Ecoaquático_Castro e Pires 2001

Prévia do material em texto

BULLETIN OF MARINE SCIENCE, 69(2): 357–371, 2001
357
BRAZILIAN CORAL REEFS: WHAT WE ALREADY KNOW
AND WHAT IS STILL MISSING
Clovis B. Castro and Débora O. Pires
ABSTRACT
The last thorough qualitative description of the unique Brazilian coral communities
was elaborated in the 1960’s. This paper reviews the status of knowledge of Brazilian
reefs, with emphasis on coral communities. Larger reef coral communities were recorded
from near the Equator (00∞53'S) to Cape Frio (23∞S), but the southernmost true reefs are
in the Abrolhos area (18∞S). Fifteen scleractinian species from Brazilian reefs (five
endemics) have been described. In Northern Brazil, the scattered information indicates
mostly offshore or oceanic communities dominated by coralline algae, with Siderastrea
stellata as the main stony coral. However, ‘Parcel do Manuel Luiz’ seems to harbor a
unique coral community established on a rocky structure. Northeastern reefs have mostly
long lines of fringing reefs (many on beach rocks), with submerged parallel lines off-
shore. Eastern reefs are uniquely characterized by the major reef builder species
Mussismilia braziliensis (endemic in this region) and pinnacles (chapeirões) as an im-
portant reef structure (sometimes fused to form bank reefs). Although there is a lack of
information on almost all aspects of reef studies, the current (1990s) surge of papers
indicate they are gaining attention from the scientific community. Current researchers
also have been collaborating with conservation efforts.
Brazil has the only true coral reefs known in the South Atlantic (Paulay, 1997). Leão et
al. (1988) stated that Eastern Brazilian reefs show distinct growth forms, reef building
fauna, and depositional setting. The number and distribution of Brazilian reefs are still
poorly known. The first comprehensive descriptions of Brazilian reefs were by Hartt (1870)
and Branner (1904). However, the most thorough qualitative description of Brazilian reef
coral communities was elaborated in the 1960s by Laborel (1969, 1970). Although Laborel
visited many reefs, he was not able to cover the whole coastline and many offshore reef
areas. After Laborel’s work, there were mostly regional evaluations of reefs (e.g., Leão et
al., 1988; Leão, 1996). Maida and Ferreira (1997), in a field trip guide to some Brazilian
reefs, summarized the history of Brazilian reef studies, and provided a brief account of
reef areas and weather and oceanographical regimes prevailing in Brazil.
Larger reef coral communities were recorded in Brazil from Parcel do Manuel Luiz
(Coura, 1994; Moura et al., 1999), Maranhão State (00∞53'S, 044∞16'W) to Arraial do
Cabo, Rio de Janeiro State (23∞S, 042∞W), and in the oceanic Atol das Rocas and Fernando
de Noronha Archipelago (Laborel, 1970). The most southern true reefs recorded are in
the Abrolhos Reef Complex (18∞S), Bahia State (Leão, 1982). Although there are reefs
for more than 2400 km, not all of these are true biogenic reefs.
The linear distance between the northernmost and southernmost reefs in Brazil is equiva-
lent to the length of Australia’s Great Barrier Reef (approximately 2600 km, Wolanski,
1994). However, the latter has a high reef coral diversity and occupies an area of 350,000
km2 (Norse, 1993), with its reefs relatively close to each other (more than 2500 individual
reefs, Wolanski, 1994). Brazilian reef coral communities are very different. In many cases
they do not form true coral reefs, just reef coral communities flourishing on hard bottoms.
Although not yet estimated, the reef area is incomparably smaller than in the Great Bar-
rier Reef, partly because the usual Brazilian reef areas are narrow. This area is also re-
358 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
stricted because the continental shelf is very narrow in most reef areas, sometimes as
little as 15 km (Milliman and Summerhayes, 1975). Also, very few scleractinian reef
coral species from Brazilian reefs (15, with five endemics) have been described, although
there may be undescribed species.
We intend to review here the status of knowledge of Brazilian reefs, centered on coral
communities. In the light of current knowledge, regional differences in coral reefs and
communities within the Brazilian coast will be shown, indicating gaps in information
about these reefs (Fig. 1).
STATUS OF KNOWLEDGE OF BRAZILIAN REEFS
NORTHERN BRAZIL 1—PARCEL DO MANUEL LUIZ AREA.—The most northern coral com-
munities known in Brazil are off the town of São Luís. There are two main reef areas
described: Parcel do Manuel Luiz (69 km2 centered on 00∞50'S, 044∞15'W) and Banco do
Álvaro (30 km2 centered on 00∞17.5'S, 044∞49.5'W) (M. Coura, pers. comm.). The former
is located some 86 km offshore, 180 km from the town of São Luís (the nearest large
settlement), and 50 km from the edge of the continental shelf. The latter is located 90 km
NW from the former. Coura (1994, pers. comm.) and Moura et al. (1999) briefly de-
scribed these formations. They occur in an area of clear waters (visibility >30 m) and are
constituted by a concentration of isolated pinnacles, each up to 300 m in diameter. These
pinnacles may reach the low tide level, but the top of most of them are in depths of up to
14 m. Their bases usually lie on a bottom at a depth of 25–45 m. They probably form a
coral community flourishing on a rocky substrate, but there have not been any drillings to
confirm this. Coura (pers. comm.) observed that in some areas of Banco do Álvaro
milleporids predominate on the reefs walls, followed by the octocoral Phyllogorgia dilatata.
In the southwestern part of Parcel do Manuel Luiz, Coura described a predominance of P.
dilatata on top and walls in depths of up to 15 m, with corals, sponges, and algae domi-
nating the slopes (approx. 30∞) below this depth. In the Museu Nacional (Rio de Janeiro),
there are voucher specimens of the stony corals Agaricia agaricites, A. fragilis, Favia
gravida, Meandrina braziliensis, Montastrea cavernosa, Mussismilia hispida, Porites
astreoides, Scolymia wellsi, and Siderastrea stellata, of the fire-coral Millepora alcicornis,
and of the gorgonians Plexaurella dichotoma and Phyllogorgia dilatata. Hudson et al.
(1999) added Favia leptophylla, Porites branneri, Scolymia sp., Millepora braziliensis,
and Stylaster roseus to this list and Moura et al. (1999) reported the occurrence of Madracis
decactis. Also, there are specimens of two undescribed new species of large Millepora
(fire coral) and Muriceopsis (gorgonian) (Amaral, 1997; Medeiros, 1998). In the vicinity,
there are records of gorgonians not recorded before in other areas of Brazil (Bayer, 1959;
Castro, 1990; Medeiros, 1998; Medeiros and Castro, 1999).
