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61 Rapid Assessment Program
A Rapid Marine Biodiversity Assessm
ent of the Coral Reefs of Northeast M
adagascar
RAP
61ofBulletinBiological
Assessment
A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northeast Madagascar
EditorsDavid Obura, Giuseppe Di Carlo, Ando Rabearisoa and Thomas Oliver
CONSERVATION INTERNATIONAL – MADAGASCAR
CORDIO
THE JOHN D. AND CATHERINE T. MACARTHUR FOUNDATION
1514647819349
ISBN 978-1-934151-46-490000 >
A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northeast Madagascar
CONTENTSParticipants and Authors .............................................................. 4Organization Profiles ..................................................................... 5Acknowledgements ....................................................................... 7Executive Summary ....................................................................... 8Report at a Glance ....................................................................... 10Chapters ......................................................................................... 17Map and Photos ........................................................................... 25Appendices .................................................................................. 68
Conservation International2011 Crystal Dr., Suite 500Arlington, VA 22202 USA
TELEPHONE: +1 703 341-2400
WEB: www.conservation.org
with generous support from:
RAP61
of Bu
lletinB
iologicalA
ssessmen
t
CONSERVATION INTERNATIONAL – M
ADAGASCAR
CORDIO
THE JOHN D. AND CATHERINE T. MACARTHUR
FOUNDATION
Rapid Assessm
ent Program
A Rapid M
arine Biodiversity
Assessm
ent of the Coral Reefs of N
ortheast Madagascar
Ed
itors
David
Ob
ura, G
iusep
pe D
i Carlo
, A
nd
o R
abeariso
a and
Th
om
as Oliver
Th
e RA
P B
ulletin
of Biological A
ssessmen
t is pu
blish
ed b
y:C
onservation
Intern
ational
2011 C
rystal Drive, Su
ite 500
Arlin
gton, V
A 2
2202
USA
Tel: +
1 7
03-3
41-2
400
ww
w.con
servation.org
Editors: D
avid O
bu
ra, Giu
sepp
e Di C
arlo, An
do R
abearisoa, Th
om
as Oliver
Design
: Kim
Meek
Map: K
ellee Koen
ig, Con
servation In
ternation
alC
over photographs: [top to b
ottom] T
urbin
aria irregularis; sailin
g vessel, Diego b
ay; anem
one fi sh
. All p
hotos
by K
eith E
llenbogen
/ILC
P.
Con
servation In
ternation
al is a private, n
on-p
rofi t organization
exemp
t from fed
eral incom
e tax un
der section
501 c (3
) of the In
ternal R
evenu
e Cod
e.
ISBN
: 978-1
-934151-4
6-4
©2011 b
y Con
servation In
ternation
alA
ll rights reserved
.
Th
e design
ations of geograp
hical en
tities in th
is pu
blication
, and
the p
resentation
of the m
aterial, do n
ot
imp
ly the exp
ression of an
y opin
ion w
hatsoever on
the p
art of Con
servation In
ternation
al or its sup
portin
g
organization
s concern
ing th
e legal status of an
y coun
try, territory, or area, or of its auth
orities, or concern
ing
the d
elimitation
of its frontiers or b
oun
daries.
An
y opin
ions exp
ressed in
the R
AP
Bu
lletin of B
iological Assessm
ent are th
ose of the w
riters and
do n
ot n
ecessarily refl ect those of C
I.
Prin
ted b
y MY
E M
adagascar
RA
P B
ulletin
of Biological A
ssessmen
t was form
erly RA
P W
orking P
apers. N
um
bers 1
–13 of th
is series were
pu
blish
ed u
nd
er the p
revious title.
Suggested citation
: Obu
ra D., D
i Carlo, G
., Rab
earisoa, A. an
d O
liver, T. (ed
itors). 2011. A
Rap
id M
arine
Biod
iversity Assessm
ent of th
e coral reefs of north
east Mad
agascar. RA
P B
ulletin
of Biological A
ssessmen
t 61.
Con
servation In
ternation
al. Arlin
gton, V
A.
3A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Participants and Authors .......................................................4
Organization Profi les ..............................................................5
Acknow
ledgements ................................................................7
Executive Summ
ary ................................................................8
Report at a Glance .................................................................10
Chapter 1 ..................................................................................17Corals of northeast M
adagascar
David Obura
Map and Photos .....................................................................25
Chapter 2 ..................................................................................29Reef fi shes of northeast M
adagascar
Melita Sam
oilys & Bem
ahafaly Randriamanantsoa
Chapter 3 .................................................................................40Echinoderm
s from the north-east coast of M
adagascar
Jean Maharavo
Chapter 4 ..................................................................................44Seagrasses and algae of north-eastern M
adagascar
Giuseppe Di Carlo & M
onica Tombolahy
Chapter 5 ..................................................................................53Coral-associated exosym
bionts of northeast Madagascar
Sea McKeon
Chapter 6 ..................................................................................56Coral reef health and status
David Obura and Tom Oliver
Table of Contents
Appendix 1 ...............................................................................68
Coral species recorded by site, during surveys in northeast M
adagascar
Appendix 2 ...............................................................................76
Coral species lists from northeast M
adagascar, Madagascar
and the western Indian Ocean
Appendix 3 ...............................................................................85
List of coral species collected for taxonomy/genetic
analyses
Appendix 4 ...............................................................................87
Reef fi sh species list, northeast Madagascar
Appendix 5 ...............................................................................92
Echinoderm species list, northeast M
adagascar
Appendix 6 ...............................................................................94
Echinoderm distribution by site, northeast M
adagascar
Appendix 7 ...............................................................................98
Algae and seagrass species distribution by location, northeast M
adagsacar
4Rapid Assessm
ent Program
Gisèle B
akaryC
entre N
ational d
e Rech
erches O
céanograp
hiq
ues (C
NR
O)
P.O B
ox 68
Hell-V
ille, 207 N
osy-Be
Mad
agascar
Giu
sepp
e DiC
arloC
onservation
Intern
ational
2011 C
rystal Drive, Su
ite 500
Arlin
gton V
A 2
2202 U
SA
Jean M
aharavo, P
hD
Cen
tre Nation
al de R
echerch
e sur l’E
nviron
nem
ent
(CN
RE
)39, R
asamim
anan
a St., Fiad
anan
aP
O B
ox 1739
An
tanan
arivo 101
Mad
agascar Sea M
cKeon
Florid
a Mu
seum
of Natu
ral History
Gain
esville, FL
32601
USA
David
Ob
ura
CO
RD
IO E
ast Africa
#9 K
ibaki F
lats, Ken
yatta Beach
, Bam
bu
ri Beach
P.O.B
OX
10135 M
ombasa 8
0101, K
enya
Em
ail: dob
ura@
cordioea.org
Kirsten
L.L
. Oleson
, Ph
.D.
NSF
Intern
ational P
ostdoctoral Sch
olarc/o C
enter on
Eth
ics in Society
Stanford
Un
iversity482 G
alvez Mall
Stanford
, CA
94305-6
079
USA
Tom
An
drew
Oliver
Hop
kins M
arine Station
P
acifi c Grove, C
A 9
4305 U
SA
Participants and Authors
An
do R
abearisoa
Con
servation In
ternation
alM
adagascar C
oun
try Program
P.O B
ox 5178, E
xplorer B
usin
ess Park
An
korond
rano, 1
01 A
ntan
anarivo
Mad
agascar E
rcilla Rad
yM
aster Stud
ent in
Econ
omics
Un
iversity of Toam
asina
P.O B
ox 591
501 T
oamasin
aM
adagascar
Bem
ahafaly R
and
riaman
antsoa
Wild
life Con
servation Society
Mad
agascarP
O B
ox 8500 Soavim
bah
oakaA
ntan
anarivo 1
01
Mad
agascar M
elita Samoilys
CO
RD
IO E
ast Africa
P.O.B
OX
24562, K
aren 0
0502, N
airobi, K
enya
Em
ail: melita.sam
oilys@gm
ail.com
Mon
ica Lu
cie Tom
bolah
yC
onservation
Intern
ational M
adagascar
P.O B
ox 5178, E
xplorer B
usin
ess Park
An
korond
rano, A
ntan
anarivo 1
01
Mad
agascar
5A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
CON
SERVATION
INTERN
ATION
AL
Con
servation In
ternation
al is an in
ternation
al non
profi t organ
ization b
ased in
Virgin
ia, U
nited
States of Am
erica. At C
onservation
Intern
ational w
e are comm
itted to restorin
g and
m
aintain
ing th
e world
’s natu
ral systems. Th
rou
gh scien
tifi c rigor, creative thin
king an
d
aggressive action, w
e are provin
g that sp
ecies can b
e saved, lan
dscap
es and
seascapes can
be
protected
and
peop
le can th
rive. CI’s m
ission: B
uild
ing u
pon
a strong fou
nd
ation of scien
ce, p
artnersh
ip an
d fi eld
dem
onstration
, CI em
pow
ers societies to respon
sibly an
d su
stainab
ly care for n
ature, ou
r global b
iodiversity, for th
e well-b
eing of h
um
anity.
Con
servation In
ternation
al (HQ
)2011 C
rystal Drive, Su
ite 500
Arlin
gton, V
A 2
2202 U
SAh
ttp://w
ww
.conservation
.org C
onservation
Intern
ational (M
adagascar)
PO
Box 5
178
Batim
ent C
2, E
xplorer B
usin
ess Park
An
korond
rano, A
ntan
anarivo 1
01
Mad
agascar
CORD
IO
CO
RD
IO E
ast Africa is a research
organisation
focused
on m
arine an
d coastal ecosystem
s in
the W
estern In
dian
Ocean
. Its mission
is to generate an
d sh
are scientifi cally sou
nd
know
ledge
for develop
ing solu
tions to th
e prob
lems an
d ch
allenges facin
g coastal and
marin
e environ
-m
ents an
d p
eople in
the W
estern In
dian
Ocean
.
#9 K
ibaki F
lats, Ken
yatta Beach
, Bam
bu
ri Beach
P.O.B
OX
10135
Mom
basa 8
0101, K
enya
ww
w.cord
ioea.org
Organization Profi les
Organization Profiles
6Rapid Assessm
ent Program
NATIO
NA
L CENTER FO
R ENVIRO
NM
ENTA
L RESEARCH
(CEN
TRE NATIO
NA
L DE RECH
ERCHE SU
R L’ENVIRO
NN
EMEN
T)
Th
e Nation
al Cen
ter for En
vironm
ental R
esearch or C
entre
Nation
al de R
echerch
e sur l’E
nviron
nem
ent (C
NR
E) is
one of M
adagascar ‘s n
ational cen
ters un
der su
pervision
of th
e Min
istry of Nation
al Ed
ucation
and
Scientifi c R
esearch
or Min
istère de l’E
du
cation N
ationale et d
e la Rech
erche
Scientifi q
ue (M
EN
RE
S). It is a pu
blic in
stitution
of an
ind
ustrial an
d com
mercial n
ature. Its m
ission is to carry
out m
arine as w
ell as terrestrial environ
men
tal research
program
s, to provid
e tools for environ
men
tal man
agemen
t in
Mad
agascar. C
entre N
ational d
e Rech
erche su
r l’En
vironn
emen
t (CN
RE
)P
O B
ox 1739
Fiad
anan
a, An
tanan
arivo 101
Mad
agascar
NATIO
NA
L CENTER FO
R OCEA
NO
GRA
PHIC RESEA
RCH
(CENTRE N
ATION
AL D
E RECHERCH
ES OCEA
NO
GRA
PHIQ
UES)
Th
e Nation
al Cen
ter for Ocean
ograph
ic Research
or Cen
tre N
ational d
e Rech
erches O
céanograp
hiq
ues (C
NR
O) is in
ch
arge of the m
arine research
in M
adagascar. A
mon
g its m
ain activities are m
arine b
iodiversity in
ventorys, stock
assessmen
ts of exploited
and
un
der-exp
loited m
arine
resources, an
d th
e mon
itoring of th
e states of marin
e and
coastal ecosystem
s, particu
larly coral reefs. Th
e Nation
al C
enter for O
ceanograp
hic R
esearch is b
ased in
Nosy-B
e, in
North
western
Mad
agascar. C
entre N
ational d
e Rech
erches O
céanograp
hiq
ues
PO
Box 6
8N
osy-Be 2
07
Mad
agascarE
mail : cn
ronosyb
e@m
oov.mg
7A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Th
is work w
as a collaborative eff ort b
etween
Con
servation In
ternation
al, CO
RD
IO, th
e N
ational C
enter for E
nviron
men
tal Research
, the N
ational C
enter for O
ceanograp
hic
Research
and
the W
ildlife C
onservation
Society. Con
servation In
ternation
al wish
es to than
k all th
e particip
ants to th
e RA
P exp
edition
, the cap
tain an
d crew
aboard
the R
/V A
ntsiva an
d
Keith
Ellen
bogen
(Blu
e Reef) for h
is ph
otograph
ic contrib
ution
. CI also w
ishes to th
ank
all the staff in
volved in
several stages of the p
roject, particu
larly Kellee K
oenig an
d L
eeann
e A
lonso. W
e also than
k Dr D
avid K
ingston
from V
irginia P
olytechn
ic Institu
te and
State U
niversity an
d P
rogram L
eader of IC
BG
Mad
agascar, and
An
driam
bolan
tsoa Rasoloh
ery and
F
inoan
a Rollan
de from
CI M
adagascar for th
e Map
pin
g. Fin
ally we are gratefu
l to the review
-ers for th
eir invalu
able feed
back on
the ch
apters of th
is report.
Th
is work w
as mad
e possible by th
e support of th
e Th
e John
D. an
d C
atherin
e T. M
acArth
ur
Foun
dation
.
Acknow
ledgements
8Rapid Assessm
ent Program
EXECUTIVE SU
MM
ARY
In an
eff ort to assess new
biod
iversity hotsp
ots and
climate resilien
t areas, Con
servation In
ter-n
ational con
du
cted a th
ird m
arine R
apid
Assessm
ent P
rogram (R
AP
) along th
e north
eas coast of M
adagascar, su
rveying th
e coastline from
the B
ay of Am
bod
ivahib
e to Voh
emar, coverin
g ap
proxim
ately 130 km
of coast. Th
e two p
rior marin
e RA
P su
rveys in M
adagascar exten
ded
from
Nosy B
e to Nosy H
ara on th
e north
west coast in
2002 (M
cKen
na et al. 2
005, R
AP
#31),
and
from C
ap d
’Am
bre to A
mbod
ivahib
e, along th
e north
east coast, in 2
007 (M
aharavo et
al. in p
ress). Th
is third
RA
P w
as aimed
at extend
ing th
e know
ledge on
Mad
agascar’s marin
e biod
iversity along th
is previou
sly un
-surveyed
stretch of coast as w
ell as to better u
nd
erstand
th
e vuln
erability of th
is region’s marin
e coastal environ
men
ts to climate ch
ange.
