Download lizard radiations

FURTHER READING
Agardy, D. T. . Scientific research opportunities at Palmyra atoll. A
report submitted to The Nature Conservancy.
Cobb, K. M., C. D. Charles, H. Cheng, and R. L. Edwards. . El
Niño–Southern Oscillation and tropical Pacific climate during the last
millennium. Nature : –.
Davis, A. S., L. B. Gray, D. A. Clague, and J. R. Hein. . The Line
Islands revisited: New Ar/Ar geochronologic evidence for episodes
of volcanism due to lithospheric extension. Geochemistry, Geophysics,
Geosystems: doi: ./GC.
Dawson, E. Y. . Changes in Palmyra atoll and its vegetation through
the activities of man –. Pacific Naturalist : .
Dinsdale, E. A., O. Pantos, S. Smriga, R. A. Edwards, F. Angly, L. Wegley,
M. Hatay, D. Hall, E. Brown, M. Haynes, L. Krause, E. Sala, S. A.
Sandin, R. Vega Thurber, B. L. Willis, F. Azam, N. Knowlton, and F.
Rohwer. . Microbial ecology of four coral atolls in the northern
Line Islands. PLoS ONE : e.
Handler, A. T., D. S. Gruner, W. P. Haines, M. W. Lange, and K. Y.
Kaneshiro. . Arthropod surveys on Palmyra atoll, Line Islands,
and insights into the decline of the native tree Pisonia grandis (Nyctaginaceae). Pacific Science : –.
Keating, B. H. . Insular geology of the Line Islands, in Geology and offshore mineral resources of the central Pacific basin, Circum-Pacific Council for Energy and Mineral Resources Earth Science Series, vol. . B.
H. Keating and B. R. Bolton, eds. New York: Springer-Verlag, –.
Sandin, S. A., J. E. Smith, E. E. DeMartini, E. A. Dinsdale, S. D. Donner,
A. M. Friedlander, T. Konotchick, M. Malay, J. E. Maragos, D. Obura,
O. Pantos, G. Paulay, M. Richie, F. Rohwer, R. E. Schroeder, S. Walsh,
J. B. C. Jackson, N. Knowlton, and E. Sala. . Baselines and degradation of coral reefs in the northern Line Islands. PLoS ONE : e.
UNESCO. . Central Pacific World Heritage Project, International
Workshop Report.
Woodrofe, C. D., and R. F. McLean. . Pleistocene morphology and
Holocene emergence of Christmas (Kiritimati) Island, Pacific Ocean.
Coral Reefs : –.
LIZARD RADIATIONS
MIGUEL VENCES
Technical University of Braunschweig, Germany
Lizards belong to the clade Squamata, together with snakes,
and among nonflying terrestrial vertebrates, they are the ones
most commonly observed on islands. Lizards are characterized by a great facility in colonizing islands and adapting
to novel ecological circumstances by changes in their morphology, physiology, and reproductive biology. They have
consequently become an important model group for the
inferential and experimental study of adaptive radiations.
LIZARDS ON ISLANDS
On major land-bridge islands with favorable climates
(i.e., in the tropical, dry, and temperate zones) both liz-
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FIGURE 1 Emblematic island lizards. (A) Gallotia stehlini, Gran Canaria.
(B) Gallotia galloti, Tenerife. (C) Chalcides sexlineatus, Gran Canaria.
(D) Tarentola delalandii, Tenerife. These species and their relatives have
originated on the Canary Islands. Photographs by Miguel Vences.
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ards and snakes are commonly encountered, with snake
species richness often being similar to lizard species
richness. On  major islands and island groups of the
Mediterranean Sea, there are  occurrences of  species of lizards and  species of snakes. However, native
extant snakes are missing on many smaller islands and on
oceanic archipelagoes such as the Macaronesian Islands
(Canary and Cape Verde Islands, Savage Islands, Madeira,
and the Azores), where native species and even endemic
radiations of lizards are present (Fig. ). The most remote
oceanic islands (e.g., Hawaii) are devoid of both native
lizards and snakes.
