Open Journal of Marine Science, 2011, 1, 50-68
doi:10.4236/ojms.2011.12006 Published Online July 2011 (http://www.SciRP.org/journal/ojms)
Scleractinian Corals and Reefs of Vietnam as a Part of th e
Pacific Reef Ecos ys tem
Yuri Ya Latypov
A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Rus si an Aca demy of Sci e nces, Vladivostok, Russia
Received May 12, 2011; revised June 1, 2011; accepted June 10, 2011
Abstract
The paper analyzes both published and unpublished results of the investigations of Vietnamese reef building
corals and reefs performed in the last decades of the twentieth century and first decades twenty-first. The
state of the art in the study of reef-building scleractinian corals and reefs is presented. The scleractinian
fauna of Vietnam is shown to match in species diversity (350 species of 80 genera) the tropical coral fauna of
the Indonesian–Malacca fertile center, from which Indo-Pacific reef-building corals originated. The whole
Vietnam coast from the Gulf of Tonkin to the Gulf of Siam is a biogeographically single whole and is a part
of the Indo-Polynesian Province of the Indo-Pacific Area.
Keywords: Vietnam, Reefs, Reef-Building Corals, Pacific Reef Ecosystem
1. Introduction
1.1. Brief the Ecological Characteristic and a
History of Studying
The coastline of Vietnam is over 3200 km long and cov-
ers 15 degrees in latitude, from the Gulf of Siam in the
south (8°N) to the Chinese border in the north (23°N).
The near shore water area (up to 50-m deep) of Vietnam,
including some 3000 islands, is about 206,000 km2.
Vietnam and its coastline are divided into 5 parts, the
Gulf of Tonkin, Central and Southern Vietnam, Gulf of
Siam, and Spratly Islands [1]. Reef-building corals and
reef accumulations are confined to hard grounds, typical
of the Vietnam coast. Between 16° and 19°N, the coast-
line is formed mostly by moving sand with a minor
presence of hard substrates. The temperature varies be-
tween 18˚C - 32˚C, and the salinity, 28‰ - 40‰. One
hundred and fourteen rivers are registered along the
coastline. The spread of the reef is limited near the
mouths of two large rivers, the Red River in the north
and the Mekong in the south, due to adverse conditions.
The ecosystems of the coral reefs of Vietnam feature
high bioproductivity, with a primary production of up to
30 - 100 mg·C/m3 per day, which is almost 100 times that
in open waters [2,3].
Vietnam is situated in the tropics, affected by two
sorts of monsoons: the wet southwest, lasting from May
till September, and the dry northeast, occurring in Octo-
ber-April. Heavy rain showers during the wet monsoon
period result in a huge (5 - 400 million m3) freshwater
influx and a substantial (up to 200 thous. tons) terri-
genous sediment influx into the sea. The daily suspended
matter precipitation rate in the reef reaches 70 - 100 g/m2
and increases tenfold during typhoons [4,5]. This results
in a remarkable decrease in water transparency, affecting,
together with other factors, the development of coral
settlement in this region.
The reef-building corals and reefs of Vietnam attracted
scientific attention as early as the first half of the twenti-
eth century [6-8] was the first to analyze the zonation of
reef-building corals in reefs of the Khanh Hoa province.
He distinguished four scleractinian-dominated facieses.
These investigators determined the species composition
of scleractinians and demonstrated its similarity to that of
Australia and Indonesia. Beginning in 1980, systematic
studies of Vietnam corals and reefs have been performed
in joint expeditions by the Institute of Marine Biology
(Vladivostok), Nha Trang Institute of Oceanography,
Haiphong Institute of Oceanology, and WWF (World
Wide Fund for Nature). The published results were
mainly related to scleractinian composition and distribu-
tion, with some papers analyzing common accompanying
macrobenthos species and a few publications providing
the general characteristics of the reefs. Part of the data
obtained was presented only in unpublished reports.

Y. Y. LATYPOV
51
Some findings were published in difficult-to obtain re-
gional works, including Vietnamese ones.
To date, in a region bordered by the Gulf of Tonkin in
the north, the Gulf of Siam in the south, and the Spratly
Islands in the South China Sea, all reef-building areas
including large islands and shoals have been studied
(Figure 1).
It is thus topical to review the data available on the
coral composition and distribution and development of
Vietnam’s reefs, as well as on their relation to the Indo-
Pacific reef ecosystem. The rapid growth of the country’s
population of 70 million, together with travel industry
development and marine aquaculture intensification [9],
has resulted in increased anthropogenic pressure on this
unique ecosystem. Generalizing data on the composition,
structure, and environmental conditions of Vietnam’s
reef communities will allow one to evaluate the degree of
their degradation and the outlook for their conservation
and recovery.
1.2. Morphological Features of the Vietnamese
Reefs
Both along the coastline and around the islands, reef-
building corals form diverse reef topographies. These
include small fringing reefs along the coastline, barrier
reefs separated from the continent (Re Island and Jang
Bo Reef), and atolls (Spratly Islands) in the open South
China Sea [10-14]. Vo and Hodgson [5] reported Viet-
nam’s reefs to include both true reef frameworks and
coral gardens. Various calcareous structures occurring on
reefs might be formed by coral settlements, usually
called coral layers, communities, or specialized settle-
ments. Such structures are typical of early reef develop-
ment stages and lack any geomorphological and vertical-
biological zonation [15-19]. All Vietnam reefs display
distinct biological and more or less prominent mor-
phostructural zonation. By the specificity of geomor-
phological and climate conditions, Vietnam’s fringing
reefs are clearly distinguished into two types (Figure 2).
The Gulf of Siam
Figure 1. Schematized map of the surveyed regions. 1: Bai
Tu Long Archipelago; 2: Ze Island; 3: Cape Danang, Cham
and Son Tra islands; 4: Re Island; 5: reefs of the Khanh
Hoa Province; 6: Thu Island; 7: Ca Thuik Islands; 8: Con
Dao Islands; 9: Tho Chu Island; 10: An Thoi Archipelago
and Namsu Islands; 11: Rach Gia bay; 12: Royal Bishop
and Astrolab shoals; 13: Spratly Islands.
Figure 2. Schematized profiles of structur al (a) and unstr uc tur ed (b) reefs. See the text for commen ts .