These reefs are located in an area that Laborel (1970) indicated as the northern limit of
occurrence of hermatypic corals in Brazil, at the outer limit of a northern zone of coral
diversity impoverishment. He reported only S. stellata, F. gravida, and M. alcicornis.
However, current checklists include almost all coral species in the northeastern coast of
Brazil (Table 1). The main absence seems to be Mussismilia harttii, which may form
large colonies in reefs in northeastern and eastern Brazil.
Studies are currently being undertaken sponsored in the main by the Maranhão State
Coastal Management Program (M. Coura, pers. comm.). A Marine State Park, covering
359CASTRO AND PIRES: BRAZILIAN CORAL REEFS
Figure 1. Map of the Brazilian coast, indicating the main known reef areas of northern, northeastern
and eastern regions and the mouth of major rivers. MA = Maranhão State; CE = Ceará State; RN =
Rio Grande do Norte State; PB = Paraíba State; PE = Pernambuco State; AL = Alagoas State; BA =
Bahia State; ES = Espírito Santo State; RJ = Rio de Janeiro State; SP = São Paulo State; SC = Santa
Catarina State. Dotted lines indicate State borders. Sectors 1–5 according to Leão (1996).
360 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
aírrevehc
E,)2991(.late
seriP,)9691(leroba
L
no
desab
atad
noitubirtsiD
.saera
feer
nailizar
B
no
slaroc
diropelli
m
dna
nainitcarelcs
fo
noitubirtsi
D.1
elba
T
orienaJ
ed
oi
R
,lanoica
N
uesu
M
eht
ni
detisoped
sne
miceps
no
dna
,9991
,.la
te
aruo
M
,9991
,.la
te
nosdu
H
,)7991(
lara
m
A
,)7991(
atse
T
,)7991(
.la
te
A
C.)J
R
N
M(
=
L
M
;naebbira
C
=
RS
;
’ziu
L
leuna
M
od
lecraP
‘,lizar
B
nrehtro
N
=
F
R
;euqo
R
oãS
fo
epa
C,lizar
B
nrehtro
N
=
sad
lot
A,lizar
B
nrehtro
N
C
R
;ahnoro
N
ed
odnanreF
dna
saco
R
=
B
N
;tsao
C
sfee
R
,lizar
B
nretsaehtro
N
=
1
srotces
,6991
,oãe
L
gnidulcni(
aiha
B
nrehtro
N
,lizar
B
nretsa
E
-
BS
;)2
=
.)5
rotces,6991,oãe
L
gnidulcni(
aiha
B
nrehtuoS,lizar
B
nretsa
E
seicepS
A
C
L
M
RS
F
R
C
R
B
N
BS
noitubirtsid
fo
ti
mil
nrehtuoS
seticiraga
aicirag
A
X
X
X
X
X
X
X
)J
R
N
M(
S
E,zur
C
atnaS
siligarf
aicirag
A
X
X
X
X
)9691,leroba
L(
A
B,sohlorb
A
adivarg
aiva
F
X
X
X
X
X
X
)9691,leroba
L(
S
E,iraparau
G
allyhpotpel
aiva
F
X
X
X
)9691,leroba
L(
A
B,sohlorb
A
sitcaced
sicarda
M
X
X
X
X
)J
R
N
M(
CS,dnalsI
oderovr
A
sisneilizarb
anirdnae
M
X
X
X
X
X
X
)9691,leroba
L(
J
R,oirF
epa
C
ot
tso
ml
A
asonrevac
aertsatno
M
X
X
X
X
X
X
ti
V
ó
)9691,leroba
L(
)erohsffo(
S
E,air
sisneilizarb
aili
msissu
M
X
X
)9691,leroba
L(
A
B,sohlorb
A
iittrah
aili
msissu
M
X
X
X
X
X
)9691,leroba
L(
]seinoloc
gnivil[
A
B,sohlorb
A
adipsih
aili
msissu
M
X
X
X
X
X
)J
R
N
M(
PS,sotnaS
sedioertsa
setiro
P
X
X
X
X
X
X
X
)J
R
N
M(
CS,dnalsI
ela
G
irennarb
setiro
P
X
X
X
X
X
X
X
)J
R
N
M(
J
R,oirF
epa
C
islle
w
ai
mylocS
X
X
X
X
X
)9691,leroba
L(
]seinoloc
gnivil[
A
B,sohlorb
A
atallets
aertsarediS
X
X
X
X
X
X
)J
R
N
M(
J
R,oirF
epa
C
inilehci
m
aineoconahpetS
X
X
X
X
X
)9691,leroba
L(
A
B,sohlorb
A
sinrocicla
aropelli
M
X
X
X
X
X
X
X
)9691,leroba
L(
J
R,oirF
epa
C
sisneilizarb
aropelli
M
X
X
X
X
X
X
r
T
ê
)0791,leroba
L(
S
E,sahlI
s
aditin
aropelli
M
X
X
)7991,lara
m
A
;9691,leroba
L(
A
B,sohlorb
A
asorrauqs
aropelli
M
X
?
?
)7991,lara
m
A(
)?(
A
B,rodavlaS
aropelli
M
.ps
X
)7991,lara
m
A(
A
M,ziu
L
leuna
M
lato
T
11
51
9
51
31
41
81
361CASTRO AND PIRES: BRAZILIAN CORAL REEFS
354 km2 and including at least three different formations, has protected this area since
1991. The main conservation issue is the heavy traffic of large ships in the nearby São
Marcos Bay, estimated at more than 1000 ships yr-1 (M. Coura, pers. comm.).
NORTHERN BRAZIL 2—UNEXPLORED NORTHERN AREA.—There is evidence of other reefs
between Parcel do Manuel Luiz and the Cape of São Roque (Branner, 1904; Laborel,
1970). Coastal beach rock reefs in Ceará were found to be less than a meter thick (Branner,
1904). On ‘coral reefs’, Branner (1904) stated that “these banks are the same as the coral
reefs further south along the coast of Pernambuco, Parahyba and Rio Grande do Norte”.
However, there are no scientific records of these structures. This gap of information cov-
ers an area extending for more than 1000 km. According to Laborel (1970), this area
presents sandy coastlines and, to this day, is still badly mapped (regarding coral commu-
nities) and little known. There are scarce scientific records of structure and biota, except
for scattered dredgings on the shelf, indicating mostly coralline algae banks; these depos-
its are modern, as indicated by living specimens (Barreto et al., 1975). Based on reports
of local divers, Laborel (1970) mentioned submerged banks dominated by S. stellata and,
on a secondary level, by M. alcicornis. Laborel also mentioned reports of other unex-
plored banks in this area. Spix and Martius (Branner, 1904) reported the use of corals for
making lime in Ceará State, but their place of origin is uncertain. As in other little known
areas of Brazil, most of these bank reefs are probably visited only by local fishermen. A
Marine State Park (Pedra da Risca do Meio) was recently (1997) created including sub-
merged [‘submerged’ will be used here only to indicate that the top is more than 5 m deep
at low tide] beach rocks (10 km off Fortaleza), which seems to present at least one species
of ‘brain coral’ (probably M. hispida) (M. Torres, pers. comm.). There are no research
groups with projects centered on coral communities in this area.