Five location
s were su
rveyed, A
mbod
ivahib
e, Loky B
ay, An
kao island
, An
dravin
a Bay an
d
Voh
emar, w
hich
inclu
ded
a mix of d
eep an
d sh
allow b
ays, and
an islan
d system
, separated
by stretch
es of exposed
linear coastlin
e. Coral reefs in
the region
were gen
erally health
y, with
coral cover averagin
g 50%
in ap
prop
riate locations, an
d seagrass b
eds ap
pearin
g in good
con
dition
. Th
e two m
ain en
vironm
ental d
rivers in th
e coastal environ
men
t in th
e areas were
sedim
entation
- particu
larly in th
e bays - an
d h
igh exp
osure to w
ind
and
waves from
the
south
ern In
dian
Ocean
.Th
e su
rvey covered coral reefs an
d associated
fi sh, ech
inod
erms, an
d algae an
d seagrass b
eds
and
foun
d th
e followin
g species rich
ness in
targeted grou
ps: algae (9
1), seagrasses (10), h
ard
corals (276), coral sym
bion
ts (67), ech
inod
erms (7
0) an
d fi sh
(271, ou
t of a restricted set of
19 fam
ilies).Th
e rich
ness of h
ard corals w
as less than
the 3
23 sp
ecies foun
d on
a broad
er stretch of th
e n
orthw
est coast usin
g more com
preh
ensive taxon
omic m
ethod
s by V
eron an
d T
urak (2
005).
At a region
al scale, usin
g species rich
ness p
redicted
from accu
mu
lation cu
rves, North
east Ma-
dagascar grou
ps w
ith oth
er sites of high
est diversity, in
clud
ing n
orthw
est Mad
agascar, north
M
ozambiq
ue an
d sou
th T
anzan
ia (> 2
50 sp
ecies) comp
ared to 1
70-2
40 sp
ecies foun
d in
the
island
group
s of the Seych
elles and
Com
oros, and
north
ern K
enya to D
jibou
ti. Th
ese fi nd
ings
lend
sup
port to th
e existence of a h
igh d
iversity center for th
e Western
Ind
ian O
cean (W
IO)
encom
passin
g the n
orthern
tip of M
adagascar an
d exten
din
g westw
ards across th
e north
ern
Mozam
biq
ue ch
ann
el. Fish
diversity w
as similarly h
igh for th
e region, record
ing 7
4%
of the
367 sp
ecies reported
for all of Mad
agascar since 1
891. Th
e rich
ness of coral exosym
bion
ts was
high
for the am
oun
t of samp
ling con
du
cted, w
ith u
nrecogn
ized in
divid
uals p
otentially from
u
nd
escribed
species, p
ointin
g to an u
nkn
own
and
poten
tially rich b
iodiversity of cryp
tic spe-
cies in th
e region.
Alth
ough
generally in
good h
ealth, th
e surveyed
hab
itats are not p
ristine. A
lgae, echin
oderm
an
d fi sh
species p
resence w
ere ind
icative of hu
man
imp
acts. Sea urch
ins an
d exten
sive macro/
epip
hytic algae are in
dicative of d
isturb
ed con
dition
s, and
were m
ost abu
nd
ant in
the b
ay of V
ohem
ar du
e to urb
an p
ollution
causin
g high
nu
trient levels. Th
e ab
sence of sea cu
cum
-bers an
d large fi sh
(sharks, grou
pers, N
apoleon
wrasse, B
um
ph
ead p
arrotfi sh) w
as ind
icative
Executive Summ
ary
Executive Summ
ary
9 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
of high
levels of exploitation
from local villages as w
ell as m
igrant fi sh
ers based
in th
e larger centers su
ch as D
iego and
V
ohem
ar. A
coral bleach
ing even
t was u
nd
erway d
urin
g the exp
edi-
tion, w
ith 2
010 p
otentially b
illed as th
e worst b
leachin
g even
t since 1
998 in
the W
estern In
dian
Ocean
. Th
is enab
led
a test of the h
ypoth
esis of protection
from w
armin
g at sites su
ch as A
mbod
ivahib
e and
Loky B
ay. Ap
art from a sin
gle sh
allow site w
ith 8
0%
cover of staghorn
Acropora in
Loky
Bay w
ith 5
0%
of corals bleach
ed, b
leachin
g was clearly
wid
espread
bu
t at low overall rates (5
% of colon
ies aff ected),
with
almost zero m
ortality observed
. Th
is was com
parab
le to m
ost parts of th
e WIO
and
less than
the w
orst-aff ected sites
nearby in
the C
omoros (M
ayotte), so there is h
igh likeli-
hood
that b
leachin
g protection
is active in th
e region.
Con
servation recom
men
dation
s inclu
de th
e followin
g:
1)
ES
TA
BL
ISH
new
Marin
e Protected
Areas, focu
sed on
rep
resentatiion
of diff eren
t hab
itats and
locations,
taking ad
vantage of clim
ate protection
from u
pw
elling
and
with
a prim
ary objective to en
han
ce and
sustain
local livelih
oods.
2)
EN
HA
NC
E fi sh
eries man
agemen
t. Th
e imp
act of local an
d m
igrant fi sh
ing, for food
and
comm
ercial markets,
were clear even
in th
e most rem
ote and
least accessible
locations on
the su
rvey.
3)
PR
OM
OT
E w
atershed
man
agemen
t to redu
ce sedi-
men
t imp
acts on m
arine system
s.
4)
IMP
LE
ME
NT
species con
servation p
lans, for fl agsh
ip
species (e.g. sh
arks, sea turtles, N
apoleon
wrasse, B
um
p-
head
parrotfi sh
and
large group
ers) as well as evolu
tio-n
arily distin
ct species en
dem
ic to the W
estern In
dian
O
cean (e.g. th
e corals Craterastrea laevis, A
nom
astrea irregu
laris, Horastrea in
dica and
Gyrosm
ilia interru
pta).
5)
EX
PL
OR
E eff ective p
artnersh
ips for con
servation,
engagin
g with
local governm
ent, local com
mu
nities an
d
private sector in
terests in th
e area.
6)
CO
ND
UC
T fu
rther b
iological assessmen
ts to fi ll gaps
in in
formation
, most evid
ent for on
the N
orthw
est coast to th
e Barren
Island
s in th
e Mozam
biq
ue ch
ann
el, a stretch
of 1,0
00 km
of coastline.
7)
DE
SIG
N gu
idelin
es and
best p
ractices for coastal and
foresh
ore develop
men
t plan
nin
g relevant to th
is region
of Mad
agascar.
10Rapid Assessm
ent Program
INTRO
DU
CTION
To fi ll th
e gap in
biological in
formation
, both
nation
ally for Mad
agascar and
for the b
roader
Western
Ind
ian O
cean (W
IO) region
, Con
servation In
ternation
al cond
ucted
a marin
e Rap
id
Assessm
ent P
rogram (R
AP
) in th
e North
eastern C
oast of Mad
agascar from th
e Bay of A
mbod
i-vah
ibe to V
ohem
ar. Th
is was a th
ird in
itiative followin
g two oth
er marin
e RA
Ps. Th
e fi rst cov-
ered ap
proxim
ately 200 km
of coastline from
Nosy B
e to Nosy H
ara on th
e north
west coast in
2002 (M
cKen
na et al. 2
005). Th
e secon
d R
AP
covered ap
proxim
ately 50 km
of coastline from
C
ap d
’Am
bre at th
e north
ern tip
of Mad
agascar to Am
bod
ivahib
e, dow
n th
e North
east coast, in
2007 (M
aharavo et al. in
press). Th
is su
rvey covered ap
proxim
ately 130 km
of coast. For a
thorou
gh review
of the existin
g marin
e biod
iversity stud
ies in M
adagascar, an
d th
e institu
tional
and
policy con
text for the th
ree exped
itions in
sup
port of M
PA
plan
nin
g and
selection, see
McK
enn
a et al. (2005).
Mad
agascar is the w
orld’s fou
rth largest islan
d, coverin
g 587,0
45 sq
uare kilom
eters, and
with
a coastlin
e length
exceedin
g 5,0
00 km
and
an estim
ated len
gth of all reefs (in
clud
ing frin
ging
reefs, island
s, platform
s, and
both
emergen
t and
subm
erged b
arrier reefs) of 3,4
59 km
(Cooke
et al. 2000). C
oastal marin
e hab
itats are pred
omin
antly m
angroves, seagrasses an
d coral reefs,
with
an exp
osed rocky sh
ore along th
e eastern ocean
-facing coast. P
ast work h
as coun
ted over
320 coral sp
ecies, and
over 750 reef-associated
fi sh sp
ecies (McK
enn
a et al. 2005). In
1999,
an ecoregion
al app
roach w
as prop
osed b
y the A
ssociation N
ationale p
our la G
estion d
es Aires
Protégées (A
NG
AP
2001) for th
e establish
men
t of Marin
e Protected
Areas, later exp
and
ed
on th
rough
the estab
lishm
ent in
2006 of an
inter-m
inisterial b
ody, th
e Com
mission
En
vi-ron
nem
ent-P
eche (C
EP
). In p
artnersh
ip w
ith in
ternation
al conservation
NG
Os, 2
0 p
otential
MP
As w
ere initially id
entifi ed
based
on th
eir extraordin
ary biod
iversity or the p
resence of
special organ
isms su
ch as tu
rtles, lemu
rs, and
sea bird
s. E
xisting th
reats to Mad
agascar’s marin
e biod
iversity inclu
de fi sh
ing, sed
imen
tation from
rivers, coastal d
evelopm
ent an
d p
ollution
, and
increasin
gly oil and
gas exploration
and
mi-
nin
g (WW
F 2
010). G
rowth
of coastal pop
ulation
s is taking p
lace more rap
idly th
an across th
e cou
ntry as a w
hole, w
ith w
omen
giving b
irth to an
average of > 6
child
ren in
coastal provin
ces. M
ost coastal comm
un
ities are poor an
d h
ighly d
epen
den
t up
on fi sh
eries for survival an
d liveli-
hood
s. Large-scale d
evelopm
ent p
rograms are b
eing p
lann
ed for oil an
d gas exp
loitation in
the
western
Mad
agascar and
Mozam
biq
ue C
han
nel, w
ith exp
loration con
cessions alread
y parceled
ou
t. Sedim
entation
from d
eforestation on
land
has lon
g been
recognized
as a prim
ary threat
to marin
e resources. Sin
ce coastal and
off shore m
arine resou
rces are major con
tribu
tors to the
nation
al econom
y, as a source of food
for their p
eople, an
d th
rough
tourism
, comm
ercial fi -sh
ing agreem
ents, an
d reven
ues from
off shore oil an
d gas d
evelopm
ent, p
ressures are exp
ected
to increase w
ith b
oth p
opu
lation grow
th an
d in
creasing d
evelopm
ent, an
d th
e likely increase in
in
equ
itable d
istribu
tion of w
ealth an
d resou
rces.C
limate ch
ange h
as becom
e an overarch
ing con
cern for coral reefs glob
ally (Hoegh
-Gu
lberg
et al. 2007). In
1998, th
e WIO
show
ed th
e high
est levels of coral bleach
ing in
respon
se to clim
ate chan
ge globally (W
ilkinson
2004). M
adagascar’s coral reefs b
leached
du
ring th
e Ind
ian
Report at a Glance
Report at a Glance
11 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Ocean
-wid
e event of 1
998 as w
ell as in sm
aller episod
es in
the last 3
-5 years (L
ind
en et al. 2
002, W
ilkinson
2004).
How
ever the p
attern of b
leachin
g has varied
, with
high
levels of b
leachin
g and
mortality in
the sou
thw
est, bu
t low or
variable m
ortality recorded
in th
e north
west an
d n
ortheast
(Qu
od et al. 2
002, W
ebster et al. 2
002). A
reef suscep
tibi-
lity mod
el that b
roadly m
atches p
atterns of coral b
leachin
g in
other p
arts of the W
IO (M
aina et al. 2
008, M
cClan
ahan
et al. 2
009), p
redictin
g lower b
leachin
g suscep
tibility in
cooler w
aters to the sou
th is con
trary to the p
attern fou
nd
in
Mad
agascar. Th
ere are several possib
le reasons for th
is d
isagreemen
t as man
y factors infl u
ence m
ultip
le stages of the
bleach
ing p
rocess (Obu
ra 2005). T
wo key on
es that m
ay be
imp
ortant to th
is region are a) reefs in
the sou
thw
est were
already m
ore high
ly degrad
ed th
an reefs in
the n
orthw
est from
local anth
ropogen
ic stresses, wh
ich can
increase vu
lne-
rability to b
leachin
g, and
b) w
hile sea su
rface temp
eratures
(SST) closer to th
e equ
ator are high
er than
those in
sub-
tropical latitu
des, corals acclim
ate locally (Coles an
d B
rown
2003), h
ence th
e absolu
te temp
erature m
ay be less im
por-
tant th
an sh
ort term variab
ility in tem
peratu
re. In
2008 a series of in
itiatives oriented
aroun
d th
e imp
acts to m
arine p
rotected area (M
PA
) plan
nin
g for climate ch
ange
were con
du
cted (C
I 2009, M
aina an
d O
bu
ra 2008). A
t the
same tim
e the secon
d m
arine R
AP
(Mah
aravo et al. in p
ress) id
entifi ed
a site poten
tially protected
from w
armin
g sea sur-
face temp
eratures b
y strong u
pw
elling, at A
mbod
ivahib
e Bay,
south
of An
tsiranan
a Bay (D
iego Suarez) on
the N
ortheast
coast. Billed
as a ‘climate p
ark’, it prom
oted fu
rther in
centive
for this R
AP
to not on
ly furth
er inven
tory and
assess Mad
a-gascar’s m
arine b
iodiversity, b
ut also to d
etermin
e if natu
ral clim
ate resilience is a gen
eral cond
ition for th
e north
west
coast, with
up
wellin
g (West an
d Salm
2003, O
bu
ra 2005)
provid
ing a refu
ge for corals un
der clim
ate chan
ge.Th
is R
AP
survey h
ad th
e followin
g objectives:
1)
High
light th
e richn
ess of marin
e biod
iversity along th
e n
ortheast M
adagascar to raise aw
areness am
ong th
e local com
mu
nities as w
ell as the glob
al general p
ublic,
2)
Collect b
iodiversity d
ata to assist in gu
idin
g conserva-
tion action
s and
marin
e priority sites id
entifi cation
in
the region
,
3)
Assess th
e vuln
erability of coral reefs in
this p
art of M
adagascar to w
armin
g sea surface tem
peratu
res, and
to u
nd
erstand
the con
nectivity w
ith oth
er marin
e ecosys-tem
s (e.g. seagrass),
4)
En
han
ce scientifi c cap
acity in M
adagascar by d
evel-op
ing collab
orations b
etween
intern
ational an
d local
scientists.