Although within-island diversification is rare in snakes
and is limited to very large islands such as Madagascar,
lizards have diversified on medium-sized islands such as
the Greater Antilles as well (see below). Of the currently
known ~ species and  families of lizards, representatives of the Gekkonidae, Iguanidae, Lacertidae, and
Scincidae are most commonly encountered on islands.
Continental islands, especially, may frequently act as
an evolutionary reservoir by enabling the survival of remnants of lineages that became extinct or very rare on the
mainland. Such is the case of the tuataras, two species
of lizard-like reptiles which are the last extant representatives of the Sphenodontia (the sister group of squamates).
At present, tuataras are confined to various small islands
off New Zealand, although fossil remains demonstrate
their past presence on the New Zealand mainland, and
that of their relatives on other continents. On the Balearic Islands in the Mediterranean Sea, the lizard Podarcis
lilfordi is present only on tiny offshore islands surrounding the larger islands of Mallorca and Menorca, where
they are extinct. On Madagascar, the radiation of snakes
in the subfamily Pseudoxyrhophiinae is very diverse, but
FIGURE 2 The largest lizard worldwide, the Komodo dragon, Varanus
komodoensis. Photograph by Thomas Ziegler.
FIGURE 3 Both of the smallest lizards worldwide occur on islands. (A)
The gecko Sphaerodactylus ariasae occurs on Isla Beata and adjacent
areas of Hispaniola. Photograph by S. Blair Hedges. (B) Adult male
Malagasy leaf chameleon of an undescribed species in the genus
Brookesia from the extreme north of Madagascar. Photograph by
Frank Glaw.
this lineage has only a few representatives in Africa, where
it probably has been replaced by other snakes.
Both the largest and smallest extant lizards occur
on islands: The largest is the Komodo dragon (Varanus
komodoensis) with a maximum snout–vent length of over
 mm (Fig. ); the smallest (Fig. ) are two species
of Sphaerodactylus geckos (S. ariasae and S. parthenopion)
from the Caribbean, with adult snout–vent lengths of
about  mm, and several species of Malagasy leaf chameleons (Brookesia) with adult snout–vent lengths of –
mm. Lizards appear to show a trend of island gigantism
and dwarfism opposite to what is generally considered as
a rule: In lineages of small lizards, the island populations
and species become even smaller, and in lineages of large
forms, the island representatives become even larger, especially in carnivorous taxa. Snakes also show size changes
in island populations and species, and snakes that evolved
to become small on islands did so to a relatively greater
degree than those that became large. The observed pattern suggests that snake body size is principally influenced
by prey size, with large snakes mainly feeding on nesting
seabirds and small snakes mainly feeding on lizards.
Many island lizards have adapted to resources that
differ from those available on the nearby mainland. The
most famous is the marine iguana from the Galápagos
(Amblyrhynchus cristatus), the only lizard that feeds on
algae while diving in the ocean. Many lizards of the family Lacertidae were originally insectivorous but became
herbivorous on islands. In fact, herbivory in mainland
lineages may be an important “preadaptation” that allows
for successful colonization of island habitats.
A further intriguing difference between island and
mainland populations of lizards is population density,
which is generally one order of magnitude higher on
islands. This phenomenon is likely driven by distinctly
lower numbers of predators and competitors. These same
factors may also have allowed island lizards to expand
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their diet to include nectar, pollen, and fruit. Indeed, in
several island ecosystems, lizards also occupy an important role as pollinators and seed dispersers.
Few studies have addressed changes in reproductive
strategy in island populations of lizards, but in species of
the family Lacertidae a trend of reduced clutch size and
larger egg size on islands has been noted.
COLONIZATION OF ISLANDS BY LIZARDS
Recent years have seen a paradigm shift in our understanding of the occurrence of many taxa on islands.
This has involved a shift from the dominance of vicariance explanations to hypotheses in which dispersal
plays at least an equally important role. In general,
the mode of reproduction of lizards and snakes, with
internal fertilization, favors overseas dispersal because
the arrival of a single gravid female to an island can
be sufficient to give rise to a new population. For lizards, there is no doubt that their dispersal capacities
are high and that they have on many occasions colonized islands over water from the mainland or from
other islands. For green iguanas, direct evidence exists
that after a hurricane in , at least  individuals
arrived on a mat of logs and uprooted trees on the eastern beaches of Anguilla and other islands in the Caribbean, and some specimens survived there for at least
three years. Molecular genetic analyses have provided
evidence for various events of long-distance dispersal
between Africa and South America (e.g., in geckos of
the genera Tarentola and Hemidactylus, and in skinks of
the genus Trachylepis). For example, Trachylepis atlantica from the Fernando de Noronha Archipelago in the
Atlantic,  km east of the Brazilian coast, belongs to
this mainly African and Malagasy genus rather than
to the related Neotropical genus Mabuya. Its ancestors
presumably colonized by overseas dispersal from Africa
rather than from nearby South America.
Native populations of lizards (and often endemic species) are found on many oceanic islands: on major archipelagos such as Macaronesia, the Galápagos, the Gulf of
Guinea islands, the Comoros, and the Mascarenes, but
also on many small and isolated islands. The Australian region, including small islands such as those of the
Solomon and Bismarck archipelagoes, harbors a massive
radiation of the scincid genus Sphenomorphus, and other
skinks (genus Emoia) have radiated on most islands in the
southwestern Pacific, including, among many others, the
Fiji, New Caledonia, Solomon, and Bismarck archipelagoes. This further demonstrates the capacity for overseas
dispersal of lizards.
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FIGURE 4 An endemic species of chameleon from the Comoro island
of Mayotte, Furcifer polleni. Photograph by Frank Glaw.
Inverse routes of colonization, from islands back to
the mainland, have occurred as well. This appears to be
the case for a Central and South American clade within
the genus Anolis, which probably originated from a West
Indian ancestor, and it is possibly also true for chameleons, which may have dispersed multiple times from
Madagascar to mainland Africa, and which certainly have
dispersed from Madagascar to Mayotte (Fig. ).
On the Gulf of Guinea islands (São Tomé, Principe, and
Annobon), a relatively high proportion of endemic burrowing species of lizards and snakes occur, indicating that the
capacity of overseas dispersal also extends to species living
in humid soil and leaf litter. A combination of ocean currents, floating islands, and reduced surface salinity caused
by freshwater discharges from large rivers may be favorable
to overseas dispersal events in general and may also enable
such soil-dwelling species to colonize islands. Eggs of some
lizards are known to be resistant to immersion in seawater.
In the case of Anolis sagrei, this may explain the survival
of populations of this lizard on small islands vulnerable to
hurricanes, but it also may allow the overseas rafting of lizard eggs in tree holes or mats of vegetation.
In some cases, commensal species of lizards have been
translocated by humans. Several species of geckos of the
genus Hemidactylus have a transcontinental distribution
that in some cases is due to natural colonization but often
may reflect deliberate or, more probably, accidental introductions. Lipinia noctua, a scincid lizard that lives alongside humans on islands of the central and eastern Pacific,
displays a phylogeographic pattern concordant with the
“express train” hypothesis: Specimens may have been
transported as stowaways on early Polynesian canoes during the rapid human colonization of Polynesian islands.
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PATTERNS OF INSULAR LIZARD RADIATIONS
The process of speciation can be either () adaptive (i.e.,
the process of an ancestral population diverging and giving rise to two daughter lineages adapted to different
niches) or () nonadaptive (e.g., the separation of the
daughter species by geographic barriers or by differentiation of features that serve for species recognition).
Most lizard radiations on smaller islands probably
belong to the category of nonadaptive and allopatric speciation on different islands. This same mode of speciation
has also taken place within some islands of sufficient size.
A few possible examples also exist for sympatric adaptive
speciation within an island.
As an instance of nonadaptive speciation on different
islands, the western Canary Islands are populated by small
radiations of skinks and geckos (Chalcides and Tarentola),
but on each island or group of islands, only one species of each genus occurs. The situation is slightly more
complex in the Canarian lacertid lizards, genus Gallotia:
Here an initial split is observed between large-sized and
small-sized species, and sympatry occurs only between
(ecologically strongly differentiated) representatives of
either group (on Hierro, Gomera, Tenerife, and probably
La Palma, if extant species and natural occurrences are
considered). Day geckos of the genus Phelsuma have radiated on the Seychelles and Mascarenes, and on each of
these two archipelagoes there is a monophyletic lineage of
various species and subspecies. At least on the Seychelles,
the available evidence favors allopatric speciation of the
three endemic taxa on different islands, with secondary
sympatry in some cases.