NORTHERN BRAZIL 3—THE CAPE OF SÃO ROQUE REEF GROUP.—Near the Cape of São
Roque, in an area extending 100 km, there are a few isolated offshore reefs known as the
Cape of São Roque Reef Group (Laborel, 1970). These reefs are also almost totally unex-
plored scientifically. Laborel (1970) visited only one of them (Maracajahú Reef—05∞24'S,
035∞16'W) and commented on the others using descriptions given by local fishermen.
They were described as oval reefs located several miles offshore, almost monospecific
regarding stony corals (S. stellata), and with vermetids and crustose coralline algae pro-
viding a great contribution to the reef cover. Recent investigations have shown the wide-
spread existence of coral reefs not appearing on nautical charts in the Cape of São Roque
area (Testa, 1997). Two main situations were described: submerged coral patch reefs and
submerged buried coral structures. The reefs Testa (1997) investigated were in shallow
(£10 m), turbid waters and reached a height of up to 6 m. According to her, shallow cores
showed mostly corallines and corals in their structure. She also indicated a dominance of
living and dead coralline algae and S. stellata on today’s cover, although together with
other scleractinian and Milleporidae species, vermetids, sponges, and seaweeds. Testa
(1997) pointed to the lack of studies on reef composition and ecology in this area, where
there are no marine parks or reserves. No research groups at the moment have projects on
reef communities in this area.
NORTHERN BRAZIL 4—THE OCEANIC ATOL DAS ROCAS AND FERNANDO DE NORONHA ARCHI-
PELAGO.—Although close (150 km apart), these oceanic islands present very different
structures and communities, occurring in clear waters (up to more than 30 m visibility).
The Atol das Rocas (03∞52' S, 033∞49' W), 266 km off the northeast coast of Brazil, is
located on top of a sea mountain 26 km long by 8 km wide; the reef itself is an annular
362 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
formation more than 3.7 km E-W by 2.5 km N-S (Kikuchi and Leão, 1997). At low tide,
the reef top is exposed and most of the lagoon dries out. The area is under the influence of
the Southern Equatorial Current, which runs Westward (into the Cape of São Roque area).
The modern reefs are more than 10 m thick (Kikuchi and Leão, 1997). Echeverría et al.
(1997) reported eight species of corals, with massive dominance by a single species, S.
stellata. Gherardi (1995) also registered the dominance of S. stellata and Maida and Ferreira
(1997) estimated its cover as anything up to 45% on the walls and bottom of some deep
pools. However, a core hole drilled in one of the sand cays showed coralline algae as the
most abundant organisms, forming more than 60% of the recovered rock; corals made up
about 10% of the rock (Kikuchi and Leão, 1997).
Although sharing most of its cnidarian fauna with Fernando de Noronha, Atol das Rocas
in comparison exhibits a lower diversity, due mostly to the absence of hydroids (Echeverría
et al.,1997). The restricted distribution of certain cnidarian species to the atoll is probably
due to specific habitats—as the coarse carbonate debris present in Atol das Rocas is ab-
sent in Fernando de Noronha (Echeverría et al., 1997).
This reef has been protected as a Federal Biological Reserve since 1979, comprising
the whole sea mountain above the 1000 m isobath, and was the first site designated as a
Brazilian ‘marine protected area’.
Although there are large gaps in our knowledge about the Cape of São Roque Reef
Group, it seems to present similarities with the better known Atol das Rocas in terms of
builders and coral cover dominance. Futurestudies may show that the atoll represents the
same kind of community as the Cape of São Roque reef group, being located upstream in
the predominant surface current in the area.
The Fernando de Noronha Archipelago, also oceanic (150 km east of Atol das Rocas),
presents only coral communities flourishing on its rocky shores (Laborel, 1970). How-
ever, reef platforms formed mainly by vermetids and Melobesioidae on the infralittoral
fringe have been described by several authors in different areas of the archipelago (Branner,
1904; Laborel and Kempf, 1966; Kempf and Laborel, 1968; Laborel, 1970; Eston et al.,
1986). All coral species reported for these islands are also found on the Brazilian north-
eastern mainland (Pires et al., 1992). Laborel (1970) indicated an estimated density of
coral colonies larger than that found on the reefs of northeastern Brazil. Laborel (1970)
and Pires et al. (1992) gave qualitative reports of deeper areas with relatively high cover
of M. cavernosa. However, this species population seems to have decreased drastically in
the last few years, being replaced by algal cover (M. Maida, pers. comm.). There is no
quantitative data on this decline and the possible causes are unknown. Maida et al. (1995)
found many colonies of S. stellata (48%) presenting signs of partial necrosis in very
shallow waters and suggested that this damage was caused by uncontrolled tourist activi-
ties. There has been a Federal Marine National Park protecting approximately 50% of the
Archipelago since 1988.
NORTHEASTERN BRAZIL—THE REEFS COAST.—The shore from Natal (approximately 6∞S)
to the mouth of the São Francisco River (approximately 10∞S) presents several coastal
reefs. These reefs present major gaps just south of Natal. The area, approximately 600 km
long, was named by Laborel as Reefs Coast. Although the most striking feature is the
long, straight, beach rock lines that emerge during low tides, there are also records of
multiple submerged lines of beach rocks parallel to the coast at different depths, formed
during different sea level still-stands (Mabesoone, 1964; Laborel, 1970). There are verbal
reports of many areas of still unmapped submerged banks (M. Maida, pers. comm.).
363CASTRO AND PIRES: BRAZILIAN CORAL REEFS
Besides this, there are superficial coral reefs, mostly developed on these submerged beach
rocks (Branner, 1904; Laborel, 1970; Maida and Ferreira, 1997). However, we do not
know of any drillings to estimate composition and thickness of ‘coral reef’ structures in
this area. Bottom depths around them rarely exceed 10 m, in turbid waters.
Coral communities were qualitatively described by Branner (1904), Laborel (1970),
and Maida and Ferreira (1997), but there are no quantitative studies so far. Most reef
studies in this area were conducted on fringing or emerging bank reefs. Nearshore reefs
usually present an emerging eroded top and an outer rim composed mostly of coralline
algae and vermetids. On the top, there are several areas covered by zoanthids and scat-
tered S. stellata and F. gravida. In the infralittoral, there is a certain zonation, with an
upper M. alcicornis zone, a middle zone of M. harttii and M. cavernosa at the bottom.
The main coral builders of the reefs in Tamandaré (south of Pernambuco) seem to be M.
cavernosa and M. harttii (Maida and Ferreira, 1997). The latter species was also reported
as covering all the available surface in areas of the Tambaú Reef, João Pessoa (Young,
1986). Maida and Ferreira (1997) described other Tamandaré Reefs including structures
formed by isolated columns, up to 5–6 m high, and expanded laterally on the top. Laborel’s
(1970) description of these reefs does not mention these columns. Instead, he reported the
occurrence of rounded structures, although with some showing the subjacent form of the
beach rocks.
According to Laborel (1970), the greater development of reef coral communities on the
Reefs Coast was between Recife (approximately 8∞S) and Maceió (approximately 10∞S),
with a gradual impoverishment towards the edges of this area. The mouth of the São
Francisco River, discharging a huge sediment load into the sea, would act as a barrier
between northeastern and eastern Brazilian reefs. Laborel (1969, 1970) affirmed that it
separated the geographical ranges of two subspecies of M.hispida and that it was also the
northern distribution limit of Mussismilia braziliensis.
Maida and Ferreira (1995) examined the recruitment of corals on settlement plates of
three reef zones of Tamandaré. They found no difference in density and mortality rates of
recruits among the reef crest, the basis of the fore reef, and the back reef zone. After one
year of submersion, these authors recorded very low coral recruitment (1.18 ± 0.41 corals
per15 ¥ 15 cm plate; 0.30 ± 0.17 hydrocorals plate-1).
An environmentally protected area was created in 1997 in the area described by Laborel
(1970) as bearing the most developed reefs. It was named Costa dos Corais [Coral Coast].
It extends over 130 km of coastline between Tamandaré (Pernambuco State) and Paripuera
(Alagoas State). There is currently a large conservation initiative under way in this unit,
with contributions from the Interamerican Development Bank, Pew Fellows Program,
Federal, State, and Municipal governments, NGOs, and Universities (M. Maida, pers.
comm). Charting and evaluating the submerged reefs is one of the objectives being pur-
sued in this protected area.
Due to the proximity of large cities and ease of access (in small boats or on foot), the
Reefs Coast has been the most affected by the influence of human activities. The main
conservation problems arise from the presence of a large coastal population, leading to
pollution, overfishing, and unorderly tourism (M. Maida, pers. comm.). These reefs in-
clude some of the most heavily impacted areas in terms of human activity on the Brazil-
ian coast. There are verbal reports of huge amounts of corals being removed in Pernambuco
and Paraíba in the past decades for the production of lime used by the sugarcane industry
and in construction (M. Maida, pers. comm.). Indeed, Branner (1904) had already de-
364 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
scribed this practice in Paraíba State in 1875, suggesting that “the marked absence of
living corals from the northern part of the [Parahyba do Norte] reef is probably due to
some extent to the fact that they have been and still are taken by the inhabitants living in
the neighboring villages for the purpose of making lime”. He stated that the burning of
the corals for lime must have been a very old custom even then.
Although there are clear signs of coral community decay, there are no published assess-
ments on the health of these reefs. Preliminary and unpublished observations in the 1990s,
seeking to promote the creation of the Costa dos Corais, showed reefs in a very bad
condition near the major cities (Recife and Maceió) and increasingly better conditions
away from them (M. Maida, pers. comm.).
 EASTERN BRAZIL—BAHIA STATE.—The State of Bahia has reefs along more than 900 km
of its shoreline; the only area apparently lacking reefs is the northernmost part of the
State, due to the influence of the São Francisco River (Laborel, 1970). The continental
shelf in Bahia averages 50 km in width (Leão et al., 1988). Leão (1996) reviewed the
knowledge on the morphology and distribution of these reefs, and described the major
environmental impacts on them. The author divided the State’s reef area into five main
sectors. Two of these are almost completely unknown; the other three had various aspects
of their geological history, geomorphology, and community structure studied. The posi-
tion and area of submerged reefs in all the sectors are still poorly known.
Northern Bahia (Leão, 1996: sector 1) shows discontinuous bank reefs, in waters usu-
ally shallower than 10 m, formed as isolated columns that may fuse together at their tops
(Leão, 1996). One study showed a reef structure 10 m thickin one of these reefs (Leão et
al., 1988). Other coral-algal communities are present as thin veneers (less than 1 m thick)
on beach rocks parallel to the coastline (Leão, 1996). Here coral diversity is less than in
areas further south, with eight species of hermatypic corals and one hydrocoral recorded
(Leão et al., 1997).
Strong natural decay in coral cover and a shift in the community structure in the north-
ern Bahia coral reefs were found (Leão et al., 1997; Kikuchi and Leão, 1998). Over the
past 3,000 years, the main building coral M. braziliensis has been replaced by S. stellata
or M. hispida. This fact was associated to a 5 m fall in sea level. The authors also reported
a living coral cover of less than 6% in northern Bahia, compared with an estimated 20%
on the exposed old tops. Atlantic and Gulf Reef Assessment Protocol (AGRA) was re-
cently started in this area (Leão et al., 1999).
The Todos os Santos Bay area (Leão, 1996: sector 2), near Salvador, presents fringing
reefs along most of the eastern and southeastern shores of Itaparica Island and the east
coast of islands in the north part of the Bay (Laborel, 1970; Leão, 1996). Seismic profiles
and a core on Itaparica Island reef showed a biogenic reef 10 m thick (Leão et al., 1988).
Laborel (1970) classified the reefs of Itaparica Island as apparently dead and associated
this to the collecting and burning of corals for lime, as previously recorded by Hartt
(1870).
Between Salvador and Abrolhos (Leão, 1996: sectors 3–4) there is a diverse zone, with
coastal reefs (biogenic and beach rocks) and isolated banks offshore. Some of these reefs
are among the least known in Brazil, such as the reefs in the Bay of Camamu (sector 3)
and the Itacolomis Reefs and groups of coral pinnacles surrounding them (approx. 17∞S)
(sector 4). Based on nautical charts and overflights, the Itacolomis Reefs were described
as large, roundish reef structures, separated from each other by irregular canals (Laborel,
1970; Leão, 1996). They are located in an area varying from a few hundred meters to
365CASTRO AND PIRES: BRAZILIAN CORAL REEFS
approximately 10 km from the shoreline. A recent visit to the area indicated a flourishing
coral fauna, with the occurrence of at least 14 of the 15 reef coral species found in Brazil
(C. B. Castro and B. Segal, unpubl. data).
The Abrolhos area (sector 5), in southern Bahia, is located on a widening section of the
continental shelf (up to 200 km). It is considered the largest and richest coral reef area in
the South Atlantic Ocean (Laborel, 1970; Leão, 1982, 1994, 1996; Leão et al., 1988;
Castro, 1994), with relatively well preserved reefs that present all the reef coral species
reported in Brazil (Laborel, 1969, 1970) (Table 1). These are the best known reefs in
Bahia (Leão, 1996) lying on bottoms usually less than 25 m deep. It is limited to the north
by a deep submarine gulf (Poço do Jucurussu, Leão, 1994) and to the south by the mouth
of the Doce River (Laborel, 1970).
There are several reef structures in Abrolhos, of varying morphologies (Leão, 1996)
including fringing reefs, isolated columns (chapeirões), and offshore bank reefs (Leão,
1996; pers. observ.). All known reefs are located from approximately 5 to 65 km offshore.
There are still unexplored areas and evidence of large areas crowded with unmapped
submerged chapeirões (G. Dias, pers. comm.; pers. observ.). The largest of these reefs
(Parcel das Paredes) reaches some 30 km in length (more than 250 km2) (Leão, 1996).
Leão and Ginsburg (1997) emphasized the occurrence of high levels of siliciclastic
sediments in the coastal reef environments of Abrolhos (as seems to be the case in several
other reefs in Brazil), suggesting that the low coral diversity could be a response to this
condition. Moreover, our knowledge of cnidarian faunal diversity is still incomplete, as
the discovery of large new species and records indicate (Castro, 1989, 1990; Marques and
Castro, 1995). Also, Brazilian scleractinian corals need taxonomic reviews to verify if
they are really different from their Caribbean relatives and the occurrence of sibling spe-
cies on today’s nominal species.
Coral covers of up to 39% were detected in some of the Abrolhos reefs (located more
offshore), with an average for the area of around 15% (Villaça and Pitombo, 1997, data
restricted to stations where corals occurred). Most of the coral cover is located only on the
reef tops, in waters usually shallower than 10 m. In extremely shallow waters (0.5 m),
Figueiredo (1997) found coralline covers of 32 to 79% in islands (rocky shores and fring-
ing reefs) of the Abrolhos Archipelago. The author indicated that this cover was similar to
those in reefs in other regions, but the mean absolute growth rates of coralline crusts in
Abrolhos was higher than reported in other areas of the tropical Atlantic.
The only studies on coral reproductive biology in Brazilian reefs, which have been
recently started in the Abrolhos area, show that most coral species have different spawn-
ing periods (Pires et al., 1999; unpubl. data).
Bahia has several marine protected areas: a Federal Marine Park (sector 5-Abrolhos,
created in 1983); a State APA (sector 5-Ponta da Baleia, Abrolhos area); and a Municipal
Marine Park (sector 4-Recife de Fora, Porto Seguro). Almost the whole Abrolhos Bank
above 25 m in depth is included in conservation units.
Leão (1996) commented that the most common anthropogenic agents that are affecting
the Bahian reefs are related to coastal zone development, tourism, predatory fishing, the
installation of industrial projects, and exploitation of fossil fuels. The author indicated
that reefs closer to the shore are under stress, due to increased runoff caused by defores-
tation for agricultural and industrial purposes. Coutinho et al. (1993) found a high density
of macroalgae on reefs closer to the shore in the Abrolhos area, suggesting that predatory
fishing and/or increased sedimentation were the cause. Overfishing was also indicated in
366 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
view of the abundance and size of top predators and larger fish species (Ferreira and
Gonçalves, 1999). In the Abrolhos area, there are no conclusive reports of any adverse
impacts of coastal development on the reefs. Preliminary studies suggested that there is a
relatively small interaction between the coastal zone and the offshore reef systems
(Knoppers, 1996; Lopes and Dutz, 1996; Marone and Camargo, 1996). However, in-
creased sedimentation may be related to coastal activities far away from the immediate
mainland. It was suggested that southern Bahia sediments come from large rivers located
about 200 km north of the reefs areas, such as the Jequitinhonha River (Melo et al., 1975;
Leão and Ginsburg, 1997). Specific examples of industrial activities include paper plants
and offshore oil drilling near reefs in southern Bahia (Leão, 1996). Leão et al. (1994)
made a qualitative description of disturbances in the Abrolhos area due to tourism. Later,
Creed and Amado Filho (1999) studied anchor damage disturbance and posterior recov-
ery of a seagrass meadow in the Abrolhos Archipelago. They observed that scars made by
tourist boat anchors had damaged yearly 0.5% of the meadow. The damage area increased
yearly from 1993 (105 m2) to 1995 (154 m2). Massive bleaching has already been re-
ported in Southern Bahia, being related to a larger scale phenomenon (Castro and Pires,
1999).
SOUTHEASTERN BRAZIL—ESPÍRITO SANTO AND RIO DE JANEIRO STATES—South of Abrolhos,
there are three mouths of large rivers nearby (Mucuri, São Mateus, and Doce). These
rivers together are comparable to the barrier formed by the São Francisco River (Laborel,
1970). The wide continental shelf of southern Bahia narrows as one moves southwards.
The shelf presents only isolated coralline algae and Bryozoa on the shelf as far as Cape
Frio (approximately 23∞S, 042∞W), Rio de Janeiro State (Melo et al., 1975; Rocha et al.,
1975). Corallines are rare southof Cape Frio (Rocha et al., 1975). This cape presents a
strong shift in oceanographical conditions, with strong upwelling bringing up much colder
waters (Castro et al., 1995).
Reef coral species have their southern limits of distribution in various localities be-
tween Abrolhos and Santa Catarina (Table 1). However, Laborel (1970) described larger
coral communities only as far south as the rocky shores of bays in Cape Frio, describing
this locality as a “coral oasis”. Coral cover on the rocky shores of this area is, however,
much lower (Castro et al., 1995) than on the rocky shores of the Abrolhos Archipelago
(Pitombo et al., 1988).
REGIONAL DIFFERENCES AND CONSERVATION OF BRAZILIAN REEFS
Brazilian reefs do not form a homogeneous entity and our knowledge of them varies
considerable. However, a few trends can be seen. The scattered information available for
reefs or banks in northern Brazil indicate offshore or oceanic communities dominated by
coralline algae, with S. stellata occurring as one of the main builders and/or the main
stony coral. This configuration was observed in higher intensity in Atol das Rocas (Kikuchi
and Leão, 1997; Echeverría et al., 1997). There are also qualitative or verbal reports of
similar situations in other reefs of the Cabo de São Roque reef group (Laborel, 1970;
Testa, 1997) and in the unexplored northern area (Laborel, 1970). The other reef group in
northern Brazil, Parcel do Manuel Luiz, seems to harbor a coral community established
on a rocky structure (Coura, 1994), but data on coral-algal cover is lacking. However, this
coral community seems to present a unique species composition. Northeastern reefs (Reefs
Coast) have mostly long lines of fringing reefs (many as coral communities growing on
367CASTRO AND PIRES: BRAZILIAN CORAL REEFS
beach rocks), with parallel lines offshore (although these are poorly mapped and de-
scribed). There are no studies on the structure or internal composition of biogenic (?)
reefs. Eastern reefs (Bahia State) occur in diverse forms, but this region is uniquely char-
acterized by the occurrence of the major reef-builder species M. braziliensis (endemic in
this region) and the presence of chapeirões as an important reef structure.
Knowledge on rates of sedimentation and sediment transport, their biological effects
on Brazilian corals, and oceanographic data on the reef areas, such as temperature, water
transparency, salinity and water circulation, is almost completely lacking or is restricted
to isolated or sparse readings in all regions. There are very few analyses of core drillings:
in Atol das Rocas (1), in Bahia (3), and shallow cores in the Cape of São Roque.
Among the three major regions mentioned here, the reefs of the northeastern and part
of the eastern regions (first group) are adjacent or near the shoreline and close to areas
with strong coastal zone development. The other regions (northern and the south part of
the eastern) present mostly offshore reefs in less populated areas. Fishing, tourism (mostly
in summer), and industrial activities are problems to be addressed in conservation efforts
for all these areas. However, urban pollution, and the presence of large numbers of visi-
tors are additional issues in the first group. As there are no comprehensive assessment
and/or monitoring programs being undertaken (except for the recent AGRA in restricted
areas of northern Bahia), most reports of reefs in an unhealthy condition are based on
subjective observations.
There is also concern about the introduction of alien coral species. In the late 80s, a few
colonies of Tubastraea were recorded on offshore oil platforms, in Campos, northern Rio
de Janeiro State (unpubl. data). This species is now found on many rocky shores south of
Rio de Janeiro (P. S. Young and F. B. Pitombo, pers. comm.). Recently, an oil platform in
Campos was found almost completely covered by corals of this genus (unpubl. data from
January 1999). Also, large colonies of soft coral (possibly Stereonephthya, L. van Ofwegen,
pers. comm.) were observed in Arraial do Cabo (near Cape Frio, Rio de Janeiro State) in
1999, which might have been transported by oil platforms also. Oil prospecting and ex-
ploration will be increasing in the Bahia State shelf, as Brazil is currently opening up
these activities to private initiatives from all over the world. There is concern about the
consequences of introducing alien species into Brazilian low diversity coral communi-
ties.
There are few marine conservation units and they usually suffer from severe funding
problems. Lack of resources forces decision makers to neglect unsustainable areas or
activities. A proper assessment is needed (and usually missing) in order to prepare sound
management plans for these units. Reef mapping and community distribution are basic
data not available for most areas. This is particularly true in the Abrolhos area, due to its
large size. It is difficult to select and to justify new areas for conservation without a prior
knowledge of their characteristics and their current health status. Restoration attempts
are just starting (APA Costa dos Corais and APA Recife de Fora) and, so far, they have
been restricted to closing areas to visitors, with little enforcement of the corresponding
regulations.
The study of Brazilian reefs has taken place mostly on a sporadic basis (very few
researchers working simultaneously). Although there is a lack of information on almost
all aspects of reef studies, the current (1990’s) surge of papers, by various researchers in
different areas and subjects (see section Literature Cited), indicate that, for the first time
in the history of Brazilian reef studies, the accumulation of knowledge has an opportu-
368 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
nity to prosper within a multidisciplinary environment, with peer discussions. Also, the
foundation of the Brazilian Society for Reef Studies (CORALLUS) has accelerated the
exchange of data and ideas among Brazilian researchers. Most researchers working to-
day in reef environments have some degree of training or experience overseas. Besides
learning from this international experience, foreign researchers may also be attracted to
Brazil, further accelerating the production of knowledge. Most current researchers have
also been trying to collaborate with conservation efforts along the Brazilian coast. All
these elements may help to strengthen the participation of science in Brazilian reef con-
servation.
ACKNOWLEDGMENTS
We thank the following persons and institutions: B. Segal, F. Pitombo, G. Dias, M. Torres, M.
Coura, M. Maida, P. Young, R. Kikuchi and Z.Leão, for information on different reef areas or coral
communities. The above-mentioned and J. Creed, M. Figueiredo, F. Amaral and V. Testa, for infor-
mation on their current research projects. G. Gilbert, for reviewing the English version. FAPERJ,
CNPq, UFRJ, and FUJB for grants and fellowships awarded to the Setor de Celenterologia,
Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro.
LITERATURE CITED
Amaral, F. D. 1997. Milleporidae (Cnidaria, Hydrozoa) do Litoral Brasileiro. Ph.D. Dissertation.
Univ. São Paulo, São Paulo. 204 p.
Barreto, L. A., J. D. Milliman, C. A. B. Amaral and O. Francisconi. 1975. Part II. Northern Brazil.
Pages 11–43 in J. D. Milliman and C. P. Summerhayes, eds. Upper continental margin sedimen-
tation off Brazil. Contr. Sediment. 4: 11–43.
Bayer, F. M. 1959. Octocorals from Surinam and adjacent coasts of South America. Uitg. natuurw.
Studkring Suriname. (Stud. Fauna Suriname) (6): 1–43, figs. 1–21.
Branner, J. C. 1904. The stone reefs of Brazil, their geological and geographical relations, with a
chapter on the coral reefs. Bull. Mus. Comp. Zool. Harv. 54: 1–285.
Castro, C. B. 1989. A new species of Plexaurella Valenciennes, 1855 (Coelenterata, Octocorallia),
from the Abrolhos Reefs, Bahia, Brazil. Revta bras. Biol. 49(2): 597–603.
__________. 1990. Revisão Taxonômica dos Octocorallia(Cnidaria, Anthozoa) do Litoral Sul–
americano: da foz do Rio Amazonas à foz do Rio da Prata. Ph.D. Dissertation. Univ. São Paulo,
São Paulo. 343 p.
__________. 1994. Corals of Southern Bahia. Pages 160–176 in B. Hetzel and C. B. Castro, eds.
Corals of southern Bahia. Nova Fronteira, Rio de Janeiro.
__________, C. A. Echeverría, D. O. Pires, B. J. A. Mascarenhas and S. G. Freitas. 1995. Distribuição
de Cnidaria e Echinodermata no infralitoral de costões rochosos de Arraial do Cabo, Rio de
Janeiro, Brasil. Revta bras. Biol. 55(3): 471–480.
__________ and D. O. Pires. 1999. A bleaching event in a Brazilian Reef. Revta bras. Oceanogr.
47(1): 87–90.
Coura, M. F. 1994. Contribuição ao Plano de Manejo do Parque Estadual do Parcel de Manuel Luiz.
Graduate Dissertation. Univ. Federal do Maranhão, São Luís. 55 p.
Coutinho, R., R. C. Villaça, C. A. Magalhães, M. A. Guimarães, M. Apolinário and G. Muricy.
1993. Influência antrópica nos ecossistemas coralinos da região de Abrolhos, Bahia, Brasil.
Acta biol. Leopoldensia 15(1): 133–144.
Creed, J. C. and G. M. Amado Filho. 1999. Disturbance and recovery of the macroflora of a seagrass
(Halodule wrightii Ascherson) meadow in the Abrolhos Marine National Park, Brazil: an ex-
perimental evaluation of anchor damage. J. Exp. Mar. Biol. Ecol. 235: 285–306.
http://www.ingentaconnect.com/content/external-references?article=0022-0981(1999)235L.285[aid=7502300]
http://www.ingentaconnect.com/content/external-references?article=0022-0981(1999)235L.285[aid=7502300]
http://www.ingentaconnect.com/content/external-references?article=0022-0981(1999)235L.285[aid=7502300]
http://www.ingentaconnect.com/content/external-references?article=0343-4125(1975)4L.11[aid=7502302]
http://www.ingentaconnect.com/content/external-references?article=0343-4125(1975)4L.11[aid=7502302]
http://www.ingentaconnect.com/content/external-references?article=0343-4125(1975)4L.11[aid=7502302]
369CASTRO AND PIRES: BRAZILIAN CORAL REEFS
Echeverría, C. A., D. O. Pires, M. S. Medeiros and C. B. Castro. 1997. Cnidarians of the Atol das
Rocas, Brazil. Proc. 8th Int’l. Coral Reef Symp., Panama. 1: 443–446.
Eston, V. R., A. E. Migotto, E. C. Oliveira Filho, S. A. Rodrigues and J. C. Freitas. 1986. Vertical
distribution of benthic marine organisms of the Fernando de Noronha Archipelago, Brazil. Bolm
Inst. oceanogr. 34: 37–53.
Ferreira, C. E. L. and J. E. A. Gonçalves. 1999. The unique Abrolhos reef formation (Brazil): need
for specific management strategies. Coral Reefs 18: 352.
Figueiredo, M. A. O. 1997. Colonization and growth of crustose coralline algae in Abrolhos, Brazil.
Proc. 8th Int’l. Coral Reef Symp., Panama 1: 689–694.
Gherardi, D. F. M. 1995. A windward coralline algal ridge on Rocas atoll, Atlantic Ocean. Coral
Reefs 14: 34.
Hartt, C. F. 1870. Geology and physical geography of Brazil. Osgood and Co., Boston, Fields.
620 p.
Hudson, M. M., F. D. Amaral, and M. F. Coura. 1999. Atualização do levantamento dos cnidários
bentônicos do Parque Estadual Marinho do Parcel do Manuel Luiz—MA. Page 180 In XII
Encontro de Zoologia do Nordeste, Abstracts.
Kempf, M. and J. Laborel. 1968. Formations de vermets et d’algues calcaires sur les cotes du
Brésil. Recl. Trav. Stn mar. Endoume. 43(50): 9–23.
Kikuchi, R. K. P. and Z. M. A. N. Leão. 1997. Rocas (southwestern equatorial Atlantic, Brazil): an
atoll built primarily by coralline algae. Proc. 8th Int’l. Coral Reef Symp., Panama 1: 731–736.
______________ and _____________. 1998. The effects of Holocene sea level fluctuations on reef
development and coral community structure, Northern Bahia, Brazil. Anais Acad. bras. Ciênc.
70(2): 159–171
Knoppers, B., J. 1996. Particulate organic matter and nutrients. Pages 36–37 in W. Ekau and B.
Knoppers, comps. Sedimentation processes and productivity in the continental shelf waters of
East and Northeast Brazil. Joint Oceanographic Projects. JOPS II. Cruise report and first re-
sults. Center for Tropical Marine Ecology, Bremen.
Laborel, J. 1969. Madreporaires et hydrocorallaiaires récifaux des côtes Brasiliennes. Systématic,
écologie, répartition verticale et geographique. Annls Inst océanogr. Paris. 47: 171–229
________. 1970. Les peuplements de madréporaires des cotes tropicales du Brésil. Annls Univ.
Abidjan (série E) 2(3): 1–260.
________ and M. Kempf. 1966. Formações de vermetos e algas calcárias nas costas do Brasil.
Trabs Inst. oceanogr. Univ. Fed. Pe. 7/8: 33–50.
Leão, Z. M. A. N. 1982. Morphology, geology, and developmental history of the southernmost
coral reefs of Western Atlantic, Abrolhos Bank, Brazil. Ph.D. Dissertation. Univ. Miami, Mi-
ami, Florida. 218 p.
______________. 1994. The coral reefs of Southern Bahia. Pages 151–159 in B. Hetzel and C. B.
Castro. Corals of southern Bahia. Nova Fronteira, Rio de Janeiro.
______________. 1996. The coral reefs of Bahia: morphology, distribution and the major environ-
mental impacts. Anais Acad. bras. Ciênc. 68(3): 339–452.
______________, T. M. F. Araújo and M. C. Nolasco. 1988. The coral reefs off the coast of Eastern
Brazil. Proc. 6th Int’l. Coral Reef Symp., Australia 3: 339–347.
______________ and R. N. Ginsburg. 1997. Living reefs surrounded by siliciclastic sediments: the
Abrolhos coastal reefs, Bahia, Brazil. Proc. 8th Int’l. Coral Reef Symp., Panama 2: 1767–1772.
______________, R. K. P. Kikuchi, M. P. Maia and R. A. L. Lago. 1997. A catastrophic coral cover
decline since 3,000 years B. P., Northern Bahia, Brazil. Proc. 8th Int’l. Coral Reef Symp., Panama
1: 583–588.
______________, R. K. P. Kikuchi, V. Testa, M. Telles, J. Pereira, L. Dutra and C. Sampaio. 1999.
First coral reef assessment in the Southern Hemisphere applying the AGRA rapid protocol
(Caramuanas Reef, Bahia, Brazil). Page 122 in National Coral Reef Institute and NOVA South-
eastern University Oceanographic Center, Int’l. Conf. Scientific Aspects of Coral Reef Assess-
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1999)18L.352[aid=7502304]
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1999)18L.352[aid=7502304]
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1995)14L.34[aid=6258106]
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1995)14L.34[aid=6258106]
370 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 2, 2001
ment, Monitoring, and Restoration, 14–16 April, 1999, Program and Abstracts. Fort Lauder-
dale, Florida.
______________, M. D. Telles, R. Sforza, H. A. Bulhões and R. K. P. Kikuchi. 1994. Impact of
tourism development on the coral reefs of the Abrolhos area, Brazil. Pages A22–A28 in R. N.
Ginsburg, comp. Proc. colloq. on global aspects of coral reefs: health, hazards and history,
1993. Rosenstiel School of Marine and Atmospheric Sciences, Univ. Miami, Miami, Florida.
Lopes, R. and J. Dutz. 1996. Zooplancton. Pages 38–39 in W. Ekau and B. Knoppers, comps.
Sedimentation processes and productivity in the continental shelf waters of East and Northeast
Brazil. Joint Oceanographic Projects. JOPS II. Cruise report and first results. Center for tropi-
cal marine ecology, Bremen.
Mabesoone, J. M. 1964. Origin and age of the sandstone reefs of Pernambuco (northeastern Brazil).
J. Sed. Petrol. 34(4): 715–726.
Maida M. and B. P. Ferreira. 1995. Estudo preliminar sobre o assentamento de corais em um recife
na Baía de Tamandaré—PE. Bolm téc. cient. CEPENE 3(1): 23–36.
_______ and ____________. 1997. Coral reefs of Brazil: an overview. Proc. 8th Int’l. Coral Reef
Symp., Panama 1: 263–274.
_______, ____________ and C. Bellini. 1995. Avaliação preliminar do recife da Baía do Sueste,
Fernando de Noronha, com ênfase nos corais escleractíneos. Bolm téc. cient. CEPENE 3(1):
37–47.
Marone, E. and R. Camargo. 1996. Leg 3: physical oceanography. Pages 35–36 in W. Ekau and B.
Knoppers, comps. Sedimentation processes and productivity in the continental shelf waters of
East and Northeast Brazil. Joint Oceanographic Projects. JOPS II. Cruise report and first re-
sults. Centerfor Tropical Marine Ecology, Bremen.
Marques, A. C. S. J. and C. B. Castro. 1995. Muricea (Cnidaria, Octocorallia) from Brazil, with
description of a new species. Bull. Mar. Sci. 56: 161–172.
Medeiros, M. S. 1998. Octocorallia (Cnidaria: Anthozoa) da plataforma continental e ilhas oceânicas
brasileiras: famílias Paramuriceidae e Plexauridae. M.Sc. Dissertation. Museu Nacional, Univ.
Federal do Rio de Janeiro, Rio de Janeiro. 157 p.
_____________ and C. B. Castro. 1999. Paramuriceidae e Plexauridae (Cnidaria, Octocorallia) do
Brasil: batimetria e distribuição geográfica. Bolm Mus. nac. Rio de J. (398): 1–20.
Melo, U., C. P. Summerhayes and J. P. Ellis. 1975. Part 4. Salvador to Vitória, Southeastern Brazil.
Pages 78–116 in J. D. Milliman and C. Summerhayes, eds. Upper continental margin sedimen-
tation off Brazil. Contrib. Sediment. 4.
Milliman, J. D. and C. P. Summerhayes, eds. 1975. Upper continental margin sedimentation off
Brazil. Contrib. Sediment. 4. 175 p.
Moura, R. L., M. C. M. Rodrigues, R. B. Francini0-Filho and I. Sazima. 1999. Unexpected richness
of reef corals near the southern Amazon River mouth. Coral Reefs 18: 170.
Norse, E. A., ed. 1993. Global Marine Biological Diversity. A strategy for building conservation
into decision making. Washington, D. C., Center for Marine Conservation, Island Press. 383 p.
Paulay, G. 1997. Diversity and distribution of reef organisms. Pages 298–353 in C. Birkeland. Life
and death of coral reefs. Chapman and Hall, New York.
Pires, D. O., C. B. Castro, A. E. Migotto and A. C. Marques. 1992. Cnidários Bentônicos do
Arquipélago de Fernando de Noronha, Brasil. Bolm Mus. nac. Rio de J. (354): 1–21.
_________, __________ and C. C. Ratto. 1999. Reef coral reproduction in the Abrolhos reef com-
plex, Brazil: the endemic genus Mussismilia. Mar. Biol. 135(3): 463–471.
Pitombo, F. B., C. C. Ratto and M. J. C. Belém. 1988. Species diversity and zonation pattern of
hermatypic corals at two fringing reefs at Abrolhos Archipelago, Brazil. Proc. 6th Int’l. Coral
Reef Symp., Australia 2: 817–820.
Rocha, J., J. D. Milliman, C. I. Santana and M. A. Vicalvi. 1975. Pages 111–150 in J. D. Milliman
and C. Summerhayes, eds. Upper continental margin sedimentation off Brazil. Contrib. Sedi-
ment. 4
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1999)18L.170[aid=7507248]
http://www.ingentaconnect.com/content/external-references?article=0722-4028(1999)18L.170[aid=7507248]
http://www.ingentaconnect.com/content/external-references?article=0022-4472(1964)34L.715[aid=7507249]
http://www.ingentaconnect.com/content/external-references?article=0022-4472(1964)34L.715[aid=7507249]
http://www.ingentaconnect.com/content/external-references?article=0025-3162(1999)135L.463[aid=7507250]
http://www.ingentaconnect.com/content/external-references?article=0025-3162(1999)135L.463[aid=7507250]
http://www.ingentaconnect.com/content/external-references?article=0007-4977(1995)56L.161[aid=7507251]
http://www.ingentaconnect.com/content/external-references?article=0007-4977(1995)56L.161[aid=7507251]
371CASTRO AND PIRES: BRAZILIAN CORAL REEFS
Testa, V. 1997. Calcareous algae and corals in the inner shelf of Rio Grande do Norte, Brazil. Proc.
8th Int’l. Coral Reef Symp., Panama 1: 737–742.
Villaça, R. C. and F. B. Pitombo. 1997. Benthic communities of shallow–water reefs of Abrolhos,
Brazil. Revta bras. Oceanogr. 45(1/2): 35–43.
Wolanski, E. 1994. Physical oceanographic processes of the Great Barrier Reef., CRC Press, Boca
Raton, Florida. 194 p.
Young, P. S. 1986. Análise qualitativa e quantitativa da fauna associada a corais hermatípicos (Co-
elenterata, Scleractinia) nos recifes de João Pessoa, PB. Revta bras. Biol. 46(1): 99–126.
ADDRESS: Museu Nacional, Universidade Federal do Rio de Janeiro, Departamento de Invertebrados,
Quinta da Boa Vista, s/n∞, Rio de Janeiro, RJ, 20940–040, Brasil. E-mail: <dopires@openlink.com.br>.
CORRESPONDING AUTHOR: (C.B.C.) E-mail: <cbcastro@pobox.com>.