Th
is exped
ition d
eployed
a mu
lti-discip
linary team
of 9 m
arine scien
tists and
coastal resource exp
erts to sur-
vey selected areas, an
d w
as in th
e fi eld from
28 M
arch to
14 A
pril 2
010. Su
rveys were focu
sed in
bay an
d islan
d sys-
tems alon
g the n
ortheast coast, at A
mbod
ivahib
e Bay, L
oky B
ay, Nosy A
nkao, A
nd
ravina B
ay and
Voh
emar.
Survey sitesTh
e stu
dy region
is app
roximately 1
30 km
from n
orth to
south
, along th
e north
ern en
d of th
e north
east coast of M
adagascar. A
total of 24 stu
dy sites w
ere surveyed
for coral reef fau
na w
ith an
add
itional 2
-3 reef fl at su
rveys for echi-
nod
erms. A
lgae and
seagrass assessmen
ts were con
du
cted at
some 1
5 sites ad
jacent to th
e coral reef survey sites (T
able 1
). Th
e sites grou
ped
natu
rally into 5
geograph
ic locations alon
g th
e coast, from n
orth to sou
th:
Ambodivahibe Bay is a n
arrow b
ay in th
e form of a d
eep
canyon
, pen
etrating in
to the lan
d ab
out 3
km. Its sid
es are steep
ly slopin
g, and
the b
ottom varies from
abou
t 200m
d
eep at th
e mou
th, to 4
0 m
just b
efore its end
. Man
groves an
d seagrasses are fou
nd
along b
oth sid
es of the b
ay. On
e sm
all village is located w
ithin
the b
ay.Loky Bay is a large d
eep b
ay, over 10 km
in len
gth
with
extensive frin
ging reefs an
d islan
ds at its m
outh
, and
exten
sive man
grove fringe an
d seagrass m
eadow
s at various
locations. Th
e m
ain river fl ow
ing in
to the b
ay drain
s a large catch
men
t and
man
y add
itional sm
all rivers add
up
to a h
ighly sed
imen
t-infl u
enced
area. Th
ere are several villages in
the b
ay and
temp
orary fi shin
g camp
s.N
osy Ankao is part of a set of islan
ds an
d p
atch reefs on
a ban
k (Leven
Ban
k), high
ly exposed
to ocean w
aves and
with
low
infl u
ence from
rivers. Th
e island
s and
reefs are leased
to an algae aq
uacu
lture com
pan
y that h
as several hu
nd
red
workers livin
g on th
e island
s. Lim
ited fi sh
ing occu
rs and
extern
al fi shers are d
iscouraged
from accessin
g the reef areas.
Andravina Bay is a small sh
allow circu
lar bay w
ith tw
o sm
all river emp
tying in
to it. It show
s very high
infl u
ence of
sedim
ents, w
ith coral com
mu
nities on
limited
hard
substrate
aroun
d th
e mou
th an
d a cen
tral islet. Th
e extensive san
dy
bottom
likely has seagrass b
eds w
here th
e dep
th an
d w
ater clarity is ap
prop
riate. A sm
all village and
fi shin
g camp
(s) are located
in th
e bay.
Vohemar is a large b
ay almost com
pletely taken
up
by an
exten
sive shallow
and
intertid
al reef ban
k, a well d
eveloped
exp
osed fore reef w
ith sp
ur an
d groove d
evelopm
ent, an
d
relatively limited
lagoon an
d ch
ann
el reefs near th
e town
in
the sou
th. A
large river drain
s into th
e bay. C
lear hu
man
im
pacts are visib
le in th
e bay an
d sh
eltered reefs, from
eutro-
ph
ication an
d fi sh
ing.
PRINCIPA
L FIND
ING
S AN
D RECO
MM
END
ATION
S
Reef areasO
f the fi ve location
s, three are h
eavily sedim
ent-aff ected
bay system
s: Am
bod
ivahib
e Bay, L
oky Bay an
d A
nd
ravina.
Of th
ese, the fi rst tw
o are large enou
gh to h
ave areas with
in
them
that are less sed
imen
t-aff ected an
d allow
for comp
lex reef stru
cture (i.e. A
02/A
04 in
Am
bod
ivahib
e, A06/A
11A
Report at a Glance
12Rapid Assessm
ent Program
in L
oky Bay) w
hile th
e small size of A
nd
ravina resu
lted
in a very h
eavily sedim
ent-aff ected
system w
ith low
coral ab
un
dan
ce and
small corals. Th
e sou
thern
end
of our su
rvey, V
ohem
ar, has a very exp
osed fore-reef an
d h
eavily sedim
ent-
and
hu
man
-infl u
enced
back-reef. F
inally, N
osy An
kao is fou
nd
on a seaw
ard b
ank b
athed
by cleaner w
ater and
less in
fl uen
ced by sed
imen
tation, w
hich
is also the site for an
in
tensive seaw
eed farm
ing op
eration.
All com
pon
ents of th
e RA
P fou
nd
sedim
ent in
fl uen
ce and
fi sh
ing to b
e overwh
elmin
g factors aff ecting th
e reefs. Th
us,
wh
ile conservation
tactics may vary am
ong th
e locations, w
e recom
men
d fi rst, lim
iting fi sh
ing p
ressure in
priority sites to
main
tain a stron
g comm
un
ity of grazing fi sh
, and
second
, m
anagin
g up
stream w
atershed
s to min
imize sed
imen
t d
elivery to the reefs. Th
ese strategic goals w
ill both
main
tain
existing reef stru
ctures, an
d m
aintain
and
/or imp
rove coral recru
itmen
t, a key process for su
stainin
g a reef’s ability to
recover from d
isturb
ance.
Corals281 h
ard corals w
ere iden
tifi ed d
urin
g the su
rveys, wh
ich
usin
g Mich
aelis-Men
ten eq
uation
s to pred
ict a total spe-
cies nu
mber resu
lted in
an estim
ate of 313 sp
ecies. Th
is is sligh
tly lower th
an th
e high
est diversity sites in
North
west
Mad
agascar inclu
din
g Nosy B
e and
Nosy M
itsio (323, b
y M
cKen
na et al. 2
005, V
eron an
d T
urak 2
005), b
ut h
igher
than
the N
osy Hara area ju
st W of D
iego on th
e North
west
coast (217, O
bu
ra 2009).
At a region
al scale, North
east Mad
agascar group
s with
the
high
est diversity sites in
North
west M
adagascar, N
Mozam
-biq
ue an
d S T
anzan
ia, lend
ing su
pp
ort to the existen
ce of a h
igh d
iversity center for th
e WIO
in th
is region.
Key sp
ecies of interest for N
ortheast M
adagascar in
clud
e th
e followin
g:
–
the region
al end
emic m
onosp
ecifi c genera –
Craterastrea
laevis, An
omastrea irregu
laris, Horastrea in
dica and
Gyros-
milia in
terrupta
–
Signifi can
t range exten
sions of In
don
esian sp
ecies are rep
orted in
this su
rvey, of An
acropora pillai and
T
urbin
aria irregularis (p
reviously record
ed in
North
west
Mad
agascar, Veron
& T
urak 2
005).
–
A recen
tly describ
ed rare sp
ecies, Psam
mocora albopicta
was fou
nd
here, as w
ell as a poten
tially un
describ
ed sp
e-cies of E
chinopora.
Am
bod
ivahib
e and
Loky B
ay have th
e high
est level of coral d
iversity of the location
s surveyed
du
e to their d
epth
an
d com
plexity, so h
ave the h
ighest valu
e for hab
itat and
sp
ecies conservation
. How
ever ind
ividu
al sites in all 5
loca-tion
s were im
portan
t for some of th
e above key sp
ecies, su
ggesting b
road-b
ased con
servation across th
e entire region
, w
ith focal p
rotection sites an
d strategies is n
ecessary.
Th
e two sites in
An
dravin
a had
the low
est species d
iver-sity of all th
e sites samp
led, sh
owin
g high
infl u
ence of
sedim
entation
. How
ever, they h
ad th
e most u
nu
sual sp
ecies assem
blage of corals, in
clud
ing th
e only sites w
ith A
nom
as-trea irregu
laris, reasonab
ly comm
on C
raterastrea laevis and
com
mon
Horastrea in
dica.
FishTh
e d
iversity of reef fi shes w
as relatively high
bu
t abu
nd
ance
was low
– a total of 2
71 sp
ecies (from ju
st 19 fam
ilies) were
seen on
this N
ortheast coastlin
e, 74%
of the 3
67 sp
ecies rep
orted for M
adagascar sin
ce 1891 (A
llen 2
005). A
total cou
nt of 2
96 sp
ecies was record
ed.
Th
e deep
bays of L
oky Bay an
d A
mbod
ivahib
e Bay h
ad
the h
ighest fi sh
diversity, likely d
ue to h
abitat d
iversity, togeth
er with
sites at Nosy A
nkao w
here reefs h
ad m
ore com
plex stru
cture an
d relatively h
igh coral cover an
d w
ater clarity. M
ultip
le-zone con
servation areas in
these b
ays off er a good
opp
ortun
ity for maxim
izing con
servation of fi sh
sp
ecies.Th
e ab
sence of h
ighly vu
lnerab
le species (sh
ark species,
bu
mp
head
parrots –
Bolbom
etapon m
uricatu
m) as w
ell as low
den
sities of comm
on reef fi sh
eries species (sn
app
ers an
d grou
pers) an
d h
erbivorou
s fi shes, w
hich
are imp
ortant
in m
aintain
ing resilien
t reefs (scrapers –
parrot fi sh
and
grazers –
surgeon
fi sh), w
as noticeab
le. Alth
ough
this m
ay in
part refl ect th
e low relief reefs of th
e region, th
e low ab
un
-d
ance of target fi sh
ery species is in
dicative of fi sh
ing im
pacts.
Th
e presen
ce of small villages, th
e regional cen
ters at V
ohem
ar and
Diego Su
arez, and
activity by seasonal m
igrant
fi shers,are all likely to im
pose m
oderate to h
igh levels of
pressu
re even on
seemin
gly remote location
s. Targeted
spe-
cies/resource m
anagem
ent p
lans w
ith local com
mu
nities are
recomm
end
ed to facilitate fi sh
recovery. On
a positive n
ote, th
e Nap
oleon w
rasse (Cheilin
us u
ndu
latus) w
as seen at several
sites, thou
gh n
one w
ere large termin
al ph
ase males.
Th
e presen
ce of a comm
ercial algae farm at N
osy An
kao, w
hich
discou
rages fi shin
g in th
e area, had
resulted
in a
noticeab
ly high
er species d
iversity and
high
er abu
nd
ance
of fi shery target sp
ecies (omn
ivores) than
seen in
the oth
er location
s. Th
ough
partly attrib
uted
to natu
ral diff eren
ces in
the reefs of N
osy An
kao (low sed
imen
t, high
relief), this
examp
le provid
es an altern
ative man
agemen
t option
e.g. w
ith villages in
the oth
er locations su
rveyed, th
at can b
e con
sidered
for conservation
app
lication.
Algae/seagrasses
Nin
ety one sp
ecies of algae and
ten sp
ecies of seagrass were
foun
d alon
g the n
ortheastern
Mad
agascar. Algal d
iversity w
as dom
inated
by red
s, followed
by green
and
brow
ns
(44, 3
2 an
d 11
species, resp
ectively). Th
e seagrass spe-
cies were typ
ical of East A
frica, nam
ely Th alassoden
dron
ciliatum
, Th alassia hem
prichii, Syringodiu
m isoetifoliu
m,
Cym
odocea rotun
data and
C. serru
lata, Halodu
le un
inervis
and
H. w
rightii, Halophila ovalis an
d H
. stipulacea, Z
ostera
Report at a Glance
13 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
capensis. In
deep
er water (i.e. on
the ou
ter edge of coral
reefs) in ligh
t limited
cond
itions, on
ly Halophila sp
ecies were
foun
d.
Du
e to the ab
sence of coastal d
evelopm
ent, lim
ited b
oat-in
g operation
s and
other an
throp
ogenic stressors, seagrass
bed
s were m
ostly in good
cond
itions alon
g this coastlin
e. Th
e on
ly exception
s were fou
nd
in p
roximity of sew
age out-
falls or larger hu
man
settlemen
ts, particu
larly Voh
emar. In
th
ese areas, the h
igh organ
ic matter con
tent in
the sed
imen
t an
d w
ater turb
idity in
creases epip
hyte grow
th an
d grow
th of
macroalgal form
s. A
lthou
gh h
ealthy, seagrass ecosystem
s may b
e aff ected b
y ch
angin
g climatic con
dition
s in th
e WIO
, throu
gh in
crease in
sea surface tem
peratu
re (expected
to rise up
to 0.6
°C) an
d
sea level rise (pred
icted u
p to 5
0cm
by 2100) an
d ch
anges in
storm
s/cyclone p
atterns, freq
uen
cy and
inten
sity.
Echinoderms
Sixty eight sp
ecies of echin
oderm
s belon
ging to fi ve classes
were record
ed: 3
species of crin
oid, 1
8 h
olothu
rians,
27 op
hiu
roids, 1
0 ech
inoid
s and
10 asteroid
s, wh
ich w
ith an
ad
dition
al 2 sp
ecies previou
sly recorded
from th
e area gives a total rich
ness of 7
0 sp
ecies for the su
rvey area. Th
e h
ighest d
iversity of echin
derm
s was fou
nd
in d
eep
bays (e.g. L
oky Bay) w
here h
abitat d
iversity was h
ighest,
bu
t also in im
pacts sites (e.g. V
ohem
ar) wh
ere opp
ortun
is-tic sp
ecies typical of d
egraded
cond
itions w
ere dom
inan
t.A
s echin
oderm
species are good
ecological ind
icators,their
pattern
s of abu
nd
ance are good
ind
icators of the con
dition
of th
e reefs. Th
e pred
atory seastar Acan
thaster planci w
as not
observed
du
ring th
e survey.
Th
e pattern
s of echin
oderm
species d
istribu
tions stron
gly su
pp
ort other fi n
din
gs of the R
AP
and
the n
eed for b
iodi-
versity/ resource con
servation in
the m
ost diverse sites an
d
remed
ial action in
the m
ost degrad
ed sites.
Coral exosymbionts
On
prelim
inary an
alysis, 67 exosym
bion
t taxa from ap
proxi-
mately 4
8 stru
ctural sp
ecies were sam
pled
. A n
um
ber of
the exosym
bion
ts show
ed ch
aracteristics not kn
own
from
Table 1. Study sites visited during the RAP. Site codes correspond to the main m
ap on page 25.