Crucial to test hypotheses of radiation on islands are
robust phylogenies. However, critical data on the interplay of dispersal and vicariance can be provided by the
geological age of an island or of its last connection to
the mainland, and hence the age of evolutionary splits
in the lineage under study. For example, the two Galápagos iguanas (the terrestrial genus Conolophus and the
marine iguana Amblyrhynchus; Fig. ) occur on the same
islands and do form a monophyletic group. This could
be interpreted as an example of speciation by ecological
specialization under sympatric conditions. However, the
age of the evolutionary divergence between these species
predates the geological origin of the current Galápagos
Islands. This indicates that either () they must have
diverged on a previous, now submerged land mass, or ()
both species originated on the mainland, they colonized
the Galápagos independently, and their mainland relatives subsequently went extinct. In general, the possibility
of extinction must always be taken into account to under-
FIGURE 5 Galápagos iguanas. (A) The marine iguana, Amblyrhynchus
cristatus. Photograph by Ylenia Chiari. (B) A terrestrial iguana, Conolophus subcristatus. Photograph by Scott Glaberman.
stand the biogeographic history of lizard populations on
islands.
The best-studied case of an insular lizard radiation is
that of the Caribbean genus Anolis (Iguanidae), the anoles,
which are among the most common terrestrial vertebrates
in the Caribbean and are found on almost every island in
this region. There are over  species of anoles, of which
nearly  are Caribbean. Their origin has been estimated
at around  million years ago, and fossil specimens preserved in amber are known from the Oligocene to the
Miocene of the Dominican Republic. The patterns of
anole radiation have been intensively studied by Jonathan
B. Losos and colleagues. Summaries are found in Losos
() and Losos and Thorpe (), from where much
of the following information has been extracted.
Anoles are very good dispersers, evidenced by cases of
related taxa occurring on islands of great geographic distance. However, by far the highest proportion of Caribbean anoles are endemic to single island banks (more than
%). A few cases of natural hybridization are known, but
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in general, mismating among species of these lizards is
prevented by the throat fans (“dewlaps”) of males, which
show specific colors and patterns used in species recognition. In fact, sympatric species of anoles always differ
in the size, color, or patterning of their dewlaps. Up to
 species of anoles can coexist at a single site, and such
sympatric species almost always differ in terms of habitat
use and morphology or physiology.
The number of anole species coexisting on a certain
island is significantly correlated with island size. Considering only small islands (i.e., islands of a surface of 
km or less), the species–area relationship is stronger for
islands that were in the past connected by land bridges to
other land masses than for isolated islands, highlighting
the importance of historical effects: Land-bridge islands
probably had a higher number of species at the time of
isolation, and through subsequent extinctions, species
numbers adjusted to the island-specific ecological carrying capacity. In contrast, isolated islands depend fully on
over-water colonization as the source for species. Isolated
islands mostly are populated by a single species of anole
only, with a maximum of two species per island (which
then differ in their ecology). Apparently, colonization of
small isolated islands by anoles can be successful only if
() the island does not yet harbor any anole population or
() the island is populated by an anole species that differs
in ecological requirements from the new colonizers.
Evolutionary diversification of anoles appears to occur
on a single island when its size is above a certain threshold. In the Caribbean, within-island diversification has
occurred on the Greater Antilles (Jamaica, Puerto Rico,
Hispaniola, and Cuba). Each of these large islands harbors endemic divergent lineages, which contain various species and, hence, very probably originated on the
island. Within-island speciation can be invoked for at
least % of the Greater Antillean anoles. A few examples from smaller islands or island groups exist of cooccurrence of endemic taxa that could have arisen on the
same island, but these cases are not compelling. Hence,
a certain island area is necessary for within-island speciation, a conclusion that highlights the importance of geography for this process.
The Anolis radiations on the four Greater Antillean
islands (although phylogenetically independent) show
recurrent patterns. As was first pointed out by Ernest Williams, different types of habitat specialists (ecomorphs)
occur on all or most of the Greater Antilles. These are
usually represented by several species on each island (Figs.
–). Initially six ecomorphs were proposed, but others
have since been distinguished. Interestingly, molecular
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FIGURE 6 Ecomorphs of Caribbean Anolis. All species shown are from
Hispaniola. Names roughly denote the preferred habitat of each ecomorph. (A) Crown giant: Anolis baleatus. (B) Trunk crown: A. coelestinus. Note that the photographs are not to scale; Crown Giants are much
larger than all other ecomorphs. Photographs by S. Blair Hedges.
FIGURE 7 Ecomorphs of Caribbean Anolis, continued. (A) Trunk: A.
christophei. (B) Trunk ground: A. cybotes. Photographs are not to scale.
Photographs by S. Blair Hedges.
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Cuba was fragmented during the Miocene. The Anolis
alutaceus group, also on Cuba, contains  species with
narrow distributions, mostly centered on different mountain ranges, a pattern that is also seen in other groups.
Which prevalent pattern of species formation gave rise
to the current diversity of anoles? Adaptive speciation in
sympatry or parapatry may occur in Caribbean anoles, but
it is probably not the main driving force explaining their
diversity. In many cases, populations became isolated on
small land-bridge islands or reached isolated small islands
by overseas dispersal. Geographically and thus genetically
separated from other anole populations, they evolved different morphologies and dewlaps, probably largely because
of adaptation to new ecological conditions. On the larger
islands, species belonging to the main ecomorphs underwent allopatric speciation (e.g., on different mountain
ranges or on parts of their island that were separated by
water barriers in periods of rising sea levels). As summarized in the following section, many examples indicate that
adaptation can occur in the absence of speciation in Caribbean anoles. But it is still uncertain how the initial differentiation of ecomorphs on each of the Greater Antillean
islands took place.
PHYLOGEOGRAPHY AND EXPERIMENTAL
TESTS OF SELECTION
FIGURE 8 Ecomorphs of Caribbean Anolis, continued. (A) Stream: A.
eugenegrahami. (B) Grass: A. semilineatus. (C) Twig: A. placidus. Photographs are not to scale. Photographs by S. Blair Hedges.
data show that, with two exceptions, the ecomorphs arose
independently on the different islands: Different ancestors diversified independently and gave rise to the same
ecological and morphological adaptations.
Species belonging to different ecomorphs usually
occur sympatrically, but species belonging to the same
ecomorph generally are geographically separated within
an island (and have different dewlap colors or patterning). In addition to the six main ecomorphs, many islands
harbor further habitat specialists, but these usually occur
on a single island only.
In several cases, the different species of one ecomorph
occur in geographically separated populations scattered
across an island. In the Anolis carolinensis group, three evolutionary lineages can be distinguished and have ranges
corresponding to three paleo-archipelagoes into which
Deciphering radiations is possible by looking at general
patterns across a whole group or by examining in more
detail the microevolutionary processes. Comparison
of DNA sequences allows phylogeographical analyses
where chiefly the geographical distribution of differentiated alleles (haplotypes) is mapped, and the phylogenetic
relationships among these haplotypes is determined. The
assumption is that haplotypes evolve through mutation,
and different haplotypes get fixed in genetically isolated
populations. In various studies on anoles and Canarian
lizards, Roger S. Thorpe and colleagues have found evidence for discordance between historical and adaptive
patterns. For example, in Gallotia lizards on the Canarian
island of Tenerife, a historical boundary of mitochondrial
haplotype lineages exists between western and northeastern areas, whereas within both groups, morphological
differences were found between northern and southern populations, reflecting strong ecological differences
between the humid north and arid south of the island.
On Dominica, Anolis oculatus shows a complex phylogeographical structure that is not fully concordant with the
phenotypic variability encountered.
These examples demonstrate that morphological
adaptations to local conditions, especially in terms of col-
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oration, can evolve very fast in island lizards. This is also
witnessed by the large variability of lacertid lizard species
inhabiting Mediterranean islands (e.g., Adriatic islands,
satellite islands of the Balearics, or Tyrrhenic islands in
Greece). From many of these archipelagoes, a plethora of
subspecies have been described based on color patterns
and partly on variation in scale numbers, but molecular
studies have rarely found any significant differentiation
between these populations, indicating that the external
differences evolved extremely rapidly, on a geological
timescale. Other work has yielded evidence that in Anolis
sagrei, the number of body scales increases with increasing precipitation and with decreasing temperature in
open arid habitats, and the variation in scale numbers is
probably heritable. In further experiments, the effects of a
potential predator (the ground-dwelling lizard Leiocephalus carinatus) on the behavior of Anolis sagrei was tested
by introducing the potential predator on six small islands
on the Bahamas and using six other predator-free islands
as control sites. As a result, anoles altered their behavior
by using the ground less often, but in addition, a strong
selection took place: Surviving specimens on the experimental islands had larger body sizes and longer hindlimbs
than those on control sites, probably reflecting their better capacities to escape.
Evidence for strong selection pressures acting on
island lizards also comes from further experimental
studies. The Dominican Anolis oculatus displays various
ecomorphological variants related to different conditions between the east and west coasts and the montane
regions of the island. In experiments, lizards were translocated to large lizard-proof enclosures in regions occupied by other habitat types than those in their source
population. Morphology (coloration, scale counts, body
proportions) of the translocated lizards were scored,
and each lizard individually marked. Several months
later, survivors were collected and identified. Morphological differences were found between survivors
and non-survivors (e.g., of specimens of the montane
population in enclosures of the relatively xeric west
coast), and the intensity of selection was dependent on
the magnitude of ecological change experienced by the
specimens in the enclosures.
How these intraspecific processes of fast morphological
variation relate to the actual process of species formation
and adaptive radiation is not clear. Evidence of parapatric
forms with restricted gene flow among them comes from
the islands of Dominica and Martinique; on Martinique
this may constitute evidence for adaptive (ecological) species
formation because the forms are distinguished by current
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habitat and not by historical allopatry. It seems clear that
these lizards have a strong potential to adapt to new ecological conditions by changes in morphology and coloration,
and this may have favored adaptive speciation (mostly under
allopatric conditions). This may also be a factor explaining
the recurrent evolution of similar ecomorphs.
SEE ALSO THE FOLLOWING ARTICLES
Adaptive Radiation / Convergence / Dispersal /
Komodo Dragons / Snakes
FURTHER READING
Losos, J. B. . Integrative approaches to evolutionary ecology: Anolis
lizards as model systems. Annual Reviews of Ecology and Systematics :
–.
Losos, J. B. . Ecological and evolutionary determinants of the speciesarea relationship in Caribbean anoline lizards, in Evolution on islands. P.
R. Grant, ed. Oxford: Oxford University Press, –.
Losos, J. B., and R. S. Thorpe. . Evolutionary diversification of
Caribbean Anolis lizards, in Adaptive speciation. U. Dieckmann, M.
Doebeli, J. A. J. Metz, and D. Tautz, eds. Cambridge: Cambridge University Press, –.
Olesen, J. M., and A. Valido. . Lizards as pollinators and seed dispersers: an island phenomenon. Trends in Ecology and Evolution : –.
Williams, E. E. . Ecomorphs, faunas, island size, and diverse end
points in island radiations of Anolis, in Lizard ecology. R. B. Huey, E.
R. Pianka, and T. W. Schoener, eds. Cambridge, MA: Harvard University Press, –.
LOPHELIA OASES
SANDRA BROOKE
Marine Conservation Biology Institute, Bellevue, WA
The deep-water stony coral Lophelia pertusa (Linnaeus
) creates extensive and complex structures on hardbottomed areas in the deep sea, including continental
shelf bedrock, lithified sediment mounds, volcanic basalt,
and (microbially mediated) authigenic carbonate. Large
colonies of L. pertusa have abundant tangled branches
that provide habitats for diverse and abundant associated
communities. These long-lived and slow-growing coral
ecosystems are currently under threat globally from negative human impact, and although some areas have been
placed under protective legislation, continued international effort is needed to ensure the future of these valuable resources.
CORAL BIOLOGY
There are several species of “framework-building”
deep-water corals (Lophelia pertusa, Oculina varicosa,
LO P H E L I A OA S E S
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