Y. Y. LATYPOV
52
Reefs of the first type have a distinct zonation (reef
lagoon, reef-flat, etc.) with a developed carbonate frame-
work, so-called structural reefs [20] common for the
tropical zone of the World Ocean. Reefs of the second
type display a weak morphostructural zonation, with
some zones occasionally absent. Carbonate deposits in
such reefs comprise only coral settlements of a low
thickness, hardly changing the overall substrate profile.
These are so-called structureless [20] or encrusting [18]
reefs. Vietnam’s structural reefs are mainly formed in
closed bights and on the organogenic base of Holocene
reefs [21,22], while structureless reefs are formed off
promontories and in open bights, mostly on stone and
rocky substrates [18,23-25].
Vietnam’s reefs pertain to the epicontinental monsoon
type [7,11,21,24,26,27]. They are situated at the South
China Sea periphery. The shoal waters of this region are
highly eutrophicated, and the grounds are highly silted
due to the huge amount of terrigenous influx. Other hy-
drological conditions are also not optimal for reef-build-
ing coral growth. Thus, in the Gulf of Tonkin, the salinity
may drop to 26‰, and winter water temperature to 16˚C.
Heavy northeast monsoon winds generate coastal waves
up to three meters high with a 6-s period. During south-
west monsoons, the Vietnam coast is struck by 5 - 10
typhoons per year with the strongest storm excitements
and strong desalination, that sometimes results in almost
utter annihilation of coral reef [5,28]. All this renders
appreciable influence on features of formation of reefs
which occurs, as a rule, on a stony, rock-stony substratum
and less often on coralogenous sediments of Holocene
reefs. Vietnam’s reefs feature a moderate vertical and
horizontal spread and low thickness of modern reef-de-
rived deposits. Their offshore spread usually does not
exceed 200 - 300 m. They rarely extend to a depth of over
20 m. Sometimes they lack distinct morphological zona-
tion. Most of Vietnam’s reefs have an indistinct reef flat
and slope. In some reefs, mostly ones on stone and boul-
der substrates, the only distinct zone is the reef slope.
However, they all have a distinct vertical biological zona-
tion, showing up in the dominant species succession and
in the change in the composition and structure of coral
communities and accompanying macrobenthos. Barrier,
platform and fringing reefs on Holocene organogenous
deposits always have well expressed morphological zon-
ality, characteristic for the majority of reefs Indo-pacific
[12,27,28].
2. Results
2.1. Species Diversity
According to the studies performed in the first decades of
the twenty first century, Vietnam’s reef-building coral
fauna comprises 350 species (see Table 1), pertaining to
80 genera (including 9 ahermatypic corals), of which 137
species, belonging to 26 genera, were not previously
known for that region, and 13 species from 6 genera
were described for the first time [11,18,29-33]). As in
most Indo-Pacific reefs [5,19,24,34-36], the species di-
versity of Vietnam’s reefs consists mainly of the mem-
bers of 5 families, Acroporidae (117 species), Faviidae
(42 species), Fungiidae (32 species), Poritidae (31 spe-
cies), and Dendrophylliidae (25 species), making up al-
together 64.48% of the total scleractinian species com-
position (Figure 3). The five genera most diverse and
widespread in all reefs comprise Acropora (90 species),
Montipora (28 species), Porites (20 species), Favia (14
species), and Fungia (12 species) are most various and
numerous on all reefs, making 47% of all specific riches
of scleractinian (Figure 4).
In all, some 20 scleractinian species form mono- spe-
cific settlements, varying from small “spots” (tens of
square meters) to extended zones (hundreds of square
meters), with a coverage reaching 60% - 100%. One fifth
of all scleractinians occur throughout the Vietnam coast
(Figure 5).
As a whole, the species diversity of reef-building
scleractinians in different areas of the Vietnam coast is
quite comparable, ranging from 190 species in the Gulf
of Tonkin to 265 in the South Vietnam (Figure 6). Simi-
lar (193 - 256) numbers of species were reported for
reefs of Indonesia, the Philippines, and Western Austra-
lia [36,37]. Central and South Vietnam reefs are most
similar in species composition and are quite comparable
to Spratly reefs). The degree of similarity of specific
composition of Scleractinian various areas of Vietnam is
resulted on the clustered diagram (Figure 7).
The peculiarity of the coral faunas of the Siam and
Tonkin gulfs as revealed by cluster analysis (Figure 8) is
consistent with their ecological peculiarities [7,23,38,39].
Their scleractinian diversity is partly caused by their
similar hydrological regimes. Both gulfs are shallows
with high water eutrophycation and turbidity, with a
predominance of clay sediments.
These factors cause a similarity of the biological and
morphostructural zonation of reefs and species composi-
tion of reef communities in the gulfs. At the same time,
certain differences in climatic and geomorphological
conditions of the gulfs result in some dissimilarities in
their scleractinian species composition. The development,
zonation, species composition, and structure of the reefs
in the gulfs were reported previously [13,23,39,40], so
here, only major similarities and differences will be
mentioned.