LocationCode
Site name
Latitute (S)Longitude (E)
Depth (m)
Ambodihavibe
A01
Am
bod
ivahib
e Inn
er12°2
2.2
75
49°2
6.4
25
10
A02
Am
bod
ivahib
e S12°2
1.5
249°2
7.6
89
10
A03
"12°2
2.3
449°2
6.9
72
9
A04
Am
bod
ivahib
e S, chan
nel
12°2
1.0
749°2
7.8
93
15
A05
Am
bod
ivahib
e N12°2
0.3
249°2
6.1
66
10
LokyA
06
Loky S
12°4
3.8
249°4
1.7
92
10
A07
"12°4
4.9
43
49°4
0.7
43
10
A08
Loky N
orth W
est12°4
3.4
52
49°4
0.0
49
A09
"12°4
2.1
19
49°3
9.9
21
10
A10
Loky N
orth12°3
9.4
48
49°3
6.6
99
9
A11A
Loky A
cropora gard
ens
12°3
9.7
74
49°3
6.4
86
2
A11B
Loky in
ner reef
12°4
3.1
04
49°3
9.1
09
0.5
A12
Loky W
est12°4
4.3
949°3
9.6
62
4
A19
Loky In
ner
12°4
5.1
88
49°4
0.2
78
12
AnkaoA
20
An
kao North
12°4
7.4
84
49°4
8.7
31
6
A21
An
kao South
12°4
9.6
48
49°4
8.7
13
6
A22
An
kao North
-East
12°4
6.0
04
49°4
9.3
51
10
A23
An
kao North
12°4
6.9
11
49°4
8.6
58
5
A24
An
kao Bay
12°4
8.1
74
49°4
7.5
36
5
AndravinaA
17
An
dravin
a Bay S p
t12°5
6.4
63
49°5
2.0
96
6
A18
An
dravin
a rock12°5
5.9
55
49°5
1.2
46
6
Vohemar
A13
Voh
emar N
orth13°1
9.5
85
50°0
.807
8
A14
Voh
emar In
ner
13°2
0.8
81
50°0
.279
6
A15
Voh
emar N
orth13°2
0.0
89
50°0
.899
11
A16
Voh
emar Sou
th13°2
1.0
83
50°1
.177
11
Report at a Glance
14Rapid Assessm
ent Program
existing sp
ecies, giving a p
ossibility of several n
ew sp
ecies. E
ven b
ased on
such
prelim
inary fi n
din
gs, the su
ggestion for
so far un
-measu
red region
al diversity is stron
g, emp
hasi-
zing th
e need
for furth
er samp
ling in
this area an
d eff ective
conservation
actions to p
revent losses b
efore species can
be
docu
men
ted an
d d
escribed
.
Reef health and coral bleachingO
verall the region
show
ed h
igh coral cover (m
ean of 4
8%
), an
d coral p
opu
lations th
at span
ned
the exp
ected size ran
ge for each
genu
s, inclu
din
g large, matu
re colonies. Th
e
north
ern th
ree locations (A
mbod
ivahib
e Bay, N
osy An
kao, L
oky Bay) sh
owed
greater coral cover, larger corals and
fewer
bleach
ed colon
ies than
the sou
thern
two sites (A
nd
ravina,
and
Voh
emar).
From
Feb
ruary to A
pril 2
010 a w
arm p
ool of water
aff ected reefs th
rough
out th
e western
Ind
ian O
cean, cau
sing
bleach
ing in
several locations. In
the stu
dy area h
owever,
coral bleach
ing w
as relatively low, sh
owin
g a mean
~5%
of colon
ies aff ected. Th
is fi n
din
g, together w
ith th
e low im
pact
detected
from th
e 1998 m
ass bleach
ing even
t, suggests th
at th
e corals in th
is region h
ave largely resisted th
e negative
eff ects of heatin
g events, an
d su
pp
orts the h
ypoth
esis that
these reefs are th
ermally resistan
t.Th
e recru
itmen
t of corals in th
e areas surveyed
was p
resent
bu
t generally low
. Th
is may b
e du
e to a nu
mber of factors,
inclu
din
g: (1) few
sources of coral larvae: m
ost reefs in th
is region
have a n
arrow an
d sh
allow reef p
rofi le, limitin
g the
total area of reef comm
un
ity and
poten
tial source colon
ies for rep
rodu
ction, an
d th
e few u
pstream
sources of larvae in
th
e central In
dian
Ocean
are qu
ite distan
t; (2) low
retention
of d
ispersin
g larvae: strong cu
rrents th
at accelerate north
-w
estward
aroun
d th
e tip of M
adagascar m
ay deliver d
isper-
sing larvae to th
e East A
frican m
ainlan
d in
stead of retain
ing
them
locally; and
(3) little good
settlemen
t substrate: m
ost reefs sh
owed
a relatively low ab
un
dan
ce of bare, h
ard su
b-
strate free of sedim
ent, tu
rf, soft coral or spon
ges. We
recomm
end
conservation
actions to en
han
ce coral recruit-
men
t, a key factor for ecological resilience, th
rough
water-
shed
man
agemen
t to limit/red
uce sed
imen
tation
GEN
ERAL RECO
MM
END
ATION
S
Of th
e fi ve locations, th
ree are heavily sed
imen
t-aff ected
bay system
s: Am
bod
ivahib
e Bay, L
oky Bay an
d A
nd
ravina.
Of th
ese, the fi rst tw
o are large enou
gh to h
ave areas with
in
them
that are less sed
imen
t-aff ected an
d allow
for comp
lex reef stru
cture (i.e. A
02/A
04 in
Am
bod
ivahib
e, A06/A
11A
in
Loky B
ay) wh
ile the sm
all size of An
dravin
a resulted
in
a very heavily sed
imen
t-aff ected system
with
low coral
abu
nd
ance an
d sm
all corals. Th
e south
ern en
d of ou
r survey,
Voh
emar, h
as a very exposed
fore-reef; the lagoon
and
back-
reef are very heavily sed
imen
t- and
hu
man
-infl u
enced
to the
extent th
at coral comm
un
ity develop
men
t is low. F
inally,
Nosy A
nkao is on
a seaward
ban
k bath
ed by clean
er oceanic
water an
d less in
fl uen
ced b
y sedim
entation
, and
is also the
site for an in
tensive seaw
eed farm
ing op
eration.
Th
e fi nd
ings of th
is RA
P su
pp
ort the recom
men
dation
s p
roposed
in p
revious su
rveys (see box b
elow) an
d p
ropose
add
itional action
s for the m
anagem
ent an
d con
servation of
marin
e resources in
North
east Mad
agascar and
the W
IO
Region
:.
1)
ES
TA
BL
ISH
new
Marin
e Protected
Areas. A
nu
mber
of opp
ortun
ities and
priority areas can
be id
entifi ed
on
this coastlin
e for protection
, throu
gh p
artnersh
ips w
ith
diff eren
t institu
tions an
d com
mu
nities to ach
ieve the
most eff ective p
rotection of b
iodiversity an
d n
atural
resources, an
d su
stenan
ce of local livelihood
s.A
mbodivahibe –
high
priority ‘clim
ate park’ w
ith
existing activities for con
servation p
lann
ing, an
d n
ested
with
in a b
roader con
servation in
itiative inclu
din
g Ivo-von
a, Ram
ena an
d D
iego Bay to th
e north
. L
oky Bay –
high
est diversity of corals an
d fi sh
du
e to com
plex h
abitats an
d large area. R
eplicate u
pw
elling/
climate p
rotection site to A
mbod
ivahib
eA
nkao –
un
usu
al island
system w
ith clear w
ater, and
w
ith th
e best fi sh
pop
ulation
s du
e to de facto p
rotection
by con
cession ow
ner of th
e island
s.A
ndravin
a – low
diversity b
ut h
ighly u
niq
ue region
al en
dem
ic coral pop
ulation
s.V
ohemar –
high
diversity an
d h
ighly d
eveloped
fore reef system
, bu
t with
heavy u
se and
imp
act of the
bay an
d reefs from
urb
an an
d p
ort develop
men
t and
sed
imen
tation.
2)
EN
HA
NC
E fi sh
eries man
agemen
t. Th
e imp
act of local an
d m
igrant fi sh
ing, for food
and
comm
ercial markets,
were clear even
in th
e most rem
ote and
least accessible
locations on
the su
rvey. It is essential to m
aintain
fi sh
pop
ulation
s to ensu
re their ecosystem
services are rea-lized
, in p
articular a stron
g comm
un
ity of grazing fi sh
es. M
aintain
ing or b
oosting th
e pop
ulation
of grazers w
ill likely aid th
e main
tenan
ce of existing ad
ult corals
(i.e. sources of coral larvae), an
d h
as strong eff ects on
m
aintain
ing th
e availability of b
are, grazed settlem
ent
substrate. F
ish ab
un
dan
ce was gen
erally low relative to
other sites in
the region
, with
some key sp
ecies absen
t. Sp
ecifi c fi shery m
anagem
ent p
lans w
ill be req
uired
to en
courage fi sh
pop
ulation
recovery. Th
ough
fi sh sp
ecies d
iversity was relatively h
igh overall, certain
families
and
sites had
notab
ly low d
iversity. Wh
ile mu
ch of th
e region
app
ears sparsely p
opu
lated, w
e repeated
ly noticed
tem
porary fi sh
ing cam
ps, an
d often
these w
ere equ
ipp
ed
with
eff ective gear such
as out-b
oard m
otors. Fu
rther
socio-econom
ic research sh
ould
look specifi cally at th
e eff ects of m
igrant fi sh
ers.
3)
PR
OM
OT
E w
atershed
man
agemen
t to redu
ce sed
imen
t imp
acts on m
arine system
s. Urgen
t steps
are need
ed to con
trol sedim
ent d
elivery into th
e reef
Report at a Glance
15 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
system. A
mod
erate level of sedim
ent is n
ot necessar-
ily detrim
ental for reefs, an
d th
is region faces n
aturally
high
levels of sedim
entation
, eviden
ced by m
any sed
i-m
ent-ad
apted
coral comm
un
ities, with
high
abu
nd
ance
of more sed
imen
t-tolerant sp
ecies. In fact, m
any of th
e region
al end
emics, e.g. H
orastrea and
Gyrosm
illia, are w
ell adap
ted to sed
imen
t heavy ecosystem
s. How
ever, in
man
y sites reef develop
men
t was ap
paren
tly limited
by
sedim
entation
, either th
rough
direct b
urial or th
rough
ligh
t limitation
in th
e turb
id w
ater. Given
the h
istory of rap
id d
eforestation in
Mad
agascar, and
the con
ti-n
ued
heavy relian
ce of the grow
ing coastal p
opu
lation
on m
angrove stan
ds for b
uild
ing m
aterial and
charcoal
cooking fi res, sed
imen
t delivery h
as likely increased
recen
tly and
, with
out in
tervention
, will con
tinu
e to in
crease. Some of th
e watersh
eds aff ectin
g the reefs,
e.g. Loky B
ay, have existin
g forest conservation
initita-
tives in w
hich
CI is a p
artner, an
d th
ese can b
e leve-raged
to broad
en th
e focus of w
ork to a ridge-to-reefs
framew
ork.
4)
IMP
LE
ME
NT
species con
servation p
lans. F
lagship
sp
ecies for protection
(e.g. sharks, sea tu
rtles, Nap
oleon
wrasse, B
um
ph
ead p
arrotfi sh an
d large grou
pers) w
ill all b
enefi t from
species con
servation p
lans if th
ey can
be estab
lished
in th
is area, or nation
ally in M
adagas-
car. Th
e poten
tial for species p
lans for u
nu
sual sp
ecies m
ay also be p
owerfu
l tools for broad
er goals, such
as for evolu
tionarily d
istinct corals en
dem
ic to the
western
Ind
ian O
cean (C
raterastrea laevis, An
omastrea
irregularis, H
orastrea indica an
d G
yrosmilia in
terrupta).
Th
ese can serve as fl agsh
ip sp
ecies for imp
lemen
tation
of other recom
men
dation
s (e.g. MP
As an
d w
atershed
m
anagem
ent).
5)
INS
TIG
AT
E eff ective p
artnersh
ips for con
servation.
With
the h
ighly d
ispersed
and
marin
e-resource-d
epen
-d
ent local com
mu
nities an
d p
opu
lation, p
artnersh
ips
will b
e the corn
erstone for eff ective con
servation an
d
marin
e man
agemen
t. Th
is has b
een clearly sh
own
by
experien
ce since th
e 2n
d RA
P an
d th
e develop
men
t of con
servation activities in
the A
mbod
ivahib
e-Ivovna-
Ram
ena-D
iego Bay system
to the n
orth. Th
ree typ
es of p
artnersh
ips m
ay be p
ossible, d
epen
din
g on th
e location
and
the b
alance of local stakeh
olders:
Local govern
men
t: In ou
r meetin
gs with
the local
governm
ent in
Voh
emar, th
ey expressed
great interest in
con
servation eff orts for th
eir area, bu
t were q
uite can
did
ab
out th
e curren
t lack of such
eff orts. Coastal zon
e m
anagem
ent, w
atershed
man
agemen
t and
pollu
tion/
mu
nicip
al man
agemen
t are critical areas for interven
-tion
in all location
s, bu
t particu
larly Voh
emar an
d L
oky B
ay.Local com
mun
ities: man
y of the villages in
the region
are q
uite rem
ote, with
poor access b
y land
and
limited
in
frastructu
re for travel by sea. Com
mu
nity-b
ased
initiatives for con
servation, p
aired w
ith in
come gen
era-tion
and
imp
roving stan
dard
s of living w
ill be n
ecessary in
these location
s. Integrated
projects in
corporatin
g h
ealth an
d ed
ucation
, two areas clearly lackin
g in som
e of th
e remote villages, m
ay be n
ecessary.P
rivate Partn
erships: private in
vestors and
ind
ustries
(e.e. seaweed
farmin
g) have b
een active in
the region
for over a d
ecade. N
osy An
kao’s reefs, for examp
le, rem
ain th
e least aff ected reefs in
our su
rvey by b
oth
sedim
entation
and
fi shin
g as seaweed
farmers are
actively discou
raging fi sh
ing on
the reefs. Th
e p
rivate sector m
ay also provid
e local fi shers w
ith an
alternative
livelihood
throu
gh algal farm
ing, th
ereby red
ucin
g fi sh
ing p
ressure. E
ff ective partn
ership
with
private
partn
ers can b
e key in ach
ieving eff ective con
servation
targets.
6)
CO
ND
UC
T fu
rther b
iological assessmen
ts. Th
is su
rvey represen
ts the th
ird of six su
rveys recomm
end
ed
followin
g the fi rst R
AP
(McK
enn
a et al. 2005). P
riority sites for ad
dition
al RA
Ps can
be id
entifi ed
. Based
on
the p
resence of th
e high
ly succesfu
al Masoala M
arine
Park ab
out 2
00 km
south
of Voh
emar, several M
PA
in
itiatives on th
e North
west coast from
Nosy H
ara to Sah
amalaza, an
d exten
sive work on
the sou
thw
est coast (th
e region ecom
passin
g An
davad
oaka to Tu
lear) a clear gap
is the coastlin
e south
of Saham
alaza dow
n
the N
orthw
est coast, aroun
d th
e ‘hu
mp’ of th
e island
in
Conservation Recomm
endations, RAP 31 (2005):
1)
Pu
rsue a vigorou
s program
of bio
logical
assessmen
t.
2)
Review
and
con
solid
ate previo
us k
no
wled
ge of M
adagascar’s m
arine b
iota.
3)
Collection
of add
ition
al data an
d lo
ng-term
m
on
itorin
g program
s essential for m
arine con
ser-vation
plan
nin
g and
imp
lemen
tation.
4)
Estab
lish a n
etwo
rk o
f marin
e pro
tected areas.
5)
En
act more eff ective law
s to regulate fi sh
ing
activities.
6)
Prom
ote and
develop
con
servation
-orien
ted
marin
e tou
rism for th
e ben
efi t of local com
mu
nities.
7)
Provid
e dive train
ing for staff of local u
niversities
and
conservation
organization
s.
8)
Set up
ou
treach an
d p
ub
lic awaren
ess program
s on
marin
e resources to all local stakeh
olders.
9)
Prom
ote com
mu
nity p
articipatio
n in
conserva-
tion p
lann
ing an
d m
anagem
ent.
10) Stren
gthen
species co
nservatio
n p
rogram
s for rare an
d en
dan
gered m
arine w
ildlife.
Report at a Glance
16Rapid Assessm
ent Program
the M
ozambiq
ue ch
ann
el, to the B
arren Islan
ds. Th
is
represen
ts a coastline of close to 1
,000 km
, and
the least
know
n of reef en
vironm
ents in
the cou
ntry.
7)
DE
SIG
N gu
idelin
es and
best p
ractices for coastal and
foresh
ore develop
men
t plan
nin
g. As coastal d
eve-lop
men
t and
pop
ulation
growth
increases alon
g this
coastline, coral reefs, m
angroves an
d seagrasses w
ill face grow
ing p
ressures su
ch as m
echan
ical distu
rban
ce for in
frastructu
re develop
men
t (hotels, m
arinas, etc)
and
from b
oating, in
creasing sed
imen
t run
off du
e to d
eforestation, an
d extraction
for food an
d com
merce.
At p
resent, th
e North
east coast is relatively remote, an
d
is protected
by the very rou
gh seas an
d h
ighly exp
osed
natu
re of the sh
orelines. Strategies to m
anage an
d lim
it th
reats as pop
ulation
increases w
ill be n
ecessary, to rep
lace the n
atural p
rotection as in
centives in
crease to exp
loit the area. A
dd
itionally, “clim
ate-smart” d
evelop-
men
t shou
ld b
e prom
oted, w
hich
takes into accou
nt th
e p
otential for in
creased storm
activity, salt water in
tru-
sion, an
d oth
er climate ch
ange im
pacts. Th
is b
est-prac-
tice develop
men
t is particu
larly timely as M
adagascar’s
nation
al Integrated
Coastal Z
one M
anagem
ent (G
estion
Intégrée d
es Zon
es Côtères –
GIZ
C) p
rocess is gainin
g p
olicy traction at th
e nation
al level.
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, G. R
. 2005. R
eef fi shes of N
orthw
estern M
adagascar.
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cKen
na, S., G
.R. A
llen an
d H
. Ran
drian
asolo (ed
s.). A R
apid
Marin
e Biod
iversity Assessm
ent of th
e C
oral reefs of North
west M
adagascar. R
AP
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lletin of
Biological A
ssessmen
t 31. C
onservation
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ational,
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GA
P, 2001. P
lan d
e Gestion
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arcs Nation
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ada-
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oles, S. and
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Brow
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003. C
oral bleach
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arine
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e imp
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iodiversity an
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aharavo, J., T
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liver and
A. R
abearisoa. In
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apid
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e Biod
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ent of N
ortheast
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servation In
ternation
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arivo. M
adagascar.
Cooke, A
., O. R
atomah
enin
a, E. R
anaivoson
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aza-fi n
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adagascar. In
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pard
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ental evalu
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olum
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egional ch
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Main
a, J. and
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ra. 2008. C
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atial d
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d M
arine E
cosystem V
uln
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ents in
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enu
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odelin
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IS an
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estern
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McC
lanah
an, T
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oham
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limate ch
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spatio-tem
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in seaw
ater temp
erature
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col. Mon
ogr. 77: 5
03–525.
McK
enn
a, S., G.R
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and
rianasolo (ed
s.). 2005. A
Rap
id M
arine B
iodiversity A
ssessmen
t of the
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orthw
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P B
ulletin
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iological Assessm
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1. C
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ational,
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bu
ra, D.O
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oral Reef R
esilience A
ssessmen
t of the
Nosy H
ara Marin
e Protected
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orthw
est Mad
agas-car. W
WF
Mad
agascar.O
bu
ra, D.O
. 2005. R
esilience an
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d b
leachin
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e Western
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oastal and
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Qu
od, J.P., Y. D
ahalan
i, L.B
igot, J. B. N
icet, S. Ah
amad
a an
d J. M
aharavo. 2
002. Statu
s of Coral R
eefs at R
éun
ion, M
ayotte and
Mad
agascar. In: L
ind
en, O
., D.
Souter, D
. Wilh
elmsson
, and
D.O
. Obu
ra (eds.). C
oral R
eef Degrad
ation In
Th
e Ind
ian O
cean. Statu
s reports
2002. C
OR
DIO
/SAR
EC
Marin
e Science P
rogram. P
p. 185-1
89.
Veron
, J. E. N
. and
E. T
urak. 2
005. R
eef corals of north
-w
est Mad
agascar. In: M
cKen
na, S., G
.R. A
llen an
d
H. R
and
rianasolo (ed
s.) A R
apid
Marin
e Biod
iversity A
ssessmen
t of the C
oral reefs of North
west M
adagascar.
RA
P B
ulletin
of Biological A
ssessmen
t 31. C
onservation
In
ternation
al, Wash
ington
DC
, USA
. Pp. 2
3-3
0.
Web
ster, F. J. and
K. M
cMah
on. 2
002. A
n A
ssessmen
t of C
oral Reefs in
North
west M
adagascar. In
: Lin
den
, O.,
D. Sou
ter, D. W
ilhelm
sson, an
d D
.O. O
bu
ra (eds.).
Coral R
eef Degrad
ation In
Th
e Ind
ian O
cean. Statu
s rep
orts 2002. C
OR
DIO
/SAR
EC
Marin
e Science P
ro-gram
. Pp. 1
90-2
00.
West, J. an
d R
. Salm. 2
003. E
nviron
men
tal determ
inan
ts of resistan
ce and
resilience to coral b
leachin
g: imp
lications
for marin
e protected
area man
agemen
t. Con
servation
Biology 1
7: 9
56-9
67.
17A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
SUM
MA
RY
North
east and
North
west M
adagascar h
ave similar levels of coral d
iversity, the h
ighest
recorded
in th
e Western
Ind
ian O
cean. Th
is su
rvey recorded
281 sp
ecies of corals, with
low
er taxonom
ic eff ort than
earlier surveys th
at recorded
323 sp
ecies of corals in N
orthw
est M
adagascar.
At a region
al scale, this coral d
iversity is equ
ivalent to sites in
N M
ozambiq
ue an
d S T
anza-
nia, d
efi nin
g a triangle of m
aximu
m d
iversity in th
e western
Ind
ian O
cean, en
comp
assing th
e n
orthern
Mozam
biq
ue ch
ann
el.Sites in
North
east Mad
agascar host som
e of the largest p
opu
lations of th
e most evolu
-tion
arily distin
ct and
ecologically rare mon
ospecifi c gen
era in th
e Ind
ian O
cean, in
clud
ing
Craterastrea laevis, A
nom
astrea irregularis, H
orastrea indica an
d G
yromsilia in
terrupta. Sign
ifi -can
t range exten
sions of In
don
esian sp
ecies are reported
in th
is survey, of A
nacropora pillai an
d
Turbin
aria irregularis (p
reviously record
ed in
North
west M
adagascar, V
eron &
Tu
rak 2005).
A recen
tly describ
ed rare sp
ecies, Psam
mocora albopicta w
as foun
d h
ere, as well as a p
otentially
un
describ
ed sp
ecies of Echin
opora.T
wo of th
e fi ve locations, A
mbod
ivahib
e and
Loky B
ay, have th
e high
est level of diversity
amon
g the location
s surveyed
du
e to their d
epth
and
structu
ral comp
lexity, so have th
e high
est valu
e for hab
itat and
species con
servation. H
owever in
divid
ual sites in
all 5 location
s were
imp
ortant for som
e of the ab
ove key species, su
ggesting b
road-b
ased con
servation across th
e en
tire region, w
here focal p
rotection sites an
d strategies is n
ecessary.
INTRO
DU
CTION
Scleractinian
corals are the arch
itects that b
uild
coral reefs, the m
ost diverse m
arine en
viron-
men
ts on th
e plan
et. Th
eir diversity at a location
is ind
icative of the d
iversity and
robu
stness of
other reef fau
na, an
d corals h
ave been
the focu
s of biod
iversity conservation
and
research for
decad
es. Th
e coral reefs of Mad
agascar have b
een th
e focus of research
for over 50 years, b
ut
hu
man
use of reef an
d lin
ked en
vironm
ents is cau
sing m
assive imp
acts and
leadin
g to biod
i-versity loss. C
orals are one of th
e focal group
s of reef biod
iversity assessmen
ts, and
this w
ork con
tinu
es that of tw
o earlier marin
e RA
Ps in
north
ern M
adagascar (M
cKen
na et al. 2
005,
Mah
aravo and
al., 2011).
Th
e fi rst marin
e RA
P (M
cKen
na et al. 2
005) estab
lished
that th
e North
west tip
of Mad
agas-car h
ad th
e high
est diversity of h
ard corals of an
y site in th
e Ind
ian O
cean, w
ith 3
23 sp
ecies ob
served on
the su
rvey and
a total of 380 sp
ecies reported
for Mad
agascar overall. 463 reef fi sh
sp
ecies were id
entifi ed
, with
a pred
iction of 5
76 as a total sp
ecies richn
ess for North
west M
ad-
agascar and
a list of 788 sp
ecies for Mad
agascar overall. Th
e main
fi nd
ing from
the se-con
d
marin
e RA
P (M
aharavo an
d al., in
press) w
as that A
mbod
ivahib
e, on th
e North
east tip of
Mad
agascar was ap
paren
tly un
-imp
acted b
y ocean-w
ide m
ass bleach
ing an
d m
ortality of corals in
1998 an
d sh
owed
characteristics of cooler w
ater du
e to up
wellin
g, poten
tially a protection
Chapter 1
Corals of northeast Madagascar
David Obura
Chapter 1
18Rapid Assessm
ent Program
mech
anism
shelterin
g reefs in th
e bay from
high
temp
erature
events cau
sing coral b
leachin
g. On
the b
asis of this featu
re, th
e bay w
as iden
tifi ed as a p
otential ‘clim
ate park’.
Th
e objectives of th
is coral survey of th
e 3rd m
arine R
AP
w
ere to extend
the coral d
iversity surveys from
the p
re-viou
s RA
Ps farth
er dow
n th
e North
east coast, and
to collect d
ata comp
atible w
ith C
OR
DIO
’s coral diversity su
rveys th
at started in
2002, b
uild
ing u
p a d
ataset of 17 location
s to u
nd
ertake a consisten
t regional an
alysis of coral biod
iversity an
d b
iogeograph
ic pattern
s (Obu
ra, in p
rep). Th
ese su
rveys w
ill help
iden
tify the exten
t and
natu
re of the cen
ter of d
iversity for the w
estern In
dian
Ocean
that in
clues n
orthern
M
adagascar as on
e of its three corn
ers.
METH
OD
S
Coral sp
ecies were id
entifi ed
in th
e fi eld, an
d a fu
ll species
list was d
eveloped
based
on fi eld
IDs u
sing d
igital ph
otogra-p
hy as a p
rimary referen
ce and
references p
rovidin
g un
der-
water p
hotograp
hs (see Sh
epp
ard an
d O
bu
ra 2004). D
ue to
other ob
jectives of the exp
edition
a full sp
ecies list was n
ot bu
ilt up
for each site, th
ough
a comp
rehen
sive species list
was attem
pted
at most sites, focu
sing on
searchin
g for new
sp
ecies not seen
on p
rior dives.
Usin
g species record
s from su
ccessive dives an
accum
ula-
tion cu
rve for the su
rvey area is establish
ed th
at asymp
totes tow
ards a total sp
ecies richn
ess for the stu
dy area. R
egio-n
ally, betw
een 1
0 an
d 3
0 d
ives have b
een record
ed in
each
stud
y area, with
30 sam
ples b
eing record
ed in
this su
rvey of N
ortheast M
adagascar (at 2
4 u
niq
ue sites). M
ichaelis-M
en-
ten en
zyme kin
etic equ
ations h
ave been
app
lied to estim
ate sp
ecies diversity in
a nu
mber region
s and
taxonom
ic group
s (K
eating 1
998). Th
e eq
uation
s app
roximate a sam
plin
g cu
rve that satu
rates after a large nu
mber of sam
ples, an
d
calculate a p
redicted
maxim
um
species rich
ness (S
max ) w
ith
infi n
ite samp
les. Th
e mu
ltivariate analysis softw
are PR
IME
R
v 6.0
(Clarke an
d G
orley 2006) in
clud
es mod
ules for estim
a-tion
of accum
ulation
curves, in
clud
ing M
ichaelis-M
enten
coeffi cien
ts, and
this is u
sed h
ere to calculate th
e expected
sp
cies diversity of th
e these sam
ples, to b
e used
in region
al com
parison
s of coral diversity.
RESULTS
Coral species richnessA
total of 281 sp
ecies of reef-bu
ildin
g (skeleton-form
ing)
corals were id
entifi ed
in th
is survey. O
f these, 2
75 sp
ecies w
ere in th
e order scleractin
ia (hard
corals), 6 sp
ecies were
non
-scleractinia (T
able 1
.1), an
d a total of 6
1 gen
era and
17 fam
ilies were record
ed. Th
is total com
pares w
ith th
e follow
ing:
Usin
g exactly the sam
e survey tech
niq
ue 2
17 sp
ecies were
recorded
at the N
osy Hara in
the N
orthw
est (just w
est of the
north
ern tip
of Mad
agascar, 16 sites). Th
ese n
um
bers in
di-
cate that th
e North
east coast surveyed
is more d
iverse than
th
e up
per p
art of the N
orthw
est coast, at Nosy H
ara. Th
e 1
st marin
e RA
P in
clud
ed m
ore comp
rehen
sive taxo-n
omic exp
ertise and
meth
ods for sam
plin
g and
iden
tifying
hard
corals (Veron
& T
urak 2
005), an
d covered
a larger area from
Nosy B
e to Nosy H
ara in 2
002. It record
ed 3
23 sp
ecies of corals (exclu
din
g non
-scleractinia, so com
parab
le with
275 scleractin
ian sp
ecies recorded
here) w
ere recorded
. Th
e h
ighest d
iversity of corals du
ring th
e 1st m
arine R
AP
was in
th
e Nosy M
itsio archip
elago, with
somew
hat low
er diversity
at Nosy H
ara.U
sing M
ichaelis-M
enten
pred
iction cu
rves (fi g. 1.2
), a p
redicted
species n
um
ber of 3
13 w
as foun
d for N
ortheast
Mad
agascar, suggestin
g an ad
dition
al 32 sp
ecies than
those
recorded
in th
e surveys w
ould
be fou
nd
in ad
dition
al sur-
veys. Samp
ling of exp
osed fore reef sites, w
hich
was h
ighly
restricted b
y weath
er on th
is trip, wou
ld likely h
ave revealed
add
itional sp
ecies and
closer concord
ance b
etween
the actu
al an
d p
redicted
richn
ess.Sp
ecies richn
ess is estaimted
by Sm
ax from M
ichaelis-
Men
ten eq
uation
s in P
RIM
ER
v6.0
. Sites in T
anzan
ia, M
ozambiq
ue an
d M
adagascar h
ave the h
ighest d
iversity, w
hile sites in
the islan
ds (Seych
elles) and
in n
orthern
Ken
ya an
d D
jibou
ti have th
e lowest d
iversity. Source: O
bu
ra, data
un
der p
reparation
.Th
e p
redicted
nu
mber of sp
ecies iden
tifi ed in
this stu
dy
can b
e comp
ared to oth
er sites in th
e region w
here th
e same
meth
od h
as been
app
lied (fi g. 1
.2). N
ortheast M
adagascar
ranks h
ighest in
this d
ataset (313 sp
ecies), followed
by M
afi a islan
d (3
07 sp
ecies, central T
anzan
ia). All of th
e locations
with
species rich
ness >
250 exten
d in
a triangle from
Ch
agos arch
ipelago in
the east to n
orthern
Mozam
biq
ue to cen
tral T
anzan
ia in th
e north
(the K
enya d
ata poin
t is based
on
sporad
ic surveys sp
read over 5
years wh
ereas all the oth
er sites w
ere samp
led w
ithin
a few w
eeks, so may n
ot be com
-p
arable). Th
ese resu
lts pin
poin
t the corn
ers of a hyp
othesized
Figure 1.1. Number of coral species identifi ed in Northeast M
adagascar in M
arch-April 2010, showing the accumulation of species with each
additional dive sample, and a logarithm
ic regression curve passing through the points (r ²=
0.98).
Corals of northeast Madagascar
19 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
triangle of h
igh d
iversity in th
e western
Ind
ian O
cean, an
alo-gou
s to the C
oral Trian
gle in sou
theast A
sia, wh
ere diversity-
generatin
g and
main
tainin
g processes are stron
gest.
Location and site comparisons
Am
ong th
e locations su
rveyed, A
mbod
ivahib
e and
Loky
Bay scored
the h
ighest n
um
ber of sp
ecies, with
212 each
(11
and
7 sam
ples, resp
ectively). Species rich
ness at th
e other
locations w
as consid
erably low
er, with
168 at A
nkao (6
sur-
veys), 155 sp
ecies at Voh
emar (4
surveys) an
d 111
species at
An
dravin
a (2 su
rveys). Interestin
gly, the n
um
ber of sp
ecies at L
oky Bay sh
owed
a strong fl atten
ing at 7
+ su
rveys wh
ile spe-
cies nu
mber at A
mbod
ivahib
e contin
ued
to increase, sh
own
by th
e logarithm
ic curve for A
mbod
ivahib
e rising h
igher
than
that for L
oky Bay (fi g. 1
.3). A
possib
le reason for
high
er diversity at A
mbod
ivahib
e over Loky B
ay is the low
er level of sed
imen
t infl u
ence, as th
e former d
oes not h
ave a m
ajor river system feed
ing in
to it, wh
ile the latter d
oes. Even
th
e larger area and
more d
iverse hab
itats of Loky B
ay don’t
app
ear to comp
ensate for th
is eff ect. How
ever the ou
ter reef at L
oky Bay cou
ld n
ot be su
rveyed, an
d clear-w
ater species
foun
d in
Am
bod
ivahib
e migh
t occur th
ere bu
t were n
ot sam
pled
. Also in
terestingly, th
e curves for A
nkao, V
ohem
ar an
d A
nd
ravina are n
early iden
tical. Th
ese locations h
ad
lower h
abitat d
iversity than
the b
ay systems, in
particu
larly A
nd
ravina, w
hich
is a very small low
-comp
lexity bay w
ith
high
sedim
ent in
fl uen
ce.A
nkao sh
owed
characteristics of b
eing a m
ore typical ou
ter reef system
for East A
frica (extensive p
atch reefs on
a clear-w
ater sand
and
rock platform
from 3
-8 m
deep
), thou
gh
samp
ling w
as only p
ossible on
patch
and
fringin
g reefs, w
ithou
t surveys on
the ou
ter reefs. Voh
emar is a large reef
system b
ut w
ith low
diversity of h
abitats an
d lim
ited area
of good h
abitat for corals. Th
e ou
ter reef is very high
ly
develop
ed w
ith h
igh cover of corals on
comp
lex bom
mies
and
pillars d
own
to 15 m
, bu
t is un
iform an
d h
omogen
eous.
Th
e chan
nel ed
ge has h
igh coral cover b
ut relatively low
d
iversity du
e to high
sedim
ent in
fl uen
ce, bu
t most of th
e lagoon
and
reef fl at are either too sed
imen
t-infl u
enced
or sh
allow an
d aerially exp
osed, resp
ectively, for corals.O
n a site b
y site basis (fi g. 1
.4), L
oky Bay h
ad th
e site w
ith th
e high
est diversity, site 6
; a reef with
high
topograp
hic
comp
lexity of spu
rs and
bom
mies on
a sand
y base at 8
m,
rising to w
ithin
1-2
m of th
e surface, th
en p
lun
ging d
own
a steep
wall in
to the d
eep (>
50 m
) center of th
e bay. L
ocated
on th
e insid
e of the sp
ur of lan
d en
closing th
e bay, th
e site is sh
eltered from
ocean w
aves, bu
t suffi cien
tly cleaned
by
wave action
to off set the h
igh sed
imen
t fl owin
g out of th
e bay. In
Am
bod
ivahib
e, high
diversity of corals w
as foun
d
Figure 1.3. Species accumulation curves for the fi ve locations surveyed
in Northeast Madagascar. Logarithm
ic regression lines for each location show predicted species num
ber with increasing samples, up to 15
samples per location.
Figure 1.2. Coral species diversity in Northeast Madagascar in relation to other sites in the W
estern Indian Ocean.
Chapter 1
20Rapid Assessm
ent Program
on th
e steep slop
es insid
e the b
ay, both
on th
e inn
er south
sid
e (site 1) an
d th
e fringin
g/patch
reef on th
e inn
er north
sid
e (sites 2 an
d 3
). Sites 10 an
d 1
6 (L
oky Bay an
d V
ohem
ar, resp
ectively) were th
e high
est-diversity exp
osed ou
ter reef sites. Sites at A
nd
ravina h
ad low
diversity, d
ue to th
e simp
le top
ograph
y and
combin
ation of w
ave and
sedim
ent stress.
Samp
ling for sp
ecies was n
ot focused
on com
pilin
g ind
i-vid
ual-site d
iversity lists, so the sites at th
e lower en
d of th
e scale are n
ot show
n as th
ese do n
ot represen
t true estim
ates of site d
iversity.Site 1
1A
, in L
oky Bay, w
as an exten
sive low-d
iversity stan
d at u
p to 6
m d
epth
, covering ten
s of hectares, of
staghorn
Acropora an
d G
alaxea astreata to >80%
cover. Th
is site w
as the largest con
tinu
ous coral assem
blage in
the su
rvey area. It w
as extensively b
leached
at the tim
e of surveys, w
ith
some m
ortality. Neverth
eless, recovery from th
e bleach
ing is
expected
to be good
.
Coral species composition
Acropora w
as the m
ost diverse gen
us in
the su
rveys, with
53 sp
ecies, followed
by Mon
tipora with
24, F
avia with
13 an
d P
avona, F
avites, Gon
iastrea and
Porites w
ith 1
0 sp
e-cies each
. More th
an h
alf the gen
era (35 ou
t of 61) h
ad 3
or few
er species p
er genu
s. Th
e species record
ed in
this stu
dy
are comp
ared w
ith p
rior records from
the fi rst m
arine R
AP
in
the n
orthw
est (McK
enn
a et al. 2005, V
eron &
Tu
rak 2005) in
Ap
pen
dices 2
and
3.
Th
is show
s a strong d
iff erence b
etween
that stu
dy an
d
this on
e, prin
cipally related
to taxonom
ic meth
odology an
d
focus of th
at stud
y pu
rely on taxon
omy, an
d th
is one on
fi eld
iden
tifi cation an
d ecology. In
this stu
dy th
e genera A
cropora, F
un
gia, Cycloseris, A
lveopora, Gon
iopora, and
Porites are less
well rep
resented
, particu
larly Porites, for w
hich
only 1
/3 th
e n
um
ber of sp
ecies were id
entifi ed
here com
pared
to Veron
&
Tu
rak 2005. Th
is is p
rincip
ally du
e to the d
iffi cult of in
-w
ater iden
tifi cation of th
ese genera. A
qu
antitative an
alysis of th
e stud
ies reveals (Ap
pen
dix 2
):
• C
oral species record
ed in
both
RA
P 1
and
this stu
dy
= 2
00
• C
oral species record
ed in
RA
P 1
bu
t not in
this stu
dy
= 1
16. O
f these, 9
3 are p
otentially id
entifi cation
issues,
most com
mon
ly of diffi cu
lt to recognize sp
ecies that
this au
thor w
ould
lum
p w
ith an
other com
mon
species,
17 w
ere not seen
in th
is stud
y bu
t well kn
own
by th
is au
thor so u
nlikely to h
ave been
lum
ped
un
der an
other
species, an
d 4
have b
een syn
onym
zed.
• C
oral species record
ed in
RA
P3 (th
is stud
y) bu
t not
RA
P1 (V
eron &
Tu
rak 2005) w
ere a total of 53 sp
ecies. O
f these, 3
1 m
ay be d
ue to syn
onym
y issues w
ith grou
p
A1, 2
0 are w
ell know
n to th
is auth
or so represen
t true
add
itional sp
ecies, and
2 sp
ecies of wh
ich on
e is new
ly d
escribed
(Psam
mocora albopicta, B
enzon
i 2006) an
d
one th
at has n
ot been
recorded
in th
e region for d
ecades
(Craterastrea laevis), m
ost likely bein
g lum
ped
with
a L
eptoseris sp. as was in
itially don
e du
ring collection
in
this stu
dy.
Figure 1.4. Coral species richness at individual sites in Northeast Madagascar. The x axis shows the site num
ber (see Site list on page 20) followed by a letter showing its location: A – Am
bodivahibe, L – Loky Bay, K – Ankao, V –Vohemar, and R – Andravina. Two sites had m
ultiple samples – sites 6 (2 sam
ples) and 1 (3 sam
ples), and the combined species richness for those sam
ples is also shown. Lowest-diversity sites are not shown, as sampling at these sites focused
on adding new species to the list, not on a whole-site diversity sample.
Corals of northeast Madagascar
21 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
• R
AP
1 rep
orted 5
7 corals from
species lists for M
adagas-
car that w
ere not record
ed u
nd
erwater. O
f these, 1
9 w
ere ob
served in
this stu
dy, an
d 3
6 w
ere also not ob
served in
th
is stud
y. Fu
rther, R
AP
1 rep
orted a fu
ll speies list for
the region
(Western
Ind
ian O
cean), of w
hich
13 w
ere n
ot observed
in eith
er RA
P1 or R
AP
3
Com
parin
g the sp
ecies iden
tifi ed in
each stu
dy: 321 in
the
fi rst RA
P an
d 281 in
this on
e, ie. a diff eren
ce of 40 species,
and th
e diff eren
ces in m
ethod
ology between
the tw
o, it is likely th
at the coral fau
na are sim
ilar between
the tw
o sides of
the n
orthern
tip of M
adagascar. F
urth
er, the exp
ected sp
ecies rich
ness from
the M
ichaelis-M
enten
curve (fi g. 1.2) su
ggests a sp
ecies richn
ess of 313 species, closer to th
at foun
d in
the fi rst
RA
P.
Key speciesK
ey species of in
terest are listed b
elow u
nd
er the h
eadin
gs region
al end
emics, u
nd
escribed
species, region
al species
with
un
certain ran
ges and
validity, ran
ge extension
s (cer-tain
and
un
certain) an
d n
otes on taxon
omy/syn
onym
y and
term
inology.
Regional endemics, highly restricted, rare
Th
ese are the m
ost high
ly restricted sp
ecies recorded
.C
raterastrea laevis; this sp
ecies is not illu
strated in
most
texts, inclu
din
g Veron
(2000), as it is very p
oorly know
n
in th
e literature. It w
as collected in
this stu
dy at V
ohem
ar an
d A
nd
ravina, b
ut n
ot recognized
un
til surveys 2
mon
ths
later by F. Ben
zoni, in
Mayotte, w
here a secon
d p
opu
lation
is abu
nd
ant in
the lagoon
. In th
e surveys h
ere it was at low
/u
ncom
mon
den
sities and
recorded
from th
ree sites (16, 1
7
and
24 in
three location
s Voh
emar, A
nd
ravina an
d A
nkao).
An
omastraea irregu
laris (Maren
zeller 1901)
; extremely
rare, foun
d at tw
o sites in A
nd
ravina (1
7, 1
8), an
d seen
p
reviously by th
is auth
or in M
asoala abou
t 200 km
south
in
Mad
agascar and
a site on th
e Ken
ya coast.
Regional endemics, broadly distributed but rare
Th
ese species are en
dem
ic to the W
estern In
dian
Ocean
, bu
t broad
ly distrib
uted
thou
gh th
ey are mostly rare at in
divid
ual
locations.
Gyrosm
ilia interru
pta (Eh
renberg 1
834); locally rare b
ut
wid
ely distrib
uted
in th
e Western
Ind
ian O
cean (W
IO), th
e largest p
opu
lations of th
is species h
ave been
recorded
on th
is su
rvey, in L
oky Bay in
particu
lar (Sites 7, 8
, 9 in
particu
lar, also sites 2
, 5 an
d 2
2). It sh
owed
extensive b
leachin
g du
ring
the su
rvey.H
orastrea indica (P
ichon
1971); also locally rare an
d w
ide-
spread
in th
e WIO
(more w
idesp
read th
an G
. interru
pta), w
as wid
ely distrib
uted
in N
ortheast M
adagascar b
ut n
ot as locally ab
un
dan
t as G. in
terrupta (Sites 4
, 5, 8
, 10, 1
7, 1
8).
Regional endemics, broadly distributed but com
mon
Th
ese species are en
dem
ic to the W
estern In
dian
Ocean
, bu
t com
mon
. Th
ey were record
ed at m
any sites in
this su
rvey
and
wou
ld likely b
e foun
d at th
e majority of sites w
ith su
f-fi cien
t samp
ling.
Acropora appressa (E
hren
berg 1
834); kn
own
main
ly from
the sou
thern
African
coast, now
recorded
as wid
espread
in
the SW
IO an
d likely exten
din
g as far north
as Ken
ya.A
cropora branchi (R
iegl 1995); d
escribed
from Sou
th A
fri-can
and
south
ern M
ozambican
coasts, is wid
espread
in th
e W
IO (see R
iegl 1995a,b
).C
oscinaraea sp.A
un
describ
ed; a very com
mon
species of
Coscin
araea in th
e WIO
, bu
t likely un
describ
ed. A
nam
e has
been
given to it b
y M. C
laerebou
dt from
the G
ulf of A
den
, bu
t not form
ally registered taxon
omically. N
oted b
y Veron
&
Tu
rak (2005).
Stylophora madagascaren
sis (Veron
2000); d
escribed
by
JEN
Veron
from M
adagascar, th
is growth
form w
as very com
mon
, how
ever there is con
siderab
le confu
sion in
n
omen
clature for th
is genu
s, and
the statu
s of this as a sp
e-cies is cu
rrently u
nresolved
(Allen
Ch
en, p
ers. comm
.).P
lesiastrea devantieri (V
eron 2
000); on
ly recently d
escribed
from
the G
ulf of A
den
, this sp
ecies is wid
espread
and
com-
mon
throu
ghou
t the W
IO.
Pectin
ia africanus (V
eron 2
000); on
ly recently d
escribed
, th
is species is w
idesp
read an
d com
mon
throu
ghou
t the
WIO
. Th
e wid
espread
P. lactuca h
as only rarely b
een seen
by th
is auth
or.
Undescribed species A
single sp
ecies from th
is survey, th
at has n
ot been
recorded
by th
e auth
or elsewh
ere, except p
erhap
s in th
e Com
oros. Its statu
s is not kn
own
. Samp
les have b
een collected
for genetic
stud
ies.E
chinopora sp.(cf. sm
all corallites) – very sm
all corallites an
d gen
erally small corallu
m, th
is species w
as seen com
-m
only on
fore reef slopes b
ut n
ot initially n
oted. It w
as record
ed p
rimarily at sites 1
4 an
d 1
5 in
Voh
emar.
Regional species, uncertain ranges and validityTh
ese fou
r species are gen
erally poorly kn
own
du
e to restricted
distrib
ution
s and
/or have b
een on
ly recently
describ
ed. In
termed
iates betw
een th
ese and
other sp
ecies in
the tw
o genera ap
pear com
mon
and
are very hard
to d
istingu
ish u
nd
erwater an
d b
etween
diff eren
t sets of surveys.
Fu
rther w
ok is need
ed on
these.
Gon
iastrea colum
ella (Crosslan
di 1
948);
Gon
iastrea deformis (V
eron 2
000)
Platygyra carn
osus (V
eron 2
000)
Platygyra crosslan
di (Matth
ai 1928)
Range extensions, certainTh
ese sp
ecies have n
ot previou
sly been
recorded
from M
ada-
gascar, nor th
e WIO
, except for T
urbin
aria irregularis, w
hich
V
eron &
Tu
rak reported
from th
e 1st R
AP
in 2
002. Th
ey are
very distin
ctive, and
the ran
ge extension
s are certain.
An
acropora pillai has on
ly recently b
een id
entifi ed
and
d
escribed
from th
e Coral T
riangle region
by JE
N V
eron
(2000, 2
002). Seen
at one site, 1
6, in
Voh
emar. A
. forbesi has
Chapter 1
22Rapid Assessm
ent Program
been
reported
from M
adagascar, b
ut w
ith a m
uch
thicker
bran
chin
g growth
form.
Echin
ophyllia echinata (Saville-K
ent 1
871); rep
orted from
th
is survey, p
reviously u
nrecogn
ized in
the W
IO.
Echin
ophyllia echinoporoides (V
eron &
Pich
on 1
980);
reported
from th
is survey, an
d also from
the C
omoros in
M
arch 2
010.
Turbin
aria irregularis ( B
ernard
); very comm
on at a few
sites in
Loky B
ay (8, 9
, 10) an
d a few
colonies in
Voh
emar,
high
ly distin
ctive.
Range extensions, uncertainD
iffi culties in
species id
entifi cation
du
e to environ
men
tal variab
ility and
pop
ulation
diff eren
ces over large distan
ces m
ean th
at some fi eld
iden
tifi cations can
not b
e fully con
-fi rm
ed w
ithou
t skeletal or genetic w
ork. Th
e followin
g iden
-tifi cation
s were m
ade w
ith h
igh con
fi den
ce, bu
t the sp
ecies h
ave not p
reviously b
een rep
orted from
this region
.A
canthastrea regu
laris A
canthastrea rotu
ndofl ora
Acan
thastrea subechin
ata; this is a d
istinctive form
that h
as been
seen by th
e auth
or in m
ultip
le locations an
d is reaso-
nab
ly certain.
Favites acu
ticolis; this is a form
of Favites th
at may b
e a colou
r morp
h of F. pen
tagona rath
er than
F. acuticolis.
Samp
les have b
een taken
for genetic id
entifi cation
.H
ydnophora pilosa; th
is species h
as not b
een rep
orted from
th
e WIO
, bu
t form com
mon
ly reported
as H. m
icroconos
here is m
ore similar to H
. pilosa than
the clean
ly hem
isph
eri-cal H
. microcon
os of the P
acifi c.M
ycedium
um
bra (Veron
2000); h
igh d
iversity in th
e form
of Mycediu
m sp
ecies is comm
on p
articularly in
the h
igh-
diversity region
iden
tifi ed h
ere (fi g. 1.2
). More th
an 2
forms
of Mycediu
m are likely p
resent, b
ut it is n
ot certain th
at M.
um
bra is the correct n
ame.
Pocillopora w
oodjonesii; sp
atulate form
s of Pocillopora are
seen, likely of th
is species.
Sandalolitha robu
sta; not yet rep
orted from
this region
.
Revized generaTh
e gen
us P
samm
ocora has u
nd
ergone m
ajor revisions, w
ith
a strong focu
s on th
e Gu
lf of Ad
en an
d In
dian
Ocean
, as w
ell as samp
les from th
e West an
d C
entral P
acifi c region
(Ben
zoni et al. 2
010). Several sp
ecies have b
een syn
ony-
mized
, and
a new
one d
escribed
.P
samm
ocora albopicta (Ben
zoni 2
006); n
ew sp
ecies, d
escribed
from sam
ples from
Om
an an
d th
e Persian
Gu
lf, an
d th
e Western
Pacifi c. F
urth
er specim
ens h
ave been
seen
by this au
thor in
Djib
outi, an
d in
this su
rvey in L
oky Bay.
Psam
mocora con
tigua (E
sper 1
797); sen
ior nam
e inclu
din
g P. obtu
sangu
la as one sp
ecies (Stefani et al. 2
008).
Psam
mocora profu
ndacella (G
ardin
er 1898); sen
ior nam
e in
clud
ing P. haim
eana as on
e species (Stefan
i et al. 2008).
Synonymy/uncertain status
Favites paraflexu
osa (Veron
2000); a very u
ncertain
form,
listed in
this su
rvey, bu
t it may b
e a morp
h of F. fl exu
osa.Favites vasta (K
lun
zinger 1
879); th
is species in
clud
es m
orph
s that resem
ble F
avia rotun
data, a more p
locoid m
or-p
hology, b
ut likely in
the sam
e species.
Blastom
ussa m
erleti (Wells 1
961); a large-p
olyped
form
of Blastom
ussa record
ed h
ere, and
also seen b
y the au
thor in
M
ayotte. It may b
e in a d
iff erent sp
ecies (see Veron
& T
urak
2005, an
d B
enzon
i, pers. com
m.).
Veron
and
Tu
rak (2005) com
piled
a list of seven sp
ecies of n
ote from th
eir surveys in
North
west M
adagascar in
2002.
Th
e followin
g notes in
dicate p
ossible con
nection
s betw
een
species sam
pled
in th
is survey an
d th
ose they m
ention
ed:
1.
a Seriatopora that is d
istinctive an
d com
mon
in som
e tu
rbid
hab
itats; likely S. guttatu
s (Veron
2000). Th
is w
as n
ot recorded
on th
is stud
y, bu
t diffi cu
lties in fi eld
iden
-tifi cation
s of Seriatopora and
Stylophora over gradien
ts of d
epth
and
sedim
ent in
fl uen
ce may h
ave resulted
in
lum
pin
g this sp
ecies with
other Seriatopora, or w
ith Sty-
lophora pistillata or madagascaren
sis. Assu
med
presen
t.
2.
an A
cropora that is com
mon
and
forms large colon
ies in
shallow
exposed
hab
itats; likely Acropora lam
arki. R
ecorded
in th
is stud
y.
3.
a Coscin
araea that is u
ncom
mon
; Likely C
oscinaraea
spA
, recorded
by th
is auth
or throu
ghou
t East A
frica and
m
ost comm
on in
north
ern K
enya, an
d likely eq
uivalen
t to a n
on-form
al species n
amed
by M
. Claereb
oud
t from
the coast of O
man
.
4.
a Turbin
aria that is rare b
ut also kn
own
from In
don
esia; L
ikely Turbin
aria irregularis. R
ecorded
in th
is stud
y, not
seen b
y this au
thor an
ywh
ere else in th
e western
Ind
ian
Ocean
.
5.
a Blastom
ussa th
at is very distin
ctive; recorded
here as
Blastom
ussa m
erletti, a distin
ctive growth
form, also
noted
from M
ayotte by a colleagu
e (F. Ben
zoni, p
ers. com
m.).
6.
a Platygyra-like faviid
that w
arrants sep
aration as a n
ew
genu
s—th
is genu
s is not en
dem
ic to Mad
agascar and
w
ill contain
two d
escribed
species cu
rrently p
laced in
P
latygyra; possib
ly P. carnosu
s, recorded
in th
is stud
y an
d m
ore broad
ly in th
e WIO
. See also comm
ents on
P
latygyra/Gon
iastrea species (ab
ove);
7.
an A
lveopora that is rare an
d restricted
to mu
dd
y sub-
strates; no ob
servations m
ade relatin
g to this sp
ecies, as A
lveopora are very diffi cu
lt to iden
tify in th
e fi eld;
In ad
dition
, they n
oted an
un
describ
ed C
yphastrea species,
bu
t no ob
servations w
ere mad
e of this on
this su
rvey.
Corals of northeast Madagascar
23 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Genetic and skeletal sam
plesF
urth
er work still n
eeds to b
e don
e to verify the valid
ity of som
e of the sp
ecies assigned
in th
is stud
y in p
articular
becau
se skeletal and
genetic sam
ples w
ere mad
e that h
ad n
ot yet b
een an
alyzed at th
e time of w
riting th
is report. Th
e list
of species an
d sam
ples is in
clud
ed in
Ap
pen
dix 4
. Processin
g an
d an
alyzing th
ese samp
les shou
ld b
e a high
priority, as se-
veral of the sam
ples are for u
ncertain
or un
describ
ed sp
ecies, w
ith th
e high
est priority b
eing th
e followin
g:
1.
Craterastrea laevis, lab
eled u
nd
er Leptsoeris sp., to
confi rm
its presen
ce and
for pop
ulation
analysis w
ith
Mayotte.
2.
Acropora sp
p, to obtain
a more com
preh
ensive an
d reli-
able list of sp
ecies
3.
An
omastrea irregu
laris, Horastrea in
dica, Gyrosm
ilia in
terrupta –
genetic in
formation
on region
al end
emic
genera.
4.
Blastom
ussa m
erletti, Coscin
araea sp. nov., P
latygyra sp
p., Stylophora spp
– u
nresolved
or un
certain sp
ecies/region
al iden
tifi cations
CON
CLUSIO
NS A
ND
RECOM
MEN
DATIO
NS
Th
e diversity of coral sp
ecies in N
ortheast M
adagascar, w
ith
281 id
entifi ed
in th
e water an
d p
redicted
to be 3
23 sp
ecies, is sligh
tly lower th
an rep
orts of the h
ighest d
iversity sites in
North
west M
adagascar in
clud
ing N
osy Be an
d N
osy Mitsio
(Veron
and
Tu
rak 2005), b
ut h
igher th
an th
e Nosy H
ara area at an
equ
ivalent latitu
de an
d scale ju
st W of D
iego on
the N
orthw
est coast. How
ever, consid
ering th
e greater taxo-n
omic resolu
tion of th
e North
west M
adagascar su
rveys, it m
ay be con
clud
ed th
at the reefs su
rveyed in
the N
ortheast in
th
is stud
y are at an eq
uivalen
t level of diversity to th
ose reefs.A
t a regional scale, N
ortheast an
d N
orthw
est Mad
agascar grou
p w
ith th
e high
est diversity sites in
S Tan
zania an
d N
M
ozambiq
ue, len
din
g sup
port to h
ypoth
esis of the exis-
tence of h
igh d
iversity center for th
e Western
Ind
ian O
cean
(WIO
) in th
is region, coverin
g the N
orthern
Mozam
biq
ue
chan
nel. A
ll other sites in
the w
estern, cen
tral and
eastern
Ind
ian O
cean, an
d th
e Red
Sea, have a low
er diversity of
hard
corals.Th
e su
rvey region h
as the largest p
opu
lations of key
regional en
dem
ic genera an
d sp
ecies – C
raterastrea laevis, A
nom
astrea irregularis, H
orastrea indica an
d G
yromsilia
interru
pta – th
at the au
thor h
as seen in
the W
IO. W
hile it
is un
clear wh
at the level of en
dem
ism is d
istingu
ishin
g nor-
thern
Mad
agascar from oth
er parts of th
e WIO
(3 of th
ese gen
era are foun
d q
uite b
roadly d
istribu
ted in
the region
), it ap
pears to su
pp
ort rare species th
at are not fou
nd
in oth
er location
s.
Am
bod
ivahib
e and
Loky B
ay have th
e high
est level of d
iversity of the location
s surveyed
, as well as th
e high
est d
iversity sites. Th
is is likely a result of th
eir dep
th th
at m
aximizes h
abitat d
iversity and
diversifi cation
of nich
es for corals to occu
py, an
d p
rovides an
escape from
the p
ervasive sed
imen
tation th
at infl u
ences sh
allower sites. N
evertheless,
the ou
ter reef sites at Voh
emar an
d th
e sedim
ent-in
fl uen
ced
sites at An
dravin
a had
distin
ctive coral assemblages.
Key focal sp
ecies for conservation
plan
nin
g in th
e survey
region in
clud
e the follow
ing:
• th
e regional en
dem
ic mon
ospecifi c gen
era - Craterastrea
laevis, An
omastrea irregu
laris, Horastrea in
dica, Gyros-
milia in
terrupta
• ad
dition
al species eith
er poorly kn
own
or represen
ting
signifi can
t range exten
sions of rare/p
oorly know
n sp
e-cies: A
nacropora pillai, T
urbin
aria irregularis, P
samm
o-cora albopicta, E
chinopora sp.(cf. sm
all corallites),
Sites with
distin
ctive coral comm
un
ities and
/or of key im
portan
ce to the ab
ove species in
clud
e the follow
ing, w
ith
the m
ost imp
ortant sites (see Sites tab
le in “R
eport at a
Glan
ce,” page 1
3) b
eing in
bold
/un
derlin
ed:
Am
bod
ivahib
e A
mb
odivah
ibe S
(2), A
mbod
ivahib
e N (5
) L
oky Bay
South
(6), n
orthw
est (8/9
), 10-n
orth,
11-A
cropora coral garden
Voh
emar
inn
er chan
nel (1
4), ou
ter north
(15), ou
ter sou
th (1
6)
An
dravin
a sou
th p
oint (1
7), A
nd
ravina rock (1
8)
An
kao A
nkao n
ortheast (2
2), A
nkao B
ay (fringin
g reef, 2
4)
REFERENCES
Ben
zoni, F. 2
006. P
samm
ocora albop
icta sp. nov., a n
ew sp
e-cies of Scleractin
ian C
oral from th
e Ind
o-West P
acifi c (Scleractin
ia; Siderastreid
ae). Zootaxa 1
358: 4
9–57.
Ben
zoni, F., F. Stefan
i., M. P
ichon
and
P. Galli. 2
010. Th
e
nam
e game: m
orph
o-molecu
lar species b
oun
daries in
th
e genu
s Psam
mocora (C
nid
aria, Scleractinia). Z
oo-logical Jou
rnal of th
e Lin
nean
Society: 1-3
6.
Clarke, K
.R. an
d R
.N. G
orley. 2006. P
RIM
ER
v6: U
ser M
anu
al/Tu
torial. PR
IME
R-E
, Plym
outh
Keatin
g, K.A
. 1998. E
stimatin
g species rich
ness: th
e M
ichaelis-M
enten
mod
el revisited. O
ikos 81: 4
11-4
16.
Mah
aravo J., T. A
. Oliver. an
d A
. Rab
earisoa. In p
ress. A
Rap
id M
arine B
iodiversity A
ssessmen
t of North
east M
adagascar. C
onservation
Intern
ational. A
ntan
anarivo.
Mad
agascar.M
cKen
na, S., G
.R. A
llen an
d H
. Ran
drian
asolo (eds.).
2005. A
Rap
id M
arine B
iodiversity A
ssessmen
t of the
Coral reefs of N
orthw
est Mad
agascar. RA
P B
ulletin
of
Chapter 1
24Rapid Assessm
ent Program
Biological A
ssessmen
t 31. C
onservation
Intern
ational,
Wash
ington
DC
, USA
.R
iegl, B. 1
995a. A
revision of th
e hard
coral genu
s Acrop
ora O
ken, 1
815 (Scleractin
ia: Astrocoen
iina: A
croporid
ae) in
south
-east Africa. Z
oological Journ
al of the L
inn
ean
Society 113: 2
49–288.
Riegl, B
. 1995b. D
escription
of four n
ew sp
ecies in th
e hard
coral gen
us A
cropora O
ken, 1
815 (Scleractin
ia: Astro-
coeniin
a: Acrop
oridae) from
south
-east Africa. Z
oologi-cal Jou
rnal of th
e Lin
nean
Society 113: 2
29–247.
Shep
pard
, C. an
d D
. Obu
ra. 2004. C
orals and
reefs of Cos-
moled
o and
Ald
abra atolls: exten
t of dam
age, assem-
blage sh
ifts and
recovery followin
g the severe m
ortality of 1
998. Jou
rnal of N
atural H
istory 39(2
): 103–121.
Stefani, F., F. B
enzon
i, M. P
ichon
, G. M
itta and
P. Galli.
2008. G
enetic an
d m
orph
ometric evid
ence for u
nre-
solved sp
ecies bou
nd
aries in th
e coral genu
s Psam
-m
ocora (Cn
idaria; Scleractin
ia). Hyd
robiologia 5
96:
153–172.
Veron
, J. E. N
. and
E. T
urak. 2
005. R
eef corals of north
-w
est Mad
agascar. In: M
cKen
na, S., G
.R. A
llen an
d
H. R
and
rianasolo (ed
s.) A R
apid
Marin
e Biod
iversity A
ssessmen
t of the C
oral reefs of North
west M
adagascar.
RA
P B
ulletin
of Biological A
ssessmen
t 31. C
onservation
In
ternation
al, Wash
ington
DC
, USA
. Pp. 2
3-3
0.
Veron
, J. E. N
. 2000. C
orals of the W
orld. A
ustralian
Insti-
tute of M
arine Scien
ce, Tow
nsville. A
ustralia.
Wallace, C
. 1999. Stagh
orn corals fo th
e World
. A revi-
sion of th
e genu
s Acrop
ora. Mu
seum
of Trop
ical Q
ueen
sland
, Au
stralia. CSIR
O P
ublish
ing.
25A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Map and Photos
Top. Map of M
adagascar, showing the northern tip (left) and the extent of the three marine RAPs (RAPs 1 and 2 shown in black boxes, the 5 locations of
RAP 3 shown in red boxes and insets below). The 5 locations in RAP 3 (from north to south ) were Am
bodivahibe in the north, which was surveyed during RAP2 (upper right inset), Loky Bay, Ankao island and Andravina Bay (m
ain inset) and Vohemar (lower left inset). Vohem
ar town is located on the southern shore of the entrance to the bay. Survey sites are shown as red triangles (reefs) and green circles (algae and seagrass, see Table 1 on page 7).
Map and Photos
26Rapid Assessm
ent Program
RAP team. Left to right - back: Keith Ellenbogen, Giuseppe di Carlo,
Monica Tom
bolahy, Seabird McKeon, David Obura. Front: Tom
Oliver, Bem
ahafaly Randriamanantsoa. M
elita Samoilys, Jean M
aharavo. Photo by Keith Ellenbogen/ILCP.
Typical exposed and slope comm
unity dominated by large tabular
Acropora spp. Photo by Keith Ellenbogen/ILCP.
Artisanal fi shermen, operating from
local canoes powered by paddle or sail are the dom
inent users of coral reefs in NE Madagascar. Photo by Keith
Ellenbogen/ILCP.
Seagrasses on shallow reef fl ats. Photo by Keith Ellenbogen/ILCP.
Coastal village, in Loky Bay. Photo by Keith Ellenbogen/ILCP.
Air roots of mangrove trees in shallow water seagrass bed. Photo by Keith
Ellenbogen/ILCP.
Map and Photos
27 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Basket star – Euryale aspera. Photo by David Obura.
Anomastrea irregularis. Photo by David Obura.
Anacropora pillai. Photo by David Obura.
Craterastrea laevis. Photo by David Obura.
Gyrosmilia interrupta. Photo by David Obura.
Horastrea indica. Photo by David Obura.
Map and Photos
28Rapid Assessm
ent Program
Apolemichthys trim
acula. Photo by Keith Ellenbogen/ILCP.
The Acropora comm
unity at site 11, the most highly bleached site found,
though with little mortality. Photo by Keith Ellenbogen/ILCP.
Cryptochirid crab. Photo by Keith Ellenbogen/ILCP.
Bleached and normal branches of Acropora palifera. Photo by Keith
Ellenbogen/ILCP.
Chlorurus strongylocephalus. Photo by Keith Ellenbogen/ILCP.
29A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Chapter 2
Reef fi shes of northeast Madagascar
Melita Sam
oilys & Bem
ahafaly Randriamanantsoa
INTRO
DU
CTION
Com
preh
ensive su
rveys and
analyses of coral reef fi sh
diversity are few
in th
e Western
Ind
ian
Ocean
(WIO
). Con
sequ
ently ou
r un
derstan
din
g of pattern
s in reef fi sh
diversity an
d sp
ecies rich
ness across th
e WIO
region is still p
oor. Detailed
species in
ventories are lim
ited to certain
location
s from M
adagascar (A
llen in
McK
enn
a et al. 2005), th
e Mascaren
e Island
s (Fricke
1999) an
d th
e Esp
arses Island
s (Ch
aban
et and
Du
rville 2005), an
d au
thoritative taxon
omic
sources b
uilt on
Mu
seum
records an
d collection
s datin
g back to th
e nin
eteen cen
tury p
ar-ticu
larly from th
e South
African
Institu
te for Aqu
atic Biod
iversity (e.g Smith
and
Heem
stra 1995).
A region
centred
aroun
d sou
thern
Tan
zania/n
orthern
Mozam
biq
ue an
d N
orthern
Mad
agas-car h
as been
prop
osed as a cen
tre of coral diversity in
the W
IO, d
riven b
y the In
dian
Ocean
cu
rrent p
atterns (O
bu
ra 2008). It is likely th
at coral reef fi shes follow
similar grad
ient p
atterns
(Ch
aban
et and
Du
rville 2005) an
d th
is hyp
othesis is cu
rrently u
nd
er investigation
by C
OR
-D
IO E
ast Africa’s “C
ore region p
roject: Is there a W
IO coral trian
gle?” fun
ded
by th
e Western
In
dian
Ocean
Marin
e Science A
ssociation, W
IOM
SA (O
bu
ra et al 2008). Th
e n
orthern
tip
of Mad
agascar represen
ts a key location in
this p
roposed
centre of coral reef d
iversity with
th
e fi rst CI m
arine R
AP
in 2
002 estab
lishin
g the n
orthw
est tip of M
adagascar as h
aving th
e h
ighest d
iversity of hard
corals of any site in
the In
dian
Ocean
, and
over 750 reef-associated
fi sh
species (M
cKen
na et al. 2
005). T
o date on
ly the n
orthw
estern sid
e of Mad
agascar has b
een
surveyed
, therefore th
e curren
t RA
P (N
o. 3) set ou
t to record sp
ecies diversity in
reef fi shes on
th
e far north
eastern coastlin
e of Mad
agascar (see Map
on p
age 25).
In con
trast to fi sh d
iversity stud
ies, man
y surveys an
d an
alyses of pattern
s in th
e pop
ula-
tion ab
un
dan
ce of coral reef fi shes across th
e WIO
have b
een d
one, w
ith a grow
ing literatu
re on
the eff ects of fi sh
ing (M
angi an
d R
oberts 2
007, M
cClan
ahan
et al 2009), th
e imp
acts of m
arine p
arks and
reserves (McC
lanah
an et al. 2
009), an
d th
e eff ects of coral reef bleach
ing
(Grah
am et al. 2
007, 2
008) on
reef fi sh com
mu
nities an
d ab
un
dan
ce. Certain
troph
ic group
s of reef fi sh
, particu
larly the h
erbivores, h
ave been
show
n to p
lay a critical role in coral reef
resilience by con
trolling m
acro-algal comm
un
ities and
preven
ting coral-algal p
hase sh
ifts (B
ellwood
et al. 2004, H
ugh
es et al. 2005). Th
e IU
CN
Clim
ate Ch
ange an
d C
oral Reefs
workin
g group
(http
://cms.iu
cn.org/cccr), h
as outlin
ed p
rotocols to qu
antify resistan
ce and
resilien
ce ind
icators for coral reefs, inclu
din
g reef fi shes (G
reen an
d B
ellwood
2009), w
hich
serve as u
seful in
dicators of reef h
ealth statu
s and
vuln
erability. Th
ese attrib
utes p
rovided
a secon
d an
d sep
arate focus of th
e RA
P3 fi sh
surveys: to collect d
ata on th
e pop
ulation
abu
n-
dan
ce of selected reef fi sh
taxa to comp
limen
t the coral an
d b
enth
ic reef resilience su
rveys (see C
hap
ter 6) in
order to assess th
e health
of the coral reefs on
this p
reviously u
n-su
rveyed coast
of north
eastern M
adagascar.
Chapter 2
30Rapid Assessm
ent Program
METH
OD
S
Th
e fi sh su
rveys involved
two d
iff erent an
d sep
arate meth
ods
to collect two d
iff erent d
ata-sets, thou
gh w
ere don
e consecu
-tively u
nd
erwater d
urin
g the R
AP
survey d
ives. Th
ese two
meth
ods are d
etailed b
elow for: a) fi sh
diversity an
d b
) fi sh
abu
nd
ance for coral reef h
ealth an
d resilien
ce.
Study sitesR
eef fi sh su
rveys were con
du
cted at th
e fi ve broad
geo-grap
hic location
s of the R
AP
: Am
bod
ivahib
e, Loky B
ay, N
osy An
kao, An
dravin
a, and
Voh
emar, over a d
istance of
aroun
d 1
25km
on th
e north
east coast of Mad
agascar (see M
ap on
page 2
5). T
wen
ty-three d
ives were d
one across
the fi ve b
road geograp
hic location
s to maxim
ise the ran
ge of h
abitats su
rveyed at each
location. W
here p
ossible 2
replicate
survey d
ives were d
one to p
rovide tw
o replicate sites for th
e ab
un
dan
ce surveys an
d to m
aximise ob
servation tim
e for sp
ecies presen
ce/absen
ce recordin
g. In total fi sh
diversity
data w
ere recorded
from 1
6 sites, an
d su
ffi cient fi sh
abu
n-
dan
ce for analysis from
8 sites (T
able 2
.1). D
ives were n
ot d
one at A
nd
ravina d
ue to ear p
roblem
s (MS) an
d th
erefore d
ata were on
ly collected on
snorkel. Sites ran
ged from
0.5
to 24m
in d
epth
.
Reef Fish Diversity
To d
ocum
ent th
e diversity of coral reef fi sh
es, we u
sed a
meth
od d
eveloped
by on
e of us (M
S) for a regional b
io-geograp
hic an
alysis of species d
istribu
tions in
the W
IO b
y C
OR
DIO
East A
frica (Obu
ra et al 2008). Th
e m
ethod
is based
on com
pilin
g a comp
lete species in
ventory of 1
9 fam
i-lies (T
able 2
.2) at each
location. Th
ese fam
ilies were selected
based
on th
e followin
g criteria: largest (of all shore fi sh
es); m
ost diverse; kn
own
ind
icators of biogeograp
hical p
atterns;
inclu
sion of en
dem
ics, rare and
vuln
erable sp
ecies (special
conservation
concern
); amen
able to U
VC
(diu
rnal, n
ot cryp-
tic); of fi shery relevan
ce/reef health
status (T
able 2
.2).
Th
is group
of 19 fam
ilies inclