus
Epizoanthus univittatus
Epizoanthus vagus
Epizoanthus vatovai
Parazoanthidae
Gerardia lamarcki
False black coral Gerardia savaglia
Yellow cluster anemone Parazoanthus axinellae
Zoanthidae
Palythoa axinellae
Palythoa marioni
Zoanthus lobatus
Antipatharia
Antipathes dichotoma mediterranea

Antipathes gracilis fragilis
Antipathes subpinnata
Bathypathes patula
Leiopathes glaberrima
Parantipathes larix
Ceriantharia
Mediterranean tube anemone Cerianthula mediterranea

Tube anemone Cerianthus lloydii
Colored tube anemone Cerianthus membranaceus
Dohm tube anemone Pachycerianthus dohrni
Pachycerianthus solitarius
Arachnantus nocturnus
Dwarf tube anemone Arachnantus oligopodus
Oculinidae
White Madrepora Madrepora oculata

Oculina patagonica
Pocilloporiidae
Star coral Madracis pharensis
[15]
The evolution of corals

in the Mediterranean
Six million years ago, towards the end of the
Miocene (Messinian) period, there were still
some remains of coral reefs from the following
genuses in the Mediterranean:15 Porites, Tarbellastrea, Siderastrea, Plesiastraea, Favites,
St ylophora o Acanthastrea. Some of them remain as submerged fossils (like at Alborn),
others are currently above sea level, and still
others straddle both environments, such as
those found in Cap Blanc (Mallorca)16 or the
gulf of Antalya and the Taurus Mountains of
southwest Turkey.17 These are examples of
fossils from both sides of the Mediterranean.
During the Messinian salinity crisis, the Mediterranean Sea underwent one of its most
drastic changes, causing the ex tinction of
many species and the coral reefs.18 Later,
when the straits of Gibraltar opened again,
the water rushed in and filling the Mediterranean Sea once again, but the reefs did not
reform. Insead, other coral forms were generated representing a great diversity.
The actual origins of corals and reefs in the
Mediterranean go far back into geological
time. There were coral formations during the
Paleocene and Eocene19 epochs, and much
older coral formations have even been discovered that date back to the Triassic, when
the Mediterranean was part of the immense,
ancient sea of Tethys, more than 200 million
years ago. Some of these sites have been excavated at the Italian location of Zorzino.20
The oldest coral reef discovered to date is

located in the state of Vermont in the United
States. It is 450 million years old.21 However,
this reef does not really fall into the category
of a coral reef as per the current definitions,
because, although some corals contributed to
its formation, it is not principally the product
Mediterranean madreporaria (Cladocora caespitosa)

OCEANA/ Juan Cuetos
[16]
of corals. Neither does it present a pervasive

ecosystem, made up of complexly interrelating habitats and the multitude of organisms
which depends on and interacts with it. In fact,
as per the modern definition of a coral reef,
the oldest known example is located in the
Monte Bolca region of Italy22, belonging to the

Eocene epoch. Here were whole communities of fish, including the first known presence
of herbivorous ones, and other organisms living in a coral-built reef. For some observers,
a shif t occurred in the ecological structure of
[17]
coral reefs some time between the Mesozoic

and Cenozoic23, 65 million years ago, when
the Mediterranean was about to be cut of f
from the Indo-Pacific zone by the formation of
the Arabian Peninsula.
The Mediterranean coral reefs from the
Eocene and the marine life which they sheltered show a strong similarity with those that
have been found in the Indo-Pacific zone,
with an even closer resemblance between
some of their fish species than with the fish
populations in the tropical Atlantic zone.24
Furthermore, the madreporaria coral that is

endemic to the Mediterranean, Cladocora
caespitosa, has also lived in this sea for a
long period of time. It is known that some
reefs date back to the Pliocene and have lasted through the Pleistocene and Holocene up
to today31. However, todays reefs are only
a small portion of their original range. The
Mediterranean madreporaria is in a progressive state of regression.
Exclusive corals and

imported corals
Just as the Mediterranean seems to be one

of the areas from which modern tropical coral
reefs have originated, some deep-sea or cold
water reefs may also come from this sea.
The deep-sea coral reef fossils formed by
species such as Lophelia per tusa, Madrepora
oculata and Desmophyllum dianthus found in
the Mediterranean date back to the end of the
Pliocene and the early Pleistocene25 (1.8 million years ago), which makes them the oldest
ones found to date.26
Coinciding with the early Miocene, it appears

that many of these reefs suf fered a serious
regression and today, only a few zones in this
sea have been shown to bear live specimens,
such as those found in the Ionian Sea27, while
their distribution in the Atlantic covers both
margins of the northern hemisphere28.
Many scientists have mentioned the possibility that the in out flow waters from the Mediterranean (known by the acronym MOW -Mediterranean Out flow Water-), larvae of these
species then invaded the Atlantic waters, thus
propagating through the whole Ocean29. As
already indicated by Oceana30 and other authors, some of the principal coral reefs in the
deep waters of the Eastern Atlantic are in the
flow path of Mediterranean water as it passes
through the straits of Gibraltar.
[18]
Oculina patagonica OCEANA/ Juan Cuetos
However, the Mediterranean appears not

only to have exported species and reefs. In
the last decades, some species of non-native
corals have set tled in the Mediterranean. This
is the case of the orange striped anemone
(Haliplanella lineate), and possibly the colonial coral Oculina patagonica.
The colonial coral Oculina patagonica is traditionally considered to be a species that was
imported into the Mediterranean32.
The first specimens of Oculina patagonica

were found in 1966 in Savona (Gulf of Genoa,
Italy).33 They are today well represented
throughout the Mediterranean, especially in

areas between Italy, France and Spain,34 and
there is also a second pocket in the Eastern
Mediterranean, between Egypt, Israel, Lebanon35 and Turkey36.
At present there is a dispute among scientists
as to whether this species is foreign or not.
At the very least, doubt exists as to whether
it should be excluded from the list of Mediterranean species. The only data on ex tra-Mediterranean samples of this species come from
sub-fossil excavations in the south of Argentina37 (hence its name) of Tertiary strata. But,
there is neither a record of any live specimens
of this scleractinia nor of any testimony from
the annals of past centuries.
There are many questions to be made regarding this species. When and how was it
introduced into the Mediterranean? Is it a relic
from the Tertiary? Is Oculina patagonica really the specimen found in the Mediterranean?
Are there living colonies of Oculina patagonica to be found in South America but which
have yet to be discovered? Many of these
questions still go unanswered.
The orange striped anemone (Haliplanella

lineate) is originally from the Pacific and is
thought to have been introduced in the 19th38
century into European Atlantic waters on the
hulls of ships39. It was discovered for the first
time in Mediterranean waters of f Corsica in
197140.
The Mediterranean also shelters some endemic species of corals. Some of them are
typical of this sea, yet may be found in surrounding zones. This is the case of the red
gorgonia (Paramuricea clavata) which may
have traveled as far as the Berlengas Islands
of Portugal. As documented by Oceana, they
are to be found in the submarine mountains
of Gorringe41.
Other species that are endemic to the Mediterranean but which may also be found in surrounding areas are Leptogorgia sarmentosa,
Maasella edwardsi, Actinia striata, Astroydes
calycularis, Balanophylla europaea, Cribrinopsis crassa, Cladocora caespitosa, Phymanthus pulcher and Corallium rubrum.
Golden cup coral (Astroides calycularis) OCEANA/ Juan Cuetos
[19]
04.
[20]
Coral habitats and

corals as habitats
Bunodeopsis strumosa on Cymodocea nodosa

OCEANA/ Juan Cuetos
The forms in which corals congregate give

rise to the creation of dif ferent types of marine habitats, or else, participate in existing
ones. Among the most significant in the Mediterranean are the following:
Coral reefs
gonia. In the Atlantic, reefs have been found

ex tending over several dozen kilometers and
reaching up to 30 meters in height43, with an
ample range of depths from 40 to over 3,000
meters44.
As mentioned above, the Mediterranean lacks

large-scale reefs. Only the biotopes formed
by some species may reasonably be classified as such.
In the Mediterranean, only a few species

belong to the scleractinians or madreporarians, which is the type of coral that forms the
great reefs of the tropics. But the great majority of these specimens are only represented
by small colonies or in isolation (or as part
of reefs created by other species). Among
these, and in the infralit toral and circumlittoral waters, the Mediterranean madreporaria
(Cladocora caespitosa) stands out as a species ex tant only in the Mediterranean and adjacent Atlantic waters. It can create large colonies up to four meters in diameter, although
it is usually found in smaller colonies of a few
hundred polyps. In some areas of the Mediterranean, they can occupy significant zones.
This can be seen with the species found in the
dif ferent reefs of the western Mediterranean,
such as in the costal lagoon of Veliko jezero
in Croatia, which measure approximately 650
square meters and between four and 18 meters deep.42 Oculina patagonica can also form
dense colonies and occupy significant areas
of the sea bed with abundant sunlight.
Reefs formed by cold water or deep water
corals is another type of coral habitat existing in the Mediterranean, mainly formed by
tuf t coral (Lophelia per tusa) and madreporaria (Madrepora oculata). This type of reef
is also found in ex tensive areas of the North
Atlantic and can generate an ecosystem capable of clustering more than 800 dif ferent
species, including corals and deep-sea gor-
Mediterranean madreporaria (Cladocora caespitosa)

Today these deep-sea reefs are rare in the

Mediterranean. Most of the findings are of
fossil or sub-fossil corals, although some live
polyps have been found in dif ferent areas.
The most important of these is in Santa Maria
di Leuca (Italy) in the Ionian Sea, where both
species appeared together with other deepsea corals, such as Desmophyllum dianthus
and Stenocyathus vermiformis, in waters from
300 to 1,200 meters deep, occupying areas
as large as 400 square kilometers45. Other
concentrations of deep-sea corals were found
in the Palams and Cap de Creus canyons of
Spain.
In both types of reefs, the temperature and
availability of nutrition seem to be the limiting
factors in distribution. Salinity, topography,
the substrate of the sea floor and light intensity seem to be the variables for the species
zooxanthelae46, not to mention the impact of
[21]
04. Corals habitats and corals as habitats
human activities such as bot tom trawling. In

some areas, it has pushed back these formations to less accessible regions such as canyons and ravines.
For deep-sea corals, the hypothesis for measuring the set tlement and grow th of a reef or
colony is influenced by oceanographic and
paleo-environmental factors. It is thought that
the microbial role in hydrocarbon seepage
during carbonate formation facilitates their
grow th47.
Coralline algae
The predominant species which form this vital ecosystem are not corals at all, but as the
name indicates, a type of algae. However, one
of the most characteristic representatives of
these formations are gorgonias, which serve
(the environmental function) as large trees.

This ecosystem is typical in the Mediterranean48, although similar formations may be
found in other areas, including the Atlantic.
Some researchers have managed to dif ferentiate up to five types of coralline algae.49
All of them have corals present. Up to 44 species have been counted in this ecosystem50.
Among the most representative are the gorgonia (Paramuricea clavata, Eunicella cavolinii, E. singularis, E. verrucosa, etc.), other octocorallians like red coral (Corallium rubrum),
dead mans fingers (Alcyonium sp.), zoantharia such as the orange colonial anemone
(Parazoanthus a xinellae) and hexacorallia
such as the yellow coral (Leptosamnia pruvotii), the orange coral (Astroides calycularis),
etc.
Pink sea fan (Eunicella verrucosa) between Astroides calycuilaris and
red gorgonian (Paramuricea clavata) OCEANA/ Juan Cuetos
[ 22 ]
The contribution of gorgonians and corals to

coralline algae is significant. They fix nutrients
and sif t sediments51, producing calcium carbonate52 and generating a large biomass53, or
can also procure substrate for the set tlement
of epibionts54.
In this respect, the larger gorgonians are often colonized by a multitude of organisms
such as bryozoans (Pentapora fascialis or Turbicellepora avicularis), hydrozoans (Eundendrinum sp., Ser tularella sp.) or sponges (Dysidea sp., Hemimycale columella). They are
also useful as an egg depository for various
species including the Cat shark (Scyliorhinus
canicula), as Oceana and other researchers55
have been able to verif y. Eggs are deposited
in species such as Paramuricea clavata and
Eunicella cavolinii in Portofino (Italy) and in
other areas of the Mediterranean.
Large concentrations
of anemones
There are some anthozoans which are usually found in large colonies, like the jewel
anemone (Cor ynactis viridis) and the yellow
Trumpet anemone colony (Aiptasia mutabilis)

amongst sponges (Ircinia variabilis) OCEANA/ Juan Cuetos
cluster anemone (Parazoanthus a xinellae),

but there are also true anemones which although normally found in small groups or in
isolation, can occasionally form large colonies, giving rise to unique habitats. This is the
case of Anemonia sulcata and Aiptasia mutabilis found on Mediterranean facies.
In the case of Anemonia sulcata, Oceana

has documented large concentrations of this
species frequently in association with brown
algae forests, such as the kelps Saccorhiza
polyschides and Laminaria ochroeleuca.
Mediterranean snakelocks anemone (Anemonia sulcata)

OCEANA/ Juan Cuetos
[ 23 ]
Concentrations of anemones with these characteristics have already been documented by

other researchers on the sea floors of some
marine reserves, such as Columbretes56 or
Alborn57.
Although Aiptasia mutabilis is of ten found
alone or in small groups, it has also been
detected in large concentrations in the company of poriferaria such as Ircinia variabilis,
or in rocky areas between prairies of marine
seagrasses.
Some species are authentic specialists when

it comes to forming a dense cover of colonies
on vertical substrates. The jewel anemone
(Cor ynactis sp.) or yellow cluster anemone
(Parazoanthus a xinellae) can occupy widespread zones of submarine walls. But some
hexacorallias also occupy this strategic high
ground. Some of them form large colonies
such as the orange coral (Astroides calycularis) and star coral (Madracis pharensis),
while others form isolated colonies yet become quite numerous, such as the yellow
coral (Leptosamnia pruvotii) and dif ferent
dendrophylliidas and caryophylliidas. Some
coral species such as Polycyathus muelerae
or Phyllangia mouchezii can mantle entire
rocks.
Other hard substrates like gravel floors, small

rocks and stones, can also serve as set tlement locations. This is the choice of many
anemones such as Diadumene cincta, Anemonia melanaster, and Metridium senile58.
Some species have taken ma ximum advantage of the hard structures which exist on
the sea floor, whether it occurs naturally or
artificially. Small zoanthidas (Epizoantus sp.)
Phyllangia mouchezii OCEANA/ Juan Cuetos
On rocks, walls
and hard substrates
The capacity to adhere to dif ferent substrates
makes corals ef ficient colonizers, even on
very steep rocks and walls. Walls, the costal
shelf and high rocks are favorite places for
many corals species. These privileged outcrops are ideal for filtering plankton-filled water which usually gets swept upward in these
zones, or else are brought here by the currents.
[ 24 ]
Eunicella cavolini, red gorgonian (Paramuricea clavata)

and Leptosamnia pruvoti in a cave OCEANA/ Carlos Surez
can colonize almost any substrate which they

find: stones, shells, coral skeletons, bot tles,
fishing lines, cables, etc.
In caves and fissures
As far as we know today, there are no anthozoans that live exclusively in caves, although
some have become more specialized in that
type of habitat. Among these are many species of dendrophylliidas, given that the majority of species belonging to this genus does
not coexist with any symbiotic algae and
therefore they do not need light. One of the
dark-water corals is the Southern cup coral
(Car yophyllia infornata) which is frequently
found in caves and under low lying overhangs59. Other species of ten associated with
walls and rocks, such as Polycyathus muellerae, Parazoanthus a xinellae or Leptosamnia
pruvoti, are also quite common in this type of
habitat60.
Not all corals like sunlight. Some of them look

for caves or hollows in which to grow. Some
species have become specialized in shady
habitats while others have added them to
an already ample habitat range, and others
still have retreated to them for lack of choice,
forced by over-exploitation of more exposed
or accessible zones.
Overgrow th on Elisella paraplexauroides
OCEANA/ Juan Cuetos
Polycyathus muellerae OCEANA/ Juan Cuetos
However, many corrals can simultaneously

exist in dark zones such as caves and fissures as well as in open zones. The Halcampoides purpurea61 anemone lives in caves, as
well as on sandy floors and on gravel. Many
daisy anemones, such as Sagar tia elegans or
S. troglody tes are not exclusive tube-dwellers of such habitats, but it is common to find
them in holes as well. One species which
used to be found in wider ranging areas has
been found limited to caves, fissures and low
lying overhangs in many areas where it is
commonly found. This is the red coral (Corallium rubrum), which has been intensively harvested in more exposed areas that are easily
accessible to fishermen, divers and robots.
[ 25 ]
even form dense colonies. In some Atlantic

European waters, Virgularia mirabilis densities62 of up to 10 individuals per square meter
have been found, making these areas virtual
sea feather forests.
Golden anemone (Condylactis aurantiaca) OCEANA/ Juan Cuetos
On muddy and sandy floors

Desert sea floors, or floors composed of fine
sediment such as sand and mud, are usually more challenging places for anthozoans
to set tle on. However, a few species have
colonized this terrain in which the colonization strategy must take into account the sof tness of the substrate. It of ten shif ts, requiring
adaptive anchoring systems or complete alternatives altogether.
Some species found on such floors are highly
specialized for this habitat. This is the case of
the sea feathers, Funiculina quadrangularis,
Virgularia mirabilis, Pennatula spp., Kophobelemnon stelliferum, etc. They are commonly
found in marine deserts and sometimes may
Many species bury part of their skeleton in

the sof t substrate in order to anchor themselves, and might even retract their bodies
entirely inside the sand or mud. This can be
seen with some sea feathers and the anemone Anemonactis mazeli. Cerianthus membranaceus can submerge its tube one meter
into the substrate with only the tentacles of its
superior end showing through63. Oceana was
able to verif y the presence of this species in
mixed communities of crinoideas (Leptometra phallangium) and sea urchin (St ylocidaris
af finis) in sandy floors below 80 meters.
Another species which is characteristic of
these floors, but only at depths of 200-300
meters, is Isidella elongata, which is an ideal
habitat for crustaceans.
The deep-water anemone (Condylactis aurantiaca) only appears on sandy bot toms64,
but it does not delve very far into them, preferring to stay close to sandy areas abut ting
rock zones. Other anemones which have
similar preferences are Cereus pedunculatus, Peachia cylindrica, and Andresia par tenopea.
There are also corrals which simply sit on the

surface of the substrate without anchoring at
all, as with Sphenotrochus andrewianus, an
infralit toral and circalit toral species.
Hermit crab (Dardanus sp.) with Calliactis parasitica

OCEANA/ Juan Cuetos
[ 26 ]
earic Islands is an example of this: It is one of

the zones poorest in nutrients, and therefore
the least turbid. Red gorgonian are not seen
above 30 meters68, while in the Gulf of Lion
and the Ligurian Sea it is not uncommon to
observe them above 15 meters deep.
Dead mans fingers (Alcyonium palmatum) OCEANA/ Juan Cuetos
In shallow waters and

at great depths
Very restricted luminosity or even with absolute lack of luminosity is sought af ter by some
species. Excluding those species which live
in symbiosis with zooxanthellas, corals are
sciaphilic animals: that is, they prefer shady
areas and even those with absence of light.
Even though the most well-known species are
found in surface waters, most of the worlds
corals grow below the photic zone or optimal light filtration zone. In fact, two-thirds of
all known coral species live in dark and cold
zones65. Deep-sea corals can live at depths of
up to 6,000 meters, although the majority live
between 500 and 2,000 meters66.
The Mediterranean is a good example of this
phobia to light. The oligotrophia of Mediterranean waters shows that species populating
shallow areas with greater turbidity do not
start to grow until depths of over 30 or 40 meters, where light has been suf ficiently filtered
by the water column. Even in the Mediterranean, the distribution of species such as the red
gorgonian (Paramuricea clavata) is strongly
influenced by the waters clarity67. The Bal-
We know that some species can even spend

part of their day out of the water in inter-tidal
zones or in tide pools.69 This is the case of the
beadlet anemone (Actinia equina). To avoid
desiccation and the ef fects of high salinity,
they retract their tentacles to reduce exposure to the air, while simultaneously storing
water reserves.
For species composed of 80%-90% water,
and as such with a low gast content70, the
challenge to resist pressure at great ocean
depths is minor. This allows them to colonize
sea abysses at depths greater than 3,000 meters.
However, there are species that seek light, in

many cases so that their resident zooxanthellae can photosynthesize it. Cladocora caespitosa is mostly found in light-filled or slightly
shaded areas, although some colonies have
also been found in deep water. The ef fects of
higher temperatures have also been noted.
It increases their calcification, so that they
usually seek water at temperatures between
11C and 25C, but if they are in temperatures
above 28 degrees they lose their zooxanthela and turn white71. But not all corals prefer
warm waters. Temperature, as we shall see
in greater detail, is also a limiting factor for
certain species of corals. Deep-sea species
such as Lophelia per tusa, prefer water below
13C72, which means Mediterranean depths
of 250-300 meters or more.
[ 27 ]
Dirty water
Dirty water is not a problem for some species
of corals. While pollution and turbidity can
be fatal for many species of anemones, corals and gorgonians, some anthozoans have
become specialists in adapting to damaged
ecosystems, such as those found in marine
ports or other places with unclean water. This
is the case of some anemones such as Diadumene cincta and Aiptasia diaphana.
Scleractinian corals are of ten the most demanding regarding environmental conditions. However, there are some that have occupied niche habitats in polluted areas, such
as Oculina patagonica, frequently seen in
commercial ports.
Some corals cluster near locations of anthropogenic spills or runof fs, such as the dead
mans fingers (Alcyonium sp.). Species which
are not too picky can find an important source
of nutrition from the out flow of residual waters, which have a high density of suspended
mat ter.
The ease with which they adapt to polluted
water has made it possible for the orange
striped anemone (Haliplanella lineate) to occupy many coastal areas of the Mediterranean. Their arrival in the waters of many European countries has been propagated by
ports73.
Bunodeopsis strumosa on Cymodocea nodosa

OCEANA/ Juan Cuetos
to being table companions. However, some

have managed a more specialized and symbiotic relationships.
On algae and seagrasses
Dif ferent anemones, usually smaller ones,

have adapted to the substrate af forded them
by marine plants and algae in order to settle. Species such as Bunodeopsis strumosa
are of ten found on the leaves of marine seagrasses such as Posidonia oceanica or Cymodocea nodosa. Paranemonia cinerea and
Paractinia striata have similar tastes. Actinia
striata is more common on seagrasses of the
Zostera genus.
Others of ten set tle on algae, such as Gonactinia prolifera. Some can grow to a large size,
like Anemonia sulcata, which manages to anchor on large kelp.
Living on the backs of others
The inherent plasticity of anthozoans has not

limited their distribution to particular physical habitats, such as rocks, sands, mud or
biological debris (shells, coral skeletons or
bryozoans) or non-organic substrates. Many
species have preferred to live on live organisms such as seagrass, algae or animals. In
some instances, the relation between anthozoan and host is an opportunistic and limited
[28]
Mediterranean snakelocks anemone (Anemonia sulcata)

on Furbellows kelp Saccorhiza polyschides OCEANA/ Juan Cuetos
Gobius xantocephalus under a colony of Epizoanthus arenaceus OCEANA/ Juan Cuetos
On living creatures
Some look for means of transportation to
carry them to new sources of food and in
exchange lend their irritating tentacles as a
form of defense for their host. The most common anemones in this type of trade are the
parasitic anemone (Calliactis parasitica), the
cloak anemone (Adamsia carciniopados) and
Hormathia alba, which are all common on
snail shells inhabited by hermit crabs (genus
Pagurus and Dardanus)74.
Oculina patagonica OCEANA/ Juan Cuetos
When it comes to giving a ride to species

such as A. carcinopados and H. alba, there is
usually one on the back of each hermit crab,
although some species, such as C. parasitica,
are limited by the surface area of the hermit
crabs shell and the capacity of the crab to
drag this weight across the sea floor. Thus,
it is not rare to see two, three, or even more
anemones on the shells of these animals.
There are also many examples of anthozoans
looking for an anchor point from which to optimize access to food sources suspended in
Orange cluster anemone (Parazoanthus a xinellae) growing

on a sponge from the A xinella genus OCEANA/ Juan Cuetos
[29]
the water, without having to go from one to

place to another; the entire bet ter if the perch
is high up. This is the vantage point of the yellow colonial anemone (Parazoanthus a xinellae) which of ten seeks the structures of fered
by branching sponges of the A xinella75 genus.
Other anemones have chosen the familiarity
of cnidarians, like Amphiantus dohrni, which
prefers the knoll of fered by the large hydrozoan, or some gorgonians such as Eunicella
verrucosa or Isidella elongate76. Protanthea
simplex seeks its fortune on reefs of Lophelia
per tusa77.
In company: corals and

symbiotic/commensal animals
The anthozoans have created wide variety of

relationships between species. Some of the
more well-knows ones, mentioned above,
are those maintained with hermit crabs, or
the close sharing between various corals and
zooxanthellae algae, a type of dinoflagellate
which depends on the light intensity and water
temperature. Zooxanthellas can be found on

the Mediterranean madreporaria (Cladocora
caespitosa)78, the snakelocks anemone (Anemonia viridis)79, the Paranemonia cinerea80,
in star coral (Madracis pharensis)81, and the
Mediterranean cup coral (Balanophyllia europaea)82. Some symbiotic algae have also
been found on Oculina patagonica, including
some endoliths of the Ostreobium genus in
the corals skeleton83. These microalgae fulfill other vital functions such as protecting the
corals from bleaching and ultraviolet radiation84.
There are dif ferent species of small crustaceans that of ten live in association with
anemones, like Telmatactis cricoides. In studies carried out on specimens from the Atlantic
Madeira and Canary Islands -which are of ten
larger than Mediterranean specimens- it was
found that 86% lived symbiotically with an average of 2-3 shrimps, usually of the Thor amboinensis85 species.
Phyllangia mouchezii catching a mauve stinger jelly fish (Pelagia noctiluca) OCEANA/ Juan Cuetos
[30]
In the Mediterranean, the shrimp that most often live in symbiosis with anemones are those
of the Periclimenes genus, such as P. sagit tifer
or P. amethysteus. They of ten live this way on
large specimens of Anemonia sulcata, Aiptasia mutabilis, Cribrinopsis crassa and Condylactis aurantiaca. It is not strange either to see
some mysidaceas, such as Leptomysis lingvura nestled between the tentacles of these
anemones.
Other crustaceans of ten live in association with corals, such as the cirripedes balanomorphas. Megatrema anglicum is of ten
found on club coral (Car yophyllia smithii),86
yellow coral (Leptopsammia pruvoti),87 Hoplangia durotrix or corals of the Dendrophyllia
and Balanophyllia genus.
Anemonia sulcata is the queen of specific

inter-species relationships. In addition to
the species mentioned above, this one can
shelter other crustaceans between its tentacles, such as the hairy crab (Pilumnus hir tellus), the lesser spider crab (Maja crispata),
or the spider crab (Inachus phalangium).88 In
the lat ter case, it prefers the more stationary
females.89 It is also the species associated
with the only anemone fish of the Mediterranean: the anemone clown fish (Gobius bucchichii).90 This Mediterranean native makes it
dif ferent from all other clown fish in the world
that belong to the Pomacentridae family, not
present in the Mediterranean.
However, in some cases roles get reversed
and it is the anthozoan which is parisitic on
other species or even on other anthozoans,
as indicated previously with Parazoanthus
a xinellae on sponges (A xinella sp.), Epizoanthus arenaceus, Calliactis parasitica, Adamsia
palliata and Hormathia alba on hermit crabs,
and Parer y thropodium coralloides and Gerardia savaglia on gorgonians.
Ascidian (Pycnoclavella taureanensis)

on Elisella paraplexauroides OCEANA/ Juan Cuetos
[31]
05.
[ 32 ]
Coral reproduction
Gerardia savaglia on red gorgonian (Paramuricea clavata)

OCEANA/ Carlos Surez
The various ways in which anthozoans reproduce has long been used as a method for
classif ying them, but recent studies91 show
that thee systems which some species use
to reproduce depend to a large ex tent on the
characteristics of the colonies and the environmental conditions to which they are subjected.
As such, the same species may use dif ferent

methods of reproduction, although there may
be a single way that is more characteristic for
some speices, or at least one observed as
such in laboratories or the natural environment. This is a topic which has generated
much debate and controversy as well as numerous scientific studies.
Using the current knowledge we have on this
mat ter, we can make the following observations:
Sexual and
asexual reproduction
Sexual reproduction, that is, reproduction

which requires the participation of males and
females for the production of eggs (oocy tes)
and sperm, has always been considered the
most normal way for anthozoans. In addition
to oviparous fertility, viviparous has also been
noted, whereby the female coral becomes
pregnant through internal fertilization of the
ovocy te resulting in polyps which are then
expulsed or delivered into the environment.
Asexual or vegetative reproduction, on the

other hand, is carried out by an isolated polyp
and new individuals are created without the
participation of the other gender. In this way,
colonies (or species) arise with only one sex
present. Furthermore, asexual reproduction
can take various forms. There are descriptions of budding, transversal and longitudinal
fission, basal laceration, fragmentation, en-
cystment, and some more typical methods

of sexual reproduction, whether oviparous or
viviparous, through parthenogenesis.
Some species which appear to primarily use

sexual reproduction can also select the vegetative mode under conditions of stress or
as an opportunistic preference, or due to the
isolation of the colonies or polyps.
Synchronized reproduction
One of the most spectacular methods of sexual reproduction among corals is synchronized reproduction. This is characterized by
the synchronization of various colonies of
corals (sometimes, from dif ferent species),
expulsing their sperm and eggs at the same
time. This strategy is very common among
scleractinians in coral reefs92, and has also
been seen among zoanthias93 and even octocorallias94.
With mass production of male and female
gametes over a short period of time, the
chances that conception will be successful increases and along with the possibility
that at least a certain percentage will escape
predators.
Red gorgonian (Paramuricea clavata) and yellow cluster anemone

(Parazoanthus a xinellae) OCEANA/ Carlos Surez
[ 33 ]
Moreover, as researchers have observed,

some species have demonstrated strategies
that increase the chances that their gametes
will fuse, by means of a chemical substance
which at tracts semen.95
In the Mediterranean this type of reproduction
exists among certain gorgonians, like Paramuricea clavata96, and among madreporarian
coral colonies such as Oculina patagonica.
In the case of the red gorgonian (P. clavata),

the synchronization sessions are not unique
or concentrated (that is, not all colonies participate in reproduction at the same time), but
rather they repeat themselves over several
months in spring and summer, three to six
days af ter the full or new moon97.
It is believed that these time periods may give

rise to hybrids.98 In tropical reefs, some cases
have been observed among the Acropora99
and Montanstraea100 genus. In the Mediterranean, no such occurrence has yet to be
documented.
Daisy anemone (Cereus pedunculatus) OCEANA/ Juan Cuetos
[ 34 ]
Oviparous, viviparous
Depending on the way in which corals reproduce, they can create complete new polyps
or intermediary phases. Both oviparous and
viviparous reproduction, be it sexual or asexual, has been widely documented in various
anthozoans.
In sexual reproduction, the generation of a

new lineage may occur by means of eggs
and their subsequent fertilization, or else, by
fertilization of a female ovocy te in the interior
of the female polyp of a coral.
In the case of oviparous reproduction, the

eggs are fertilized outside the coral. In some
species, they are expulsed outward, while
in others they remain stuck to the colony
by means of mucus produced by the polyps. They then mature there until the larvae
emerge and set tle in their surroundings101.
In most cases, oviparous and viviparous reproduction are the result of sexual reproduction, but they have also been found to occur
asexually through parthenogenesis102. This

sometimes occurs with the red anemone or
Beadlet anemone (Actinia equina), which is
found in the Mediterranean and the Atlantic103.
Budding, fission and laceration
As we have mentioned above, asexual reproduction can give rise to dif ferent means of
generation of new polyps. One method which
has been widely studied is budding. This
consists in the production of a bud or yolk by
the female polyp, which grows until another
polyp sprouts which either takes its place in
the coral that produced it or is expulsed at
such a distance that it finds a new location
to set tle. Although this type of reproduction
is more common among cnidarians, such as
hydrozoans or scyphozoans, it has also been
reported for various anthozoans.104
Fission occurs when a polyp gives rise to a

new polyp. This may occur by transversal or
longitudinal fission, depending on whether
the polyp buds laterally or on top of the other
one. Transversal fission is more common in
other cnidarians, although it may occur in anthozoans as well, but this is thought to be exceptional and nearly always provoked under
conditions of stress.105 Lateral fission is more
common among corals and in fact has been
found in both octocorallians106 and hexacorallians107, such as the orange striped anemone
(Haliplanella lineate)108 which has invaded the
Mediterranean.
There is another of asexual reproduction
which has been found to exist, at least for
some anemones109 and black corals.110 This
is fragmentation- the generation of new polyps from a piece of coral, usually a tentacle.
Egg, larva/planula
and polyp phases
Although asexual reproduction may give rise
to new polyps, the normal process for many
anthozoans is to pass through dif ferent phases until reaching the adult state.
Solitary coral (Balanophyllia europaea) OCEANA/ Carlos Surez
When those which reproduce with eggs hatch,

a ciliated larva appears. This is a planula,
and for a short period of time (normally a few
days) it lives like plankton until it once again
anchors to the substrate and gives rise to a
new polyp or initiates a colony. What makes
it dif ferent from all other cnidarians is that it
never exists in the medusa phase.
Planulas may be found in the water af ter newly expulsed eggs that were fertilized by sperm
are hatched. They may also come directly
from a polyp which was fertilized internally
and which instead of liberating an egg, waits
until it turns into a planula. In other words, the
incubation of the eggs may dif fer from one
species to another. While for red coral it is internal,111 for red gorgonian it is ex ternal.112
[ 35 ]
White gorgonian can produce an average of

four eggs per polyp, which means that a colony usually disperses an average of around
6000 planulas113 into the water. Red coral has
a very low reproduction rate of one planula
per polyp, on average.114
In some species the larva have negative floatability to avoid being swept away by currents
and to allow them to set tle in the vicinity of
the polyp or mother colony, as for example
the Mediterranean cup coral (Balanophylla
europaea).115
Hermaphroditism can define an entire colony

when polyps of both sexes are present, as in
the case of the orange coral (Astroides calycularis)122, endemic to the Mediterranean. It can
also be individual, when a polyp has both the
male and female gamete, as with the colonial
coral Madracis phaerensis123 and another endemic coral of the Mediterranean, the solitary
This negative floatability has also been found

in the eggs of zoantharia (Epizoanthus sp.),
found in the South Pacific at depths between
80 and 200 meters.
Separate sexes
and hermaphrodites
The separation of sexes, that is, the existence

of both males and females (gonochorism), is
common among corals, not only among individual polyps but also between whole colonies.
In many species in the Mediterranean, the sexes are usually separate. Thus, while a colony
is usually composed of exclusively male polyps, another will be exclusively female. This
characteristic has been documented in many
octocorallian gorgonaceas, such as Eunicella
verrucosa116, Eunicella singularis117, Paramuricea clavata118, and Corallium rubrum119, and
also in hexacorallians and deep-water species (Enallopsammia rostrata, Madrepora oculata or Lophelia per tusa)120.
Hermaphroditism is another reproductive
strategy of some anthozoans. It is mainly
seen in tropical corals of the Pocilloporidae121
family, although it is not an exclusive trait.
[ 36 ]
Parer y thropodium coralloides OCEANA/ Juan Cuetos
cup coral Balanophyllia europaea124 (this is a

rare trait among dendrophyllidas and unique
to this genus).
And of course, in the great diversity of corals morphology, there are species which may
have both gonochoric and hermaphrodite
characteristics. Hermaphroditism is less com-
mon among octocorallians. However, some

hermaphrodite colonies have been found
among coral colonies of Carijoa riisei studied
in the Pacific125 and among the Indo-Pacific
gorgonias Heteroxenia sp.126, as well as some
Mediterranean colonies of Pareritropodium
coralloides127, among others.
[ 37 ]
06.
The fight for survival
[38]
Cerianthus sp. OCEANA/ Juan Cuetos
Of great importance for the development of

corals is the energy expended on reproduction, as it can provoke a decrease in grow th
rates128. In addition, the fight for space with
other anthozoans can consume much energy. As such, bigger colonies can bet ter distribute tasks and compete for survival and
expansion129.
Trouble with the neighbours:

space
In the zone where dif ferent species of anthozoans, or even colonies of the same
species, are found, there is usually a bat tleground where polyps that are on the front
line modif y their morphology to bet ter fight
the competitor. For example, they may develop specialized tentacles130. In fact, corals
and anemones have dif ferent types of tentacles131: feeder tentacles; tentacles for capturing plankton and suspended particles;
feeler tentacles to find out whether there are
enemies lurking nearby. The lat ter may be up

to 10 times longer than the feeder tentacles.
Hunting tentacles are those which are deployed for combat.
These tentacles are present in Mediterranean species, such as the plumose anemone
(Metridium senile)132 or in the invading orange
striped anemone (Haliplanella lineate)133.
In the fight for survival, mortality rate and regeneration can play an important role. The ca-
Corals and sponges competing for space OCEANA/ Juan Cuetos
pacity of anthozoans to recover af ter sustaining wounds varies widely. While some species
need centuries to rebuild a reef or colony that
has been damaged, others regenerate much
more quickly. However, the general trend reflects the long-living and slow-growing nature
of the species, and damage, be it natural or
anthropogenic, is usually a significant challenge.
[39]
Competition for space is also found among

other marine organisms. When this happens,
dif ferent species of hydrozoans, bryozoans,
etc. colonize the coral structures and thereby
provoke their death by covering it. For species
such as Paramuricea clavata, death caused
by process represents half of the natural mortality rate of the species134. For Eunicella cavolini, it is the main cause of mortality, along
with uprooting135.
Growth
Corals must deal with various conditions in

order to grow and survive in a marine environment,
One of the determining factors on coral

grow th is the availability of food. This may
be favorable in zones with strong currents or
where dif ferent water masses mix together.
Low levels of sedimentation and turbidity also

play an important role.
Most colonies barely grow a few millimeters

a year. Such a slow grow th rate is especially common in those species which need to
grow a calcareous skeleton or create large,
branching masses. The longevity of anemones can be quite significant as well.
Unfortunately, lit tle is known about the life of
anthozoans, and their grow th rate and longevity remains a mystery for a great number
of species. However, present data seems to
corroborate the estimates regarding longevity. For example, the yellow coral (Leptosamnia
pruvoti), a colonial scleractinian with a calcium
carbonate skeleton, and gorgonian anemone
(Amphianthus dohrni), a solitary hexacorallia
without a skeleton, have estimated life spans
of between 20 and 100 years136.
Examples of grow th and longevity studied in some anthozoans137
[40]
Size matters
There is a correlation between size and the
age of anthozoans, as well as the nature
and ex tent of damages they may suf fer143.
Larger specimens are more vulnerable to incurring serious damage, although they also
show bet ter resiliency in regenerating, while
younger specimens have a limited or absent
response, although damage may be lesser.
This is why the size of colonies is an important factor in the survival of species.
Damage suf fered by the colony also brings
to light one of the main causes of mortality among anthozoans: excessive grow th of
epibionts144.
Leptosamnia pruvoti and Hoplangia durothrix
OCEANA/ Juan Cuetos
The false black coral (Gerardia savaglia)

might possibly be the longest living colonies
in the Mediterranean. It is the only zoantharia
that can produce a skeleton. In the Pacific,
colonies of other species of the genus Gerardia might reach 2,700 years of age.138 In
the Mediterranean, it is believed that some G.
savaglia formations might be more than one
thousand years old.
There are specific techniques used to measure the grow th of corals. In older colonies,
dating can be done with carbon (Ca14) or
other representative isotopes139, such as lead
(Pb210), radium (Ra226), thorium (Th230) or uranium (U234). For younger animals, the rate of
the increase in the thickness and vertical expansion of a polyp or colony can be monitored. Some of the most ef ficient methods for
some species are similar to those used for
trees: the rings that form in the skeleton of
anthozoans are counted. This has been carried out successfully with sea feathers140, gorgonians141 and scleractinian corals142.
Size is not only important for corals or a

colony, but it is also a significant factor for a
population as a whole. With species such as
P. clavata, which depend on synchronized reproduction involving numerous colonies, the
larger the population, the larger the chances
that new colonies will be generated and that
the species will survive. The mass mortality
suf fered by this species in recent years might
endanger the survival of this characteristic
Mediterranean species.
Overgrow th of algae, hydrozoans and sponges on red gorgonian

(Paramuricea clavata) OCEANA/ Carlos Surez
[41]
A seat at the table:

coral nutrition
Most anthozoans are suspension feeders.
They ex tend their tentacles and wait for small
organisms and particles floating in the water
to touch them. Their cnidos then shoot out
to catch the nutrients which are brought into
their oral disc and subsequently introduced
into their gastrovascular sac. The other common form of coral feeding is the use of a mucus layer which particles in suspension adhere to, and are then brought to the digestive
tract with the help of cilia.145
For some corals, a large part of their nutrition comes via their symbiotic relationship
with small algae living in their gastrodermis.
In most cases, these algae are dinoflagellates
(zooxanthellas), although green algae have
sometimes been found (zoochlorellas).146
Dead mans fingers (Alcyonium acaule)

[ 42 ]
Another method is to absorb the cells of organic mat ter dissolved in the water directly
through the ectoderm147.
In the case of sof t corals, such as dead mans
fingers, plankton is caught through absorbing water and filtering it like sponges.
Most corals are carnivorous. They feed on

zooplankton, although they can also combine
this nutrient with small algae or even bacteria.
Polyps can also catch prey of even greater
size, such as jelly fish. Anemones may even
try to catch crustaceans and fish.
Crustaceans are of ten an important part of
the diet of many anthozoans, although what is
actually ingested can vary greatly according
to the species. Tuf t coral (Lophelia per tusa)
has been observed catching cumaceans and
copepods148, while Mediterranean snakelocks
reduced the number of predators preying on

them, some species have become immune to
their venom and have specialized in consuming them.
Red coral (Corallium rubrum) OCEANA/ Carlos Surez
anemones (Anemonia sulcata) and Cereus

pedunculatus feed on amphipods and decapods. In the case of Actinia equina, their preferred food tends to be detritus149, while the
red gorgonian (Paramuricea clavata) ingests
eggs and larva150.
Guess whos coming

to dinner:
natural predators of corals
The cycle of nature some eat and others are

eaten. Corals also form part of the diet of certain animals. While the presence of stinging
cells and toxic substances in their bodies has
In the Mediterranean, there are not any fish

which consume corals, unlike in coral reef
where it is quite common. Here, the main
consumers of corals are mollusks, although
there are also some arthropods and worms
that choose to feed on polyps.
The most common coral-eating mollusks are
snails, especially ovulidas and coralliophilas.
As with Caribbean ovulidae such as the flamingo tongue cowrie (Cyphoma gibbossum),
they feed on gorgonians. Mediterranean mollusks have become specialists of this type
of octocorallian. One of the more common
species is Neosimnia spelta which devours
polyps and live tissue of gorgonians151 such
as Eunicella verrucosa, Eunicella singularis
and Leptogorgia sarmentosa. Other species
of ovulidae in the Mediterranean are Pseudosimnia carnea, Simnia nicaeensis, Simnia
purpurea, Aperiovula adriatica, Aperiovula
bellocqae, Globovula cavanaghi and Pedicularia sicula.
Spinimuricea atlantica
OCEANA/ Juan Cuetos
[ 43 ]
Regarding the coralliophilae snails, most are

Indo-Pacific species, but there are at least
a dozen in the Mediterranean as well152, including Coralliophila meyendor f fi and Babelomurex cariniferus153, which of ten feed on
the Cladcora caespitosa polyp. Other Mediterranean species are Babelomurex babelis,
Coralliophila brevis, Coralliophila sofiae, and
Coralliophila squamosa.
Other mollusks which have also been observed feeding on corals are the epitoniidae
snails, such as Epitonium dendrophyllidae,
which consumes orange corals (Astroides
calycularis)154, and some nudibranches, such
as Okenia elegans, which can feed on Paramuricea clavata155.
Flamingo tongue cowrie (Cyphoma gibbosum) on black sea rod

(Plexaura homomalla) in the Caribbean OCEANA/ Houssine Kaddachi
The bearded fireworm (Hermodice carunculata) is a polychaete which is found in wide

ranging areas of warm and temperate Atlantic waters156, and can reach lengths of up to
40 cm. Although their diet consists mostly of
carrion and decomposed mat ter (saprophagous), it also devours coral polyps. In the
Caribbean, it usually feeds on dif ferent species of gorgonians157 and hydrozoans158. In
the Mediterranean, its source of nutrition is
usually Oculina patagonica159.
[ 44 ]
Pycnogonids (Pycnogonum lit torale), or sea

spiders, feed on diverse actinias160, including
the plumose anemone (Metridium senile)161.
Red coral may suf fer at tacks from mollusks
and crustaceans, such as Pseudosmnia
carnea and Balssia gasti, respectively162.
Finallly, in more recent times, human beings
have also added anthozoans to their diet. In
some areas of the Mediterranean, some species of anemone are sold under the generic
name sea urchin or urchin for certain gastronomic dishes.
Corals with light
One of the adaptive advantages of some marine species is the possibility of successfully
surviving in certain habitats by carrying their
own lantern. The luminescence they create
may be used to at tract prey, avoid predators, communicate, and search for a mate,
as well as other functions that are still being
researched.
Although for many animals bioluminescence

is due to the presence of bacteria such as
Vibrio fischeri or the existence of specialized
cells, corals owe this feat to the presence of
luciferin and the enzyme luciferase, which
catalyzes its own oxidation and produces
light.163 Although, like in other cnidarians, it is
possible that some species do not need luciferase for their bioluminescence and instead
produce a photoprotein (coelenterazine),
which reacts in the presence of Ca2+ to produce the chemical reaction.164
The pennatulaceas are the octocorallians that
have been shown to demonstrate bioluminescence most of ten. In the Mediterranean,
this has been found in Cavernularia pusilla,
Veretillun cynomorium, Funiculina quadrangularis, Virgularia mirabilis, Pennatula rubra,
Pennatula phosphorea and Pteroides spinosum.165
Outside of sea feathers, bioluminescence is

quite rare among octocorallians. It has only
been observed in a single genus of Pacific
alcyonidae (Eleutherobia sp.166) and in a few
species of gorgonians of the Isididae family,167 such as Lepidisis olapa, Keratoisis sp.,
Primnoisis sp., o Isidella elongata. The lat ter
may also be found in the Mediterranean.
The only species of anemone which has been
found to have bioluminescent properties is
the Hormathia alba168, a common species in
the Mediterranean and in some areas of the
Atlantic. This species which can also live on
the shells of hermit crabs.
In other species, bioluminescence is exchanged for phosphorescence169, as with
Mediterranean snakelocks anemone (Anemonia sulcata).
Corals that move

Most anthozoans are anchored to the substrate on which they live and stay in the same
spot for their entire lifespan. Only during the
larval stage do the ciliated planula of some
coral and gorgonian species move freely in
the sea.
Others, such as the Paranemonia cinerea171,

can migrate by moving vertically to new locations according to the season or to hibernate
on the sea floor among plant detritus, awaiting a rise in the temperature for them to climb
onto the leaves of marine seagrasses.
Others, such as Telmatactis forskali172, can
swim short distances using their tentacles as
fins. This behavior has also been observed in
Bunodeopsis strumosa, especially when they
fall from a seagrass blade. This behaviour
has also been observed in anemones of the
genus Boloceroides and Gonactinia173.
Still others are crawlers that use their tentacles to creep along the sea floor, covering several meters in search of a new place
to perch themselves. Some studies174 have
shown that tube anemones of the genus Cerianthus y Pachycerianthus can move across
the sediment. This behavior has also been
observed in sea feathers af ter they have
been torn from their original anchorage point,
dragging themselves across the sea bed until
they borrow in once again, or swelling up and
let ting the current carry their bodies to a new
location174.
However, some species of anemones or sea

feathers can change their location in search
of a bet ter one. Or, with the help of a symbiont, they may travel great distances, as with
the anemone that lives on the shells of the
hermit crab.
Some of them can slowly move away from
areas that have become hazardous to their
survival, as with Actinia equina, Anemonia
Sulcata, Anthopleura ballii and Sagar tia sp.,
which can move a few centimeters with the
help of their basal disk to avoid exposure following low tide, to flee an enemy or to avoid
a light source170.
Cerianthus sp. OCEANA/ Juan Cuetos
[ 45 ]
07.
[ 46 ]
Threats to corals
Ovulid on gorgonian OCEANA/ Juan Cuetos
Coral diseases
At least 18 diseases have been identified
around the that can have a massive ef fect on
corals.176
Diseases detected in corals
ASP = Aspergillosis
DSD = Dark Spots Disease
PLS = Pink Line Syndrome
SEB = Skeletal Eroding Band
VCB = Vibrio corallilyticus Bleaching
WBD I = White Band type 1
WPD = White Pox Disease
WPL II = White Plague type 2
YBD = Yellow Band
BBD = Black Band Disease
FPS = Fungi-protozoo Syndrome
SDR = Shut Down Reaction
SKA = Skeletal Anomalies
VSB = Vibro shiloi Bleaching
WBD II = White Band type 2
WPL I = White Plague type 1
WPL III = White Plague type 3
In many cases, mortality is caused by an increase in water temperature or abnormally

long periods of high temperatures. In some
cases, the pathogen responsible for mortality has not been identified, but in at least 10
of the infections, marine and land fungi, cyanobacteria, bacteria, protozoa, nematodes,
algae and crustaceans have been the cause
of some of the documented symptoms.
In aspergiliosis (ASP), the infectious agent
is the land fungus Aspergillus sidow yi177,
which mainly af fects gorgonians of the genus Gorgonia. In black band disease (BBD),
a multitude of microorganisms such as cyanobacteria178 Phormidium corally ticum and
Trichodesmium spp., Cyanobacterium sp.,
bacteria179 (Desulvovibrio spp., Beggiatoa
spp.) and marine fungi180 have been detected.
In the Fungo-protozoico Syndrome (FPS), it
Black Band Disease (BBD) in a tropical coral

(Siderastrea sp.) OCEANA/ Houssine Kaddachi
is fungi (Trichoderma spp., Clodosporium

spp., Penicillum spp. and Humicola spp.) and
protozoa181. Pink Link Line Syndrome (PLS)
has been linked to cyanobacteria182 (Phormidium valderianum). Skeletal Eroding Band
(SEB) is at tributed to a protozoan183 (Halofolliculina corallasia). For Skeletal Anomaly Syndrome (SK A), land fungi184 such as (Aspergillus sydowii) and endolticos185 have been
shown to play a part, in addition to algae186
(Entocladia endozoica and the Syphonals),
nematodes187 (Podocot yloides stenometra)
and crustaceans188 (Petrarca madreporae).
Coral bleaching by bacteria (VCB and VSB)
is caused by the bacteria Vibrio coralliily ticus
and V. shiloi189 respectively. A bacterium190 (Vibrio charcharia) has been identified in cases
of White Band Disease. White Pox Disease
(WPD) also appears to be provoked by bacterium191 (Serratia marcescens). And in White
Plague (WPL), another bacterium, Aurantimonas coralicida192, seems to be the origin of
the disease.
Sea fan (Gorgonia ventalina) suf fering from Aspergillosis (ASP)

OCEANA/ Houssine Kaddachi
[ 47 ]
07. Threats to Corals
More than 50 species have been af fected by

high mortality rates or degradation processes which have been traced to the diseases
mentioned above. Although the majority of
these cases occur in tropical reefs, two of
them have been detected in the Mediterranean (VSB and FPS).
In the summer of 1999, a massive die-of f of
anthozoans and other animals (mollusks,
bryozoans, tunicates, and sponges) took
place during high waters temperatures down
to more than 40 meters deep in the sea of
Liguria and the French coasts of Provence193.
The corals that were af fected were red gorgonian (Paramuricea clavata), white gorgonian (Eunicella singularis), yellow gorgonian
(Eunicella cavolini), pink gorgonian (Eunicella
verrucosa), red corrals (Corallium rubrum),
(Leptogorgia sarmentosa), Mediterranean
madreporaria (Cladocora caespitosa) and
yellow colonial anemone (Parazoanthus a xinellae). In some areas, between 60% and
100% of the existing colonies were killed, resulting in the deaths of millions of gorgonians
and other anthozoans194.
Bearded fireworm (Hermodice carunculata)

OCEANA/ Juan Cuetos
[48]
For reasons still unknown, the female polyps

of the P. clavata population had an especially
high mortality rate.195
Studies have shown that gorgonians suf fered
a high degree of stress due to the high temperatures and were susceptible to a while
range of microorganisms, including fungi
and protozoans. As a result, this disease was
named as the Fungio-protozoo Syndrome
(SFP).
These occurrences have not been isolated

cases. In colonies of Parazoanthus a xinellae,
the mortality rate was repeated over various
years, considerably reducing the presence of
this species in Portofino and other zones of
the Ligurian Sea. Recently, these mortalities
have been blamed on high temperatures and
the proliferation of a number of pathogenic
agents such as cyanobacteria of the Porphyrosiphon196 genus.
In 2003, new mortalities were registered in
anthozoan populations of the Mediterranean,
which this time ex tended beyond the French
and Italian coast to Spanish waters. During
that year, a high mortality rate among red

gorgonians (Paramuricea clavata) was detected and traced to the overgrow th of mucilage197 and madreporaria (Cladocora caespitosa) with necrosis198 in the Columbretes
Islands Marine Reserve, which af fected more
than 60% of the colonies of both species.
Similar mortality rates were detected in other
coralline algae colonies from Cabo de Creus
to Cabo de Palos199. All cases pointed to abnormally high water temperatures.
The high mortality rates of red gorgonians in

the straits of Messina were also at tributed to
high water temperatures and the concurring
increase in mucilaginous algae (Tribonema
marinum and Acinetospora crinita)200. The
same causes were traced to episodes of
bleaching observed in Mediterranian madreporaria corals, (Cladocora caespitosa),
Mediterranean cup coral (Balanophyllia europaea) or Argentinean coral (Oculina patagonica) in dif ferent zones of the western
and eastern Mediterranean over the last few
decades201.
Bleaching is usually at tributed to the bacteria
Vibrio shiloi, which can cause ex tensive damage in large swathes of Oculina patagonica
colonies; however, the bacteria appears to
be sensitive to ultraviolet radiation, which has
therefore spared the more shallow coral colonies202.
Recently, the bearded fireworm (Hermodice

carunculata) has been proven to be a disease
vector for corals. This polychaete can act as a
winter reservoir for pathogens203, preventing
the bacteria from dying in cold water months
and allowing them to spread outside once
ambient temperatures rise again.
Climate change
As we have already seen, climate change has

been behind the mass die-of fs of anthozoans
that have occurred in recent years in the Med-
iterranean Sea. But aside from diseases and

bleaching, higher water temperatures have
other pernicious ef fects on anthozoans.
Due to the release of carbon dioxide emissions into the atmosphere, it is predicted
that the oceans will increase their absorption
of CO2, thereby provoking changes in the
chemical composition of the water. A larger
quantity of CO2 leads to a lower pH of water,
increasing the acidity of the oceans and reducing the availability of carbonate ions. The
consequence of this is a reduced calcification
rate204, which will af fect many marine organisms that need calcite or aragonite to form
their skeletons, like corals.
If we take into account that calculations forecasting conditions over this millennium say
that the oceans will absorb 90% of anthropogenic CO2205, we can understand the changes
this will have on the marine ecosystem. Studies indicate decreases in calcification around
40% over the nex t 50 years, and can even
decrease as much as 80% before the end of
the century.
In the Mediterranean, climate change is bringing with it other threats to coral life: changes
in the abundance of species that are sensitive
to changes in water temperature and the introduction and spread of non-native species
which may compete with the Mediterranean
ones206.
Other anthropogenic
effects on corals
Ripping out colonies

In areas that are popular with divers, the natural mortality rate of gorgonians is increased
three-fold by damages and uprooting that
occur there207. Significant amounts of death
are caused by divers ripping out gorgonians
and other large anthozoans through bad diving practices and by damage inflicted through
the anchoring of boats in areas where these
[49]
07. Threats to Corals
colonies are present. Some protected areas

in the Mediterranean, such as Medas Islands
in Spain or Port Cros in France have suf fered
damage caused by excessive diving and anchoring in vulnerable zones208.
Dredging, beach regeneration, and the construction of coastal infrastructure may move
large amounts of sediments or create a
change in the location of deposition, thereby
af fecting corals. It is known that the mechanisms which corals use to avoid burial by an
excess of sediment have a high energetic
cost associated with them211. Furthermore, in
some seas such as the Caribbean, the high
rate of sedimentation has been traced to diseases that af fect gorgonians212.
Fishing and corals
One of the greatest threats to corals are the

various fishing techniques and fishing gears
which damage colonies or rip them from the
substrate they are anchored too. Bot tom
trawling and other such methods requiring
dragging nets across the sea floor have the
gravest impact on these animals.
Ovulid on gorgonian OCEANA/ Juan Cuetos
Chemical pollution
There is very lit tle data on the ef fect of chemical pollution on corals. However, some episodes of gorgonian and red coral mortality
have been traced to it. For example, in deep
waters of the Mediterranean (80-160 meters)
in 1987, mortality trends were linked to high
rates of organochlorides such as PCB at the
mouth of the Rodan Estuary209.
Burial and colmatation
For many species of corals, sedimentation
levels are a natural limiting factor. This is why
it is not common to see gorgonians or corals close to the mouths of rivers or in horizontal seabeds where the colmatation (high
level of sedimentation) occurs at a great rate.
Conversely, they are more common on rocks
and vertical walls where sedimentation is less
likely210.
[50]
Oceana has also been able to verif y the impact of other fishing gear in contact with the
seabed (fixed nets, longlines, and traps).
While set ting or removing the gear, or while
they are dragged by marine currents, they can
snag on corals, rip them from their substrate
or lacerate them. Nevertheless, dredging and
trawling are the techniques which cause the
most damage and mortality to populations of
coral, gorgonians, sea feathers and anemones. Each day more studies verif y the damage these types of gear cause to corals and
other benthic sea life213.
It is well known that trawling is the principal

cause of the deterioration of these ecosystems
in many parts of the world.214 Scientists have
recognized that in general, wherever trawlers drag over coral reefs there is a chance
that serious damage will be caused.215 The
Secretary General of ICES, David Grif fith,
considers that dragging a heav y net over a
deep-sea reef is similar to driving a bulldozer
through a natural reser ve. The only way to
protect those reefs is to find out where they

are located and to prevent the trawlers from
dragging their nets over them.
Various studies have revealed the damage inflicted on coral reefs by dif ferent fishing techniques in zones of Atlantic of depths between
200 and 1,200 meters.216 Trawlers can destroy
up to 33 km2 of habitat on the continental shelf
in less than 15 days.217 The United States National Marine Fisheries Service (NMFS) has
estimated that in Alaska alone a single trawler
can rip out 700 kilos of deep-sea coral in a
single cast.218
In the Mediterranean, it has been shown that
fishing for crustaceans over bot toms with
fine sediments that shelter the Isidella elongata gorgonian and sea feathers (Funiculina
quadrangularis) provokes a significant loss
of biodiversity.219 Furthermore, the surfaces
harboring these two species in the western
Mediterranean have almost disappeared
completely due to this type of fishing.220
Dif ferent conclusions derived from observing

gorgonians damaged by fishing gear show
how ex tremely vulnerable their colonies are,
as well as the long recovery time they need,
sometimes exceeding a century.221
But corals do not only suf fer damage from
direct impacts. Some colonies can be buried
or suf fer severely reduced feeding capacity due to water turbidity from resuspended
sediments af ter bot tom trawling events. The
sediment stirred up by trawlers can be redeposited hundreds of meters deeper than its
original location, leading to sessile organisms
very far away being buried.222
When a colony is damaged by fishing gear

or anchors, it can lead to the opportunistic
set tlement of epibionts (hydrozoans and bryozoans) which can then finish them of f- by
either a coverage that prevents the polyps
from feeding, or by creating a greater surface

area that is resistant to wave action and currents. They can also be colonized by nematodes and polychaetes which can weaken the
colony.223
Some fisheries management measures in the

Mediterranean could avoid causing damage
to some populations of anthozoa.
One regulatory proposal for fishing in the

Mediterranean presented by the European
Commission224 seeks to prohibit trawling in
waters less than 50 meters deep and to thus
protect some of the most vital ecosystems of
this sea, such as coralline algae. These measures have been accepted by countries such
as Spain and are now part of their national
legislation225 (although there is a lack of a
consistent characterization of Mediterranean
seabeds, which prevents more ef ficient protection of these areas). Unfortunately the proposal has not yet been applied in all territorial
EU waters because it has been repeatedly
blocked by some Mediterranean countries,
provoking a legal loophole which only bnefits
destructive fishing techniques and accentuates the destruction of benthic habitat.
On the other hand, recent decisions rendered
by the General Fisheries Council for the Mediterranean (GFCM) prohibit trawling below a
depth of 1000 meters226 and in some marine
seamounts and coral reefs227 could help conserve some reefs and gorgonian gardens in
this sea.
Unfortunately, illegal fishing in prohibited areas is a common practice in the Mediterranean, which means that no species of coral is
safe from menace.
[51]
08.
Coral uses
Tree coral (Dendrophyllia ramea) OCEANA/ Juan Cuetos
[ 52 ]
Commercial exploitation
of corals
Some species of coral have been sought and
collected since antiquity because of their attractive appearance. They have been used
as elements of jewelry and costume accessories, and more recently, souvenirs.
The jewelery industry has focused on the

exploitation of so-called precious corals.
Among these are the various species of the
Corallium genus. The most sought-af ter are
those that may be found throughout the Mediterranean and adjacent Atlantic waters, such
as the red Mediterranean coral (Corallium
rubrum). Also highly prized are those of the
Pacific, like the red Pacific coral (Corallium
regale), pink corals (Corallium secundum y
C. laauense) and others (C. japonicum, C.
nobile, C. elatius). Black corals, blue coral
(Heliopora coerulea), and recently, bamboo
corals (Keratosis, Isidella, and Lepidis genuses) are also used. Golden corals (Gerardia, Narella, Calyptrophora or Callogorgia genuses) are also used, although they are less
valuable as jewels due to their porosity, small
size, fragility and other qualities which make
them unsuitable, and they generally wind up
on the souvenir market. The large colonies of
false black coral (Gerardia savaglia) ex tracted from the Sea of Marmara are also used to
make costume jewelry228.
In the Mediterranean, red coral has been the
most prized species and has given rise to an
industry specialized in ex tracting it, which
has had a large impact on the species and on
the seabed itself.
The intensive exploitation of this octocorallians has caused a nearly 70% reduction in
the last decades229. Ex traction has gone from
around 100 tons caught per year at the end
of the 1970s, to catches of barely 30 tons per
year barely 20 years later230.
Today is it a rare species despite the fact that

it once had a density of more than 1,000 colonies per square meter. That density is only
ex tant today in marine protected areas or in
places where access is dif ficult. Furthermore,
many of the colonies existing today are small.
In Spanish areas where red coral exploitation
is still permit ted, 91% of the colonies measure less than five centimeters tall231. In Italy,
66% of the colonies that were studied are not
reproductive232.
Red coral (Corallium rubrum) OCEANA/ Juan Cuetos
The ex traction of these corals has traditionally involved very destructive methods, such
as the San Andrs Cross or the Italian Bar.
The lat ter instrument consists of a metal bar
weighing over a ton with chains and crests
made of nets at tached to it. This was dragged
over the seabed and broke apart the coral.
Only a small portion of the uprooted coral
was actually captured in the nets and recuperated, while the rest lay lost and dead on the
seabed. Sometimes there were up to 2,000
boats dedicated to capturing red coral233.
In 1994, the EU234 prohibited the use of the
San Andrs Cross and similar systems for
capturing red coral, but this method has yet
to be included among those prohibited by the
Bern Convention.
[ 53 ]
08. Coral uses
Corals and medicine

Over the last decades, biomedical research
has looked to the sea as a source of new
medicine. Sponges and actinia have been
among the animals in which new composites
for pharmaceutical use have most frequently
been found, along with the cnidarians. Gorgonians, corals and anemones are supplying
raw materials and useful information to fight
diseases. The Mediterranean is seen as an
excellent place to seek these species.
Tree coral (Dendrophyllia ramea) OCEANA/ Juan Cuetos
Today, ex traction of red coral is mainly performed by divers235 who collect it at depths
of up to 120 meters, although in some areas
they use remotely operated robots with mechanical arms.
Torre del Greco (Italy) is the main market for

precious Mediterranean coral. The lack of red
Mediterranean coral has obliged the industry
to seek imports mainly from other areas, especially in the Pacific. Today the town operates a coral industry which is valued at more
than 30,000 million dollars per year236.
Antivirals have been found in the yellow gorgonian (Eunicella cavolini).237 The synthetic
composite 9--D-arabinosiladenina (ara-A)
has been found to be analogous to espongotimidina, a metabolite which is naturally
produced by this species along with its close
cousin spongouridine, analogous to 1--Darabinofuranosiluracil, (ara-U).
Gorgonias are also important as sources of

diterpenes: the eunicellin present in the white
gorgonian (Eunicella singularis)238 or palmonine found in the pink gorgonian (Eunicella
verrucosa)239 are other examples. Other anthozoa have also contributed their share, with
Given the small sizes of most colonies that

still exist today in the Mediterranean, in the
last few years a new system to exploit and sell
the smaller colonies has come into existance.
It consists of melting down the specimens
which bear lit tle market value due to their
small size, and generating a sof t paste used
to make various articles of costume jewelry.
Although there organized market, other species of anthozoa may also join the ranks of
the souvenir items, like Cladocora caespitosa
or Dendrophyllidae spp. and other scleractinians.
[ 54 ]
Eunicella cavolini OCEANA/ Juan Cuetos
drates245. For example, the paralyzing toxins

born by certain anemones, such as those in
the Anthopleura and Anemonia genus, could
be useful as local anesthetics246. The equinatoxin found in the beadlet anemone (Actinia
equina) may be useful for controlling cholesterol levels247.
Sarcodyction catenatum OCEANA/ Juan Cuetos
cembranolides being found in false red gorgonians (Parer y thropodium coralloides)240 and
sarcodictine in Stolonifer coral (Sarcodyction
catenatum).241
One must not forget the sesterterpene cladocoranes found in Mediterranean madreporaria (Cladocora cespitosa)242, which has pharmaceutical value in the treatment of various
diseases, including cancer, as it has potential
antitubercular and antibacterial properties
that inhibit the grow th of Gram-positive bacteria.
Equally important is the false black coral

(Gerardia savaglia), which has been found to
contain lectin which could potentially be used
in the treatment of the Human Immunodeficiency Virus (HIV)243.
New applications for phosphorescent proteins found in some anthozoa are also being
sought for their use in detecting and visualizing cancerous cells244.
Anthozoans have great pharmaceutical potential and one of the most singular characteristics of these animals, is their stinging
cells and the toxins they contain. As is well
known, venoms are very useful compounds in
medicine. There are very varied compounds
found in corals, including peptides, proteins,
phospholipids, phospholipase, glycoproteins, sterols, bioactive amines and carbohy-
Gerardia savaglia OCEANA/ Juan Cuetos
[ 55 ]
09.
[ 56 ]
Protected corals
Polycyathus muellerae OCEANA/ Juan Cuetos
Invertebrates are the forgot ten souls of

national, international and EC legislation.
It is even worse if they happen to live in the
oceans. This is why corals are hardly present
in conservation proposals and laws despite
their great importance to oceanic ecosystems.
Of the nearly one and a half thousand species listed in the annexes of the EU Habitats
Directive, less than 200 are invertebrates. Of
these, eight are from the sea, and only one
is an anthozoan. Furthermore, of the nearly
200 habitats listed therein, only nine are marine habitats and only one is directly related
to corals: the reefs.
In the Bern Convention, the annexes list
around 2000 species, of which 130 are invertebrates, some 40 are of marine origin, and
five are anthozoans. Moreover, the list prohibiting capture techniques and gear does not
include a single vertebrate except in the section regarding the use of explosives and venom for harvesting decapods (crustaceans).
In the Washington Convention (CITES) for

controlling the international commerce of endangered species of plants and animals, the
three appendices list nearly 35,000 ta xons,
but only 2,100 of these are invertebrates. In
Annex II, however, there are a number of anthozoans, including all of the species belonging to the orders Sclaractinia and Antipatharia, as well as the helioporidae and tubiporidae
families, which brings the total to more than
1,200 species.
The Barcelona Convention comprises several

dif ferent protocols. The Protocol Concerning Specially Protected Areas and Biological
Diversity in the Mediterranean is the one focused on marine animals and plants. Together they list some 120 species among the different annexes: 48 are invertebrates, but only
five of them are anthozoans.
OSPAR Convention does not include a single anthozoan and only two of the 14 priority
habitats have some type of relation to corals:
Lophelia per tusa reefs and the sea feather
clusters and other species which are able to
burrow.
Furthermore, the the World Conservation Union (IUCN), which has evaluated over 40,000
species,248 only includes three anthozoans
among the nearly 4,000 invertebrates that
were analyzed: The Ivell anemone (Edwardsia ivelli), the starlet anemone (Nematostella
vectensis) and the pink gorgonian (Eunicella
verrucosa). These last two are classified as
vulnerable.
In Europe, there are also some conventions

which refer exclusively or particularly to the
sea. These are the Barcelona Convention for
the protection of the Mediterranean Region
and the OSPAR Convention for the protection
of the marine environment of the North-East
Atlantic.
Red gorgonian (Paramuricea clavata) OCEANA/ Juan Cuetos
[ 57 ]
Corals that are included in international agreements and European regulations
[58]
Less than 20% of the corals living in the Mediterranean have been included under the annexes of the conventions for the protection of
animal life. Of these, most of them (85%) are
only protected under Annex II of CITES, which
does not ex tend full protection but rather regulates their commercial trade. Furthermore,
this control does not cover the fossils of these
species, despite their importance for the marine ecosystem, vital both for reef formation
as well as for serving as an optimal substrate
for the set tlement of new colonies.
This means that, excluding CITES, there are

six species of anthozoans which are protected under EC legislation or international
conventions for the protection of wildlife
which have been signed and ratified by the
EU. However, only two are included in the
lists with ma ximum protection (Annex II of
the Bern Convention and Annex II of the Barcelona Convention). The other species have
been included on the list of species for which
management plans should be established.
Moreover, these species of coral are not protected in their full range of habitat, but only in
the Mediterranean.
Jewel anemone (Cor ynactis viridis) OCEANA/ Juan Cuetos
[59]
10.
[60]
Oceana and corals
Elisella paraplexauroides OCEANA/ Juan Cuetos
Given the importance of corals, gorgonians

and anemones to the marine ecosystem and
the uncertain future of many species given
the various dangers and degree of vulnerability, Oceana proposes the development of a
Management Plan for Mediterranean Anthozoa which includes the following measures:
Prohibiting the use

of destructive fishing gear
over coral seabeds
A first necessary measure for conserving corals is prohibiting the use of trawling, dredging and other similar techniques in places
with vulnerable ecosystems, such as those
formed by the corals mentioned hereaf ter for
inclusion in the Habitats Directive. This prohibition must also be part of a definitive and
approved list used to regulate fishing in the
Mediterranean.
Equally important is the execution of resolutions that have already been approved by
the GFCM, among which are the protection
of the Eratosthenes seamount, the protection
of the Lophelia Per tusa reef at Santa Maria
di Leuca, the cold infiltration of hydrocarbons
from the Nile Delta, and the protection from
bot tom trawling at depths greater than 1,000
meters. All of these should be present in EC
legislation.
Regulating the capture of corals

To avoid non-selective and high-impact techniques for capturing coral, the definitive prohibition of trawling gear and other apparatus
for capturing coral, including robots with
mechanical arms, must be passed into law.
These methods must be included both within
Annex IV of the Bern Convention, as well as
Annex VI of the Habitats Directive.
There must also be quotas, closed areas
and minimum sizes for red coral or any other
commercialized anthozoan. The manufacture
of paste derived from small coral specimens
must be prohibited.
In the case of red coral, there must be specific limits to oversee the reduction of quotas
by 50%, together with recovery plan and a
five year monitoring plan to evaluate the evolution of the species so that a moratorium on
all captures can immediately be established
should their number keep diminishing.
Updating and improving
European legislation and
international conventions for
the protection of flora and fauna
Habitats that are generated or inhabited by
anthozoans must urgently be included in Annex I of the Habitats Directive. It should include
the dif ferent types of corral reefs, gorgonian
gardens, coralline algae and anthozoan clusters (sea feathers or gorgonians) in sof t sediment bot toms, among other habitats.
Furthermore, many species of corals should

become part of the annexes of the Habitat Directive and the Bern Conventions, BARCON
and CITES.
For example, Annex I of the Habitats Directive should be included the following species:
Cladocora caespitosa reefs, deep-sea coral
reefs, gorgonian gardens, clusters of Isidella
elongata and pennatulaceas in fine sediment
beds, gorgonian facies and other anthozoa in
coralline algae, carbonate mounds, submarine elevations (including seamountains, hills
and mounds), clif fs and walls, and fossil and
sub-fossil reefs.
Annex II of the Habitats Directive should be
listed all the anthozoan species that are vulnerable or threatened, starting with the species mentioned hereaf ter.
Annex IV of the Habitats Directive, Annex II of

BARCON and Annex II of the Bern Convention should add the following species to their
lists at the very least: Astroides calycularis,
Gerardia savaglia, Isidella elongata and Funiculina quadrangularis.
[61]
In Annex III of BARCON and in Annex III of

the Bern Convention, the following species
must be included: Paramuricea clavata, Eunicella singularis, Cladocora caespitosa, Dendrophyllia sp. In addition, all species of antipatharians should be included as well.
In Annex V of the Habitats Directive, all species of scleractinias and antipatharians should
be included as a preventive measure to avoid
overexploitation and abusive commercialization.
As mentioned above, Annex VI of the Habitats
Directive and Annex IV of the Bern Convention should include a description of prohibited anthozoan capture techniques, such as
the Italian Bar, the San Andrs Cross, and
any other trawling gear or mechanic device
created for that purpose.
Finally, CITES should include Gerardia

savaglia in Appendix I for ma ximum protection, and Corallium rubrum in Appendix II.
Evaluation and recovery plans

for threatened species
Given the great unknown that currently exists
regarding most Mediterranean anthozoans
and their conservation status, a 10-year evaluation plan must be provided for within the
Barcelona Convention in order to assess the
state of coral populations, gorgonias, and
anemones in the Mediterranean.
Once this assessment period is finished, the

species of anthozoans which are found to be
in danger or vulnerable must then be ex tended to international conventions and European legislation aimed at preventing the degradation of these populations. Furthermore, a
management and recovery plan for the species to be protected should be prepared for
inclusion in the annexes of these agreements
and laws.
[ 62 ]
As a mat ter of urgency, the first studies must

focus on those species which are in gravest
danger (such as the scleractinians), those
which are commercially exploited (such as
red coral, false black coral, anemones, etc.),
those which face the greatest mortality rates
(Paramuricea clavata, Eunicella singularis,
Cladocora caespitosa, etc.), those which form
and maintain habitats and those of which lit tle
is yet known (such as the species of the inferior circumlit toral and the deep-sea species,
such as Viminella flagellum, Elisella paraplexauroides, Paramuricea macrospina, Spinimuricea klavereni, and Callogorgia ver ticillata).
In addition, it is very possible that in the nex t

few years new corals will appear or be discovered in this sea that will be in need of
management plans. In fact, Oceana has recently found a new species in the Mediterranean whose known distribution in the Atlantic
Ocean was limited to areas like the Bay of
Biscay and the Canary islands.249 This new
species is the whip gorgonian, Spinimuricea
atlantica.
Reducing the impact
of human activities on corals
The Mediterranean countries, especially the
European ones, should be more scrupulous
in executing their international agreements for
reducing CO2 emissions into the atmosphere
in order to avoid the harmful ef fects that climate change and subsequent acidification of
the sea has on coral life. Moreover, they must
lead international ef forts to achieve more
drastic reductions of contaminating gases,
since the Mediterranean will be one of the
zones most af fected by climatic change.
Also, all necessary measures to avoid spilling

pollutants into the sea must be enacted, and
the International Convention for the Control
and Management of Ships Ballast and Sediments, under the International Maritime Or-
Leptogorgia lusitanica OCEANA/ Thierry Lanoy
ganization (IMO) charter for preventing spills

and the introduction of non-native species,
must be ratified, applied, and improved.
The development of activities which can adversely af fect corals, gorgonians, and anemones must not be permit ted. These include
coastal construction, dredging, quarrying,
etc., carried out without first performing due
and proper environmental impact studies and
sustainability plans.
Marine protected areas

All Mediterranean countries must keep anthozoans in mind as one of the most important
values when it comes to creating marine reserves and protected areas.
Control and regulations systems must also

be developed for recreational diving and the
anchoring of watercraf t in areas with vulnerable seabeds, including the development of
educational materials.
[ 63 ]
11.
Trumpet anemone (Aiptasia mutabilis) OCEANA/ Thierry Lanoy
[ 64 ]
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Car yophyllia inornata making room for

itself within the Spirastrella cunctatrix sponge
OCEANA/ Juan Cuetos
[83]
The research included in this report and its publication were carried out by Oceana with the assistance of Fondazione Zegna (the Zegna
Foundation)
Project Director | Xavier Pastor
Report Writer | Ricardo Aguilar
Editor | Marta Madina
Publication Assistants | Rebecca Greenberg, Maribel Lpez, Giorgio Contessi, Concha Martnez, Elena Alonso
Photography | Houssine Kaddachi, Juan Cuetos, Xavier Pastor, Carlos Surez, Thierry Lanoy
Cover | Tube anemone (
) OCEANA/ Juan Cuetos
The majority of the photographs included in this report were taken by Oceana during the expeditions of the Oceana Ranger in 2005 and
2006.
Design and layout | NEO Estudio Grfico, S.L.
Printer | Imprenta Roal
Photo montage | Pentados, S.A.
Acknowledgements | Oceana would like to thank the following people for the collaboration it received while in Italy: Area Marina Protta
di Portofino; the Central Institute for Marine Research (ICRAM); the University of Padova; Stefano Schiparelli from the University of
Genova; Xabi Reinare; Neptune Plongee; and Bastia Sub. It would also like show its appreciation for the support it received in Spain
from the Fundacin Biodiversidad and from all of the staff at the National Parks and Marine Conservation Reserves in Medas, Columbretes, Cabo de Gata, Alborn and Chafarinas.
Reproduction of the information gathered in this report is permitted as long as OCEANA is cited as the source.
Via Marconi 23
13835 Biella (Italia)
www.fondazionezegna.org
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Corals Mediterranean Eng

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The corals of the

The corals of the

The evolution of corals in the Mediterranean

02. Physical characteristics of corals

03. Coral species of the Mediterranean

04. Coral habitats and corals as habitats

05. Coral reproduction

Sexual and asexual

The corals of the Mediterranean

Berried anemone (Alicia mirabilis) OCEANA/ Juan Cuetos

Budding, fission and laceration

Trouble with the neighbours: space

Commercial exploitation of corals

09. Protected corals

10. Oceana and corals

06. The fight for survival

07. Threats to corals

08. Coral uses

- Prohibiting the use of destructive fishing gear over coral seabeds

The corals of the Mediterranean

There are corals which live as solitary animals

Fire coral (Millepora planata) OCEANA/ Houssine Kaddachi

Many people believe that corals are plants.

Corals belong to one of the oldest ex tant

Occupying 1.1% of the surface of the worlds

Brief explanation of the

The very term coral is ambiguous in itself,

The anthozoans, or flower animals from the

The cnidarians, or coelenterates, derive their

The hydrozoans and cubozoans spend part

The Octocorallia (or Alcyonacea) derive their

Hexacorallia (or Zoantharia) is not only the

The corals of the Mediterranean

mesenteries. They divide into seven orders:

In the second half of the 20th century, some

There is no scientific consensus as to the

White gorgonia (Eunicella singularis) OCEANA/ Juan Cuetos

The corals of the Mediterranean

A coral is a polyp which can live alone or in

Corals only have one orifice serving as both

02. Physical characteristics of corals

One of the structures most characteristic of

However, there are other cnidarians which

Clavularia sp. OCEANA/ Juan Cuetos

One must not forget that although the small

White gorgonia (Eunicella singularis) with bulging calyces

Solitary anthozoans may be found living in the

All anemones live in isolated units or in small

Although Cladocora caespitosa takes its

The corals of the Mediterranean

Maasella edwardsii OCEANA/ Juan Cuetos

Exoskeletons for all tastes

The exoskeleton of hard corals (scleractinias)

There are also examples of sof t skeletons,

When polyps live in colonies, they must find

Mediterranean madreporaria (Cladocora caespitosa) OCEANA/ Juan Cuetos

The corals of the Mediterranean

Dead mans fingers Alcyonium acaule

Yellow sea fan/gorgonia Eunicella cavolini

Tall sea pen Funiculina quadrangularis

Koster sea pen Kophobelemnon stelliferum

Sea Feather Pennatula aculeata

Sea Pen VIrgularia mirabilis

Cornucopia Cornularia cornucopiae