Y. Y. LATYPOV
53
Table 1. List species of scleractinian of vietnam.
Species Gulf of
Tonkin
Gulf of
Siam
Central
Vietnam
Nhatrang
Bay
Island
Thu
Kondao
Islands
Spratly
archipelago
Acantastrea echinata (Dana, 1846) + + + + +
Acantastrea hemprichii (Erhenberg, 1834) + + +
Acantastrea hillae (Wells, 1955) + + + +
Acropora abrolhosensis (Veron, 1985) + + +
Acropora abrotonoides (Lamarck, 1816) + + + + +
Acropora aculeus (Dana, 1846) + +
Acropora acuminata (Verril, 1864) + + + +
Acropora anthocercis (Brook, 1892) + +
Acropora aspera (Dana, 1846) + + + + + + +
Acropora austera (Dana, 1846) + + + +
Acropora brueggemanni (Brokk, 1893) + + +
Acropora cerealis (Dana, 1846) + + + + +
Acropora clathrata (Brook, 1981) + +
Acropora cytherea (Dana, 1846) + + + + + + +
Acropora dendrum (Bassett-Sminh, 1890) +
Acropora digitifera (Dana, 1846) + + + + + + +
Acropora divaricata (Dana, 1846) + + + + + + +
Acropora donei (Veron & Wallace, 1984) +
Acropora elseyi (Brook, 1892) + +
Acropora fasciculare (Latypob, 1992) +
Acropora florida (Dana, 1846) + + + + + + +
Acropora formosa (Dana, 1846) + + + + + + +
Acropora gemmifera (Brok, 1892) + + + + +
Acropora glauca (Brook, 1893) + +
Acropora grandis (Brook, 1892) + + + + + + +
Acropora granulosa (Edw. & Haime, 1860) + + + + + +
Acropora horrida (Dana, 1846) +
Acropora humilis (Dana, 1846) + + + + + + +
Acropora insignis (Nemenzo, 1967) + +
Acropora hyacinthus (Dana, 1846) + + + + + + +
Acropora latistella (Brook, 1892) + +
Acropora listeri (Brook, 1893) + + + +
Acropora longicya t h us (Edw. & Haime, 1860) + + + + +
Acropora loripes (Brook, 1892) + + + +
Acropora lutkeni (Crossland, 1952) + + +
Acropora microclados (Ehrenberg, 1834) + +
Acropora microphthalma (Verril, 1869) + + + + +

Y. Y. LATYPOV
54
Acropora millepora (Ehrenberg, 1834) + + + + + + +
Acropora monticulosa (Brüggemann, 1879) + + +
Acropora multiacuta (Nemenzo, 1967) +
Acropora nasuta (Dana, 1846) + + + + + +
Acropora nobilis (Dana, 1846) + + + + + + +
Acropora palmerae Wells, 1954 + + + +
Acropora parilis (Quelch, 1886) +
Acropora paniculata (Verrill, 1902) + +
Acropora papillare (Latypov, 1992) +
Acropora pulchra (Brook, 1891) + + + + + + +
Acropora pulchra (Brook, 1891) + + + + + + +
Acropora robusta (Dana, 1846) + + + + + + +
Acropora samoensis (Brook, 1891) +
Acropora sarmentosa (Brook, 1891) + + + + +
Acropora secale (Studer, 1878) + + +
Acropora selago (Studer. 1878) + + + + + +
Acropora solitariensi s (Veron & Wallace,1984) +
Acropora squamata (Latypov, 1992) + +
Acropora speciosa (Quelch, 1886) + +
Acropora subglabr a (Brook, 1891) +
Acropora tenuis (Dana, 1846) + + + + +
Acropora valenciennesi (Edw. & Haime, 1860) + + + + + +
Acropora valida (Dana, 1846)
Acropora vaughani (Wells, 1954) + + + +
Alveopora allingi (Hoffmeister, 1925) + + +
Alveopora marionensis (Veron & Pichon, 1982) + +
Alveopora verrilliana (Dana, 1872) + +
Anacropra forbesi (Ridley, 1884) + +
Astreopora cuculata (Lamberts, 1980) + +
Astreopora listeri (Bernard, 1896) + + + + +
Astreopora longiseptata sp nov +
Astreopora myriophthalma (Lamarck, 1816) + + + + + + +
Astreopora ocellata (Bernard, 1896) + + + + + + +
Australogyra zelli (Veron & Pichon, 1977) + +
Balanophyllia cummingii (Edw. & Haime, 1848) + +
Balanophyllia stimpsoni (Verrill, 1865) + +
Barabattoia amicorum (Edw. & Haime, 1850) + + + + + +
Barabattoia mirabilis (Yabe & Sugiyama, 1941) + + + + + +

Y. Y. LATYPOV
55
Caulastrea furcata (Dana, 1846) + + +
Caulastrea tumida (Matthai, 1928) + + + +
Coeloseris mayeri (Vaughan, 1918) + + +
Coscinaraea columna (Dana, 1846) + + + + + + +
Coscinaraea exesa (Dana, 1846) + + +
Coscinaraea mcneil li (Wells, 1962) +
Ctenactis echinata (Pallas, 1766) + + + + + +
Cycloseris cf. Sinensis (Edw. & Haime, 1851) + + +
Cycloseris costulata (Ortman, 1889) + + + +
Cycloseris cyclolites (Lamarck, 1801) + + + + +
Cycloseris densicolummelus sp. nov. +
Cycloseris patelliformis (Boschma, 1923) + + +
Cycloseris somervillei (Gardiner, 1909) +
Cycloseris tenuis (Dana, 1846) + + +
Cycloseris vaughani (Boschma, 1923) + + +
Cynarina lacrymalis (Edw. & Haime, 1848) + + + +
Cyphastrea chalcidicum (Forskål, 1775) + + + + +
Cyphastrea japonica (Yabe & Sugiyama, 1932) + +
Cyphastrea microphthalm a (Lamarck, 1816) + + + + + +
Cyphastrea serailia (Forskål, 1775) + + + + + + +
Dendrophyllia aculeata sp. nov. + +
Dendrophyllia arbuscula (Van der Horst, 1922) + + +
Dendrophyllia cornige r a (Lamarck, 1816) +
Dendrophyllia horsti (Gard. & Waugh, 1939) + + +
Dendrophyllia japonica (Regberg, 1892) + +
Dendrophyllia laborelli (Zibr. & Brilo, 1984) +
Dendrophyllia robusta (Bourne, 1905) + +
Dendrophyllia sphaerica (Nemenzo, 1981) + +
Diaseris fragilis (Alcock, 1893) + + + +
Diploastrea heliopora (Lamarck, 1816) + + + + + + +
Echinophyllia aspera (Ellis & Solander, 1786) + + + + +
Echinophyllia echinata (Saville-Kent, 1975) + + + +
E. echinoporoides (Veron & Pichon, 1980) + + + + + +
Echinophyllia nichihirai (Veron & Pichon, 1990) + + +

Y. Y. LATYPOV
56
Echinophyllia orpheensis (Veron & Pichon, 1979) + + + + + + +
Echinophyllia patu la (Hodgson & Ross, 1981) +
Echinopora gemmacea (Lamarck, 1816) + + + + + +
Echinopora hirsutissima (Edw. & Haime, 1849) + + +
Echinopora horrida (Dana, 1846) + + +
Euphyllia ancora (Veron & Pichon, 1980) +
Euphyllia cristata (Chevalier, 1971) + +
Euphyllia divisa (Veron & Pichon, 1980) + + + +
E. glabrescens (Chamisso & Eysenhard, 1821) + + + + + + +
Euphyllia grandiseptata sp. nov. +
Euphyllia yaeyamaensis (Shirai, 1980) + + +
Favia favus (Forskål, 1775) + + + + +
Favia laxa (Klunzinger, 1879) + + +
Favia lizardensis (Veron & Pichon, 1977) + + + + + + +
Favia maritima (Nemenzo, 1971) + + + + + + +
Favia matthai (Vaughan, 1918) + + + + + +
Favia maxima (Veron et al., 1977) + + + + + +
Favia pallida (Dana, 1846) + + + + + + +
Favia rotumana (Gardiner, 1889) + + + + + + +
Favia rotundata (Veron & Pichon, 1977) + + +
Favia speciosa (Dana, 1846) + + + + + + +
Favia stelligera (Dana, 1846) + + + + + + +
Favia truncatus (Veron, 2000) +
Favia veroni (Moll & Best, 1984) + +
Favites abdita (Ellis & Solander, 1786) + + + + + + +
Favites chinensis (Verrill, 1866) + + + + + +
Favites complanata (Ehrenberg, 1834) + + + + +
Favites flexuosa (Dana, 1846) + + + + + + +
Favites halicora (Ehrenberg, 1834) + + +
Favites pentagona (Esper, 1794) + + + + +
Favites russelli (Wells, 1954) +
Favites solidocolumellae sp. nov. +
Favites virens (Dana, 1846) + +
Fungia concina (Verrill, 1864) + + +

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57
Fungia corona (Döderlein, 1901) +
Fungia danai (Edwards & Haime, 1851) + + + + + +
Fungia fungites (Linnaeus, 1758) + + + + + + +
Fungia granulosa (Klunzinger, 1879) + + +
Fungia horrida (Dana, 1846) + +
Fungia repanda (Dana, 1846) + + + +
Fungia scabra (Döderlein, 1901) + +
Fungia scruposa (Klunzinger, 1879) + +
Fungia scutaria (Lamarck, 1801) + + + + + +
Fungia seychelensis (Hoeksema, 1993) +
Fungia valida (Verrill, 1864) + +
Galaxea astreata (Lamarck, 1816) + + + + + + +
Galaxea crassiseptata (Latypov, 1998) +
Galaxea fascicularis (Linnaeus, 1767) + + + + + + +
Galaxea vesiculosa sp. nov. +
Gardineroseris pavonoides sp. nov. +
Gardineroseris planulata (Dana, 1846) + + + +
Goniastrea aspera (Verrill, 1865) + + + + + + +
Goniastrea australiensis (Edw. & Haime, 1857) + + +
Goniastrea cf. palauensis (Yabe et al., 1936) +
Goniastrea edwardsi (Chevalier, 1971) + + + + + +
Goniastrea favulus (Dana, 1846) + +
Goniastrea pectinata (Ehrenberg, 1834) + + + + + + +
Goniastrea retiformis (Lamarck, 1816) + + + + + + +
Goniopora columna (Dana, 1846) + + + + +
Goniopora djiboutiensis (Vaughan, 1907) +
Goniopora fruticosa (Saville-Kent, 1893) + +
Goniopora lobata (Edwards & Haime, 1851) + + + + + + +
Goniopora pandoraensis (Veron & Pichon, 1982) + +
Goniopora stokesi (Edwards & Haime, 1851) + + + + + + +
Goniopora stutchburyi (Wells, 1955) + + +
Goniopora tenuidens (Quelch, 1886) + + + +
Halomitra pileus (Linnaeus, 1758) + +
Heliofungia actiniformis (Quoy & Gaimard,1833) +

Y. Y. LATYPOV
58
Herpolitha limax (Esper, 1797) + + + + + + +
Herpolitha weberi (Van der Horst, 1921) + + +
Heterocyathis aequicostatus (Edw. & Haime,1848) +
Heteropsammia cochlea (Spengler, 1781) + +
Hydnophora exesa (Pallas, 1766) + + + + + + +
Hydnophora microconos (Lamarck, 1816) + + + + + + +
Hydnophora rigida (Dana, 1846) + + + + +
Isopora cuneata (Dana, 1846) + + + + + +
Isopora palifera (Lamarck, 1816) + + + + + +
Leptastrea bottae (Edwards & Haime, 1849) + +
Leptastrea pruinosa (Crossland, 1952) + + + + +
Leptastrea purpurea (Dana, 1846) + + + + + + +
Leptastrea transversa (Klunzinger, 1879) + + + + +
Leptoria phrygia (Ellis & Solander, 1786) + + + + + + +
Leptoseris explanata (Yabe & Sugiyama, 1941) + + + + +
Leptoseris gardineri (Van der Horst, 1921) + +
Leptoseris hawaiiensis (Vaughan, 1907) + +
Leptoseris mycetoseroides (Wells, 1954) + + + + +
Leptoseris scaba (Vaughan, 1907) + +
Leptoseris var.mycetoseroides (Wells, 1954) + + +
Leptoseris yabei (Pillai & Scheer, 1976) + + + + +
Lithophyllon bistomatum (Latypov, 1995) +
Lithophyllon mokai (Hoeksema, 1989) + + + +
Lithophyllon undulatum (Rehberg, 1892) + + + + + +
Lobophyllia grandis sp. nov. + +
Lobophyllia corymbosa (Foskål, 1775) + + + + +
Lobophyllia flabellioformis (Veron, 2000) +
L. hattai (Yabe, Sygiyama & Eguchi, 1936) + + + + + + +
Lobophyllia hemprichii (Ehrenberg, 1834) + + + + + + +
Lobophyllia pachyse pt a (Chevalier, 1975) + + +
Lobophyllia robusta (Yabe & Sugiyama, 1936) + + +
Madracis kirbyi (Veron & Pichon, 1976) + + + + + +
Merulina ampliata (Ellis & Solander, 1786) + + + + + + +
Merulina scabriscula (Dana, 1846) +

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59
Montastrea annuligera (Edw. & Haime, 1849) + + + + + +
Montastrea colemani (Veron, 2000) +
Montastrea curta (Dana, 1846) + + + + + +
Montastrea magnistellata (Chevalier, 1971) +
Montastrea valenci ennesi (Edw. & Haime, 1848) + + + + + +
Montipora aequituberculat a (Bernard, 1897) + + + + + + +
Montipora angulata (Lamarck, 1816) + + +
Montipora australie ns i s (Bernard, 1897) + + + +
Montipora caliculata (Dana, 1846) + + + + +
Montipora crassituberl at a (Bernard, 1897) + + + + +
Montipora danae (Edwards & Haime, 1851) + + +
Montipora digitata (Dana, 1846) + + + + + + +
Montipora efflorescen s (Bernard, 1897) + +
Montipora foliosa (Pallas, 1766) + + +
Montipora grisea (Bernard, 1897 + + + +
Montipora hispida (Dana, 1846) + + + + + + +
Montipora hoffmeisteri (Wells, 1954) + + + +
Montipora informis (Bernard, 1897) +
Montipora millepora (Crossland, 1952) + + + + +
Montipora molis (Bernard, 1897) +
Montipora monasteriata (Forskål, 1775) + + +
Montipora nodosa (Dana, 1846) + + + +
Montipora porites (Veron, 2000) + +
Montipora spongodes (Bernard, 1897) + + + + + +
Montipora spumosa (Lamarck, 1816) + + +
Montipora stellata (Bernard, 1897) + +
Montipora tuberculosa (Lamarck, 1816) + + + + + + +
Montipora turgescens (Bernard, 1897) + + + + + + +
Montipora turtlensis (Veron & Wallace, 1984) + +
Montipora undata (Bernard, 1897) + + + + + +
Montipora venosa (Ehrenberg, 1834) + + +
Montipora verrucosa (Lamarck, 1816) + + + + + +
Montipora vietnamensis (Veron, 2000) +
Moseleya latistellata (Quelch, 1884) + +

Y. Y. LATYPOV
60
Mycedium elephantotus (Pallas, 1766) + + + + + +
Oulastrea alta (Nemenzo, 1981) +
Oulastrea crispata (Lamarck, 1816) + + + + + + +
Oulophyllia bennettae (Veron & Pichon, 1977) + +
Oulophyllia crispa (Lamarck, 1816) + + + + + + +
Oulophyllia levis (Nemenzo, 1959) + +
Oxypora glabra (Nemenzo, 1959) +
Oxypora lacera (Verrill, 1864) + + + + +
Pachyseris rugosa (Lamarck, 1801) + + + + + + +
Pachyseris speciosa (Dana, 1846) + + + + + + +
Pachyseris monticulosa sp. nov. + + +
Palauastrea ramosa (Yabe & Sugiyama, 1941) +
Pavona cactus (Forskål, 1775) + + +
Pavona clavus (Dana, 1846) + + + + + +
Pavona decussata (Dana, 1846) + + + + + +
Pavona distincta sp. nov. +
Pavona explanulata (Lamarck, 1816) + + + + + + +
Pavona frondifera (Lamarck, 1801) + +
Pavona maldivensis (Gardiner, 1905) + + +
Pavona minuta (Wells, 1954) + + +
Pavona varians (Verrill, 1864) + + + +
Pavona venosa (Ehrenberg, 1834) + +
Pectinia alcicornis (Saville-Kent, 1871) + + + + +
Pectinia lactuca (Pallas, 1766) + + + + + + +
Pectinia paeonia (Dana, 1846) + + + + + +
Physogyra lichtensteini (Edw. & Haime, 1851) + + + + + +
Platygyra daedalia (Ellis & Solander, 1786) + + + + + + +
Platygyra lamellina (Ehrenberg, 1834) + + + + + + +
Platygyra pini (Chevalier, 1975) + + + + + +
Platygyra sinensis (Edw. & Haime, 1849) + + + + +
Plerogyra sinuosa (Dana, 1846) + + + + + + +
Plesiastrea versipora (Lamarck, 1816) + + + + + + +
Pleuractis moluccensis (Van der Horst, 1919) + +
Pleuractis paumotensis (Stuthbury, 1833) + + + +

Y. Y. LATYPOV
61
Pocillopora capitata (Verrill, 1864) +
Pocillopora damicornis (Linnaeus, 1758) + + + + + + +
Pocillopora eydouxi (Edwards & Haime, 1860) + + + + + +
Pocillopora kelleheri (Veron, 2000) +
Pocillopora meandrina (Dana, 1846) + + + +
Pocillopora verruc os a (Ellis & Solander) + + + + + + +
Pocillopora woodjones i (Vaughan, 1918) + + + + + +
Podabacia crustacea (Pallas, 1766) + + + + + + +
Polyphyllia novaehiberniae (Lesson, 1831) +
Polyphyllia talpina (Lamarck, 1831) + + + + + + +
Porites annae (Crossland, 1952) + + + +
Porites attenuata (Nemenzo, 1955) + + +
Porites australiensis (Vaughan, 1918) + + + + + + +
Porites cylindrica (Dana, 1846) + + + + + +
Porites deformis (Nemenzo, 1955) + +
Porites densa (Vaughan, 1918) + +
Porites lichen (Dana, 1846) + + + + + + +
Porites lobata (Dana, 1846) + + + + + + +
Porites lutea (Edwards & Haime, 1860) + + + + + + +
Porites mayeri (Vaughan, 1918) + + + +
Porites monticulosa (Dana, 1846) + +
Porites mordax (Dana, 1846) + + +
Porites murrayensis (Vaughan, 1918) + + + + +
Porites nigrescens (Dana, 1848) + + + + + +
Porites rus (Forskål, 1775) + + + + + + +
Porites solida (Forskål, 1775) + + + + +
Porites sp. 1 +
Porites sp. 2 + + + + +
Porites stephensoni (Crossland, 1952) + + + + +
Porites vaughani (Crossland, 1952) + + +
Psammocora contigua (Esper, 1979) + + + + + + +
Psammocora digitata (Edwards & Haime, 1851) + + + + +
Psammocora explanulata (Van der Horst, 1922) + +
Psammocora nierstraszi (Van der Horst, 1921) + + + + + +

Y. Y. LATYPOV
62
Psammocora profundacell a (Gardiner, 1898) + + + + + +
Psammocora superficialis (Gardiner, 1898) + + + + +
Pseudosiderastrea tayamai (Yabe & Sug., 1935) + + + + + + +
Sandalolitha dentata (Quelch, 1884) + + + + + + +
Sandalolitha robusta (Quelch, 1886) + + + + + + +
Scolymia aff. Vitiensis (Brüggemann, 1877) + + + +
Scolymia australis (Edw. & Haime, 1849) + + + + +
Seriatopora calien dr u m (Ehrenberg, 1834) +
Seriatopora hystrix (Dana, 1846) + + + + + +
Stylocoeniella gue ntheri (Basset-Smith, 1890) + + + + +
Stylophora pistillata (Esper, 1797) + + + + + + +
Stylophora subseriata (Ehrenberg, 1834) + +
Symphyllia agaricia (Edwards & Haime, 1849) + + + + +
Symphyllia erythraea (Klunzinger, 1879) +
Symphyllia hassi (Pillai & Scheer, 1976) + + +
Symphyllia radians (Edwards & Haime, 1849) + + + + + + +
Symphyllia recta (Dana, 1846) + + + + + + +
Symphyllia valenciennesii (Edw. & Haime, 1849) + + + + + + +
Trachyphyllia geoffroyi + + + +
Tubastrea aurea (Quoy & Gaimard, 1833) + + + + + + +
Tubastrea coccinea (Ehrenberg, 1834) + + + + + +
Tubastrea diaphana (Dana, 1846) + + +
Tubastrea micrantha (Ehrenberg, 1834) + + + + + + +
Turbinaria bifrons (Brüggemann, 1877) + + + + + +
Turbinaria conspicua (Bernard, 1896) + + +
Turbinaria contort a (Bernard, 1896) + +
Turbinaria crater (Pallas, 1766) + + +
Turbinaria frondens (Dana, 1846) + + + + + + +
Turbinaria mesenterina (Dana, 1846) + + + + + + +
Turbinaria patula (Dana, 1846) + + +
Turbinaria peltata (Esper, 1794) + + + + + + +
Turbinaria radical is (Bernard, 1896) + + + +
Turbinaria reniformis (Bernard, 1896) + + + + + + +
Turbinaria stellulata (Lamarck, 1816) +

Y. Y. LATYPOV
63
Species composi t i on
29%
34%
12%
9%
9% 7% 6
1
2
3
4
5
Figure 3. Relative species diversity of major coral genera in
Vietnam. 1: Acroporidae; 2: Faviidae; 3: Fungiidae; 4:
Poritidae; 5: Dendrophillidae; 6: The rest genera.
Diversity
53%
26%
8% 6%4%3% 6
1
2
3
4
5
Figure 4. Reative species diversity of major coral species in
Vietnam. 1: Acropora; 2: Montipora; 3: Porites; 4: Faia; 5:
Fungia; 6: The res species.
Figure 5. Monospecific settlement on reef Myeu Island at
Nhatrang Bay, depth 2 m.
Species r i chness
0
100
200
300
400
12345678
Numb er s pec i es
Figure 6. Scleractinian species diversity in different regions
of Vietnam. 1: total number of species (347); 2: Gulf of
Tonkin (177); 3: Central Vietnam (204); 4: South Vietnam
(240); 5: Thu Island; 6: Con Dao Islands (211); 7: Spratly
Island (211), 8: Gulf of Siam (251).
Figure 7. Cluster analysis of similarity species composition
on different regions of Vietnam.
Figure 8. Similarity dendrogram of sclera ctinians faunas in
different regions of Vietnam. 1: Gulf of Tonkin; 2: Central
Vietnam; 3: South Vietnam; 4: Gulf of Siam; 5: Spratly
Island.
The reef communities in both gulfs lack members of
Palauastrea and Caulastrea and Acropora cuneata, the
latter occurring in most Vietnam reefs. Plerogyra and
Physogyra are absent in the closed part of the Gulf of
Tonkin, and Pachyseris, Mycedium, and Pectinia, in the
innermost and coastal areas of the Gulf of Siam. How-
ever, some members of the latter three genera and rare
Physogyra and Plerogyra species are found in the open
parts of both gulfs, off Hainan and Tho Chu islands.
Corals having large polyp forms and capable of self
cleaning—Galaxea, Echinopora, Lobophyllia, Echino-
phyllia, Turbinaria, Podobacia, Lithophyllon, Fungia,
and Goniopora—are widespread in both gulfs. The reefs
in both gulfs are dominated by many species of these
genera (Galaxea fascicularis, Goniopora stokesi, Echi-
nopora lamellosa, and Lobophyllia hemprichii), as well
as by Acropora cytherea, A. nobilis, Montipora hispida,
Porites lobata, and P. cylindrica, widespread in Indo-

Y. Y. LATYPOV
64
Pacific reefs. Altogether, these species cover 60% - 80%
of the substrate. Massive Porites colonies (at least 10
species) forming vast monospecific settlements are typi-
cal of both gulfs. At the same time, members of Pocil-
lopora, abundant in most Indo-Pacific reefs (5-7 species),
only rarely occur in the innermost parts of the gulfs (2
species maximum) but are common for island reefs in the
open parts of the gulfs (Tho Chu, Hainan). By and large,
the two gulfs are quite similar in coral diversity and
share 74.3% of species.
2.2. Communities
The distribution and peculiarities of benthic communities
in the coastal part of Vietnam reefs is rather constant
(See Figure 2). As a rule, these are algal-coral commu-
nities, composed of several biocenoses (zones, facieses),
dominated by individual algal or coral species or by
groups of species (Figure 9). The predominance of
Laurencia, Turbinaria, and Sargassum algae in the
coastal zones of the reefs has been reported for many
reef development areas. This may be indicative of an
increase in water eutrophication or later stages of reef
development [38,41-43].
Both along the Vietnam coast and in the whole Indo-
Pacific, in reef zones characterized by relatively stable
conditions (lagoons, deep stony and coral terraces, and
reef slopes), branched, plate, and trumpet colonies of A.
cytherea, A. hyacinthus, Montipora danae, M. foliosa,
Porites cylindrica, P. nigrescens, and others successfully
compete with differently shaped scleractinian colonies
[43-45].
A wider distribution of encrusting and plate colonies
of Euphyllia, Echinophyllia, Mycedium, Pachyseris, and
Turbinaria compared to that of branched forms is di-
rectly caused by lowered illumination. This is also the
case in many reefs of the Indo-Pacific and Caribbean
basins [46-48]. In Vietnam’s reefs, such corals are com-
mon for communities of the slope base, byoherms zone,
and fore reef platform.
Caused by abiotic factors, the vertical distribution of
reef-building corals has a strong effect on the develop-
ment of biotic zonation across a whole reef community,
beginning with settlement-site choice and ending with
interspecific trophic relationships. The relationships be-
tween the species composition of benthic communities of
some reefs as revealed by cluster analysis correlated with
the ecological and physiographical zonation of the reefs
(Figure 10).
Algal-coral lagoon and reef flat communities domi-
nated by red and brown algae and similar in coral and
common macrobenthos species composition formed a
single cluster group, that of communities developing
under similar conditions. The high similarity between
coral faunas from different sites reflected similar, some-
times extreme, conditions of reef flat and shallow-water
stone terrace. At the same time, communities of these
reef zones were set apart from those of neighboring reef
zones. Both in structural and unstructured reefs, poly-
specific reef slope communities sharing a relatively
greater number of corals also formed a distinct cluster.
Polyspecific coral communities of a reef slope of struc-
tural and unstructured reefs distinctly stands apart on the
greatest level of similarity of their specific variety.
Figure 9. Algal-coral community on reef Bathlongvi, depth
2 m.
Figure 10. Cluster dendrogram of the species composition
of benthic communities. Algal-coral community, unstruc-
tured (1) and structural (2) reefs; Acropora community,
unstructured (3) and structural (4) reefs; Acropora + Dip-
loastrea community, unstructured (5) and structural (6)
reefs; (7) bioherm community (reef slope); (8) Junceella +
Diaseris community; (9) Maleus + Junceella community. A:
lagoon, B: reef flat and terrace zone, C: reef slope, D:
for-reef platform.

Y. Y. LATYPOV
65
3. Discussion and Conclusions
To summarize the above, both structural and unstruc-
tured reefs feature vertical biological and geomor-
phological zonations. The latter is mainly determined by
peculiarities of the underwater reef slope substrate.
Similar biological zonation reflecting interzonal differ-
ences in environmental conditions (substrate, wave re-
gime, sedimentation rate, illumination) has been reported
for many of Vietnam’s reefs and various Pacific and
Caribbean reefs [21,23,40,42,43].
Shallow-water Vietnam reefs growing in highly eutro-
phic conditions lack thick reef deposits [18,23] and fea-
ture high coral diversity and distinct biological zonation,
that is, the presence of inner heterotrophic (lagoon, reef
flat) and outer autotrophic (reef slope) zones [26,49,50],
which is characteristic of typical Indo-Pacific reefs.
In reefs of Indonesia and Philippines and in the Great
Barrier Reef, a total of 360 - 410 reef-building scler-
actinians pertaining to 70 - 80 genera have been recorded
[36]. This region of the Western Pacific is considered the
center of origin of tropical coral faunas. The maximum
coral diversity is observed in the so-called Coral Triangle
[36,51,52] with apices in the Philippines, the Malacca
Peninsula, and New Guinea (Figure 11). To same fertile
the center should expense and coast of Vietnam, scler-
actinian fauna which totals 350 species belonging to 80
genera.
Vietnam’s reefs, too, obviously belong to this center,
which is evidenced by their high similarity in coral spe-
cies composition to reefs of Thailand, Indonesia, and the
Philippines (76.4%, 72.3%, and 81.6%, respectively). In
the greater Western Pacific Coral Triangle (with apices
in Vietnam, South Japan, and the Great Barrier Reef),
coral faunas are also highly similar and homogenous.
The similarities between the Vietnam coral fauna and
those of Japan and Australia are 77.5% and 86%, respec-
tively, suggesting homogeneity of the coral fauna of the
Western and Southwest Pacific. As a whole, the species
complex of Vietnam scleractinians, as well as those of
alcyonarians and gorgonarians, belongs to the tropical
fauna as the majority of Vietnam corals are also common
for the equatorial Indo-Pacific reef zone. The scleractin-
ian species composition of this area exceeds 80% of that
of the Pacific, and the alcyonarian diversity of Vietnam’s
reefs is one of the greatest in the Indo-Pacific [18,31,53].
The species composition and high diversity of Viet-
nam’s coral fauna, as well as its close similarity to the
Southwest Pacific coral fauna, allow one to refer it to the
Indonesia-Polynesian center of origin of the coral faunas
of the tropical Indo-Pacific (Figure 12) The whole Viet-
nam coast, from the Gulf of Tonkin to the Gulf of Siam,
is a biogeographically single whole and is part of the
Indo-Polynesian Province of the Indo-Pacific Area.
Figure 11. Schematized map of the generic diversity of reef-building corals in different regions of the Indo-Pacific (partly
after Veron, 1995). The dotted line and arrow indicate new and old 70 genera diversity isolines, respectiv ely.

Y. Y. LATYPOV
66
Figure 12. Optimal Vietnamese coral reef.
4. Acknowledgements
The author is sincerely grateful to the domestic and for-
eign colleagues V. Brykov, I. Budin, N. Selin, Yu.
Yakovlev, Lang Van Ken, Нгуен Van Tien, Vo Si Tuan,
assisting to me in researches and sent prints of their pub-
lications, special gratitude to Dr. Douglas Fenner for
editing of the English text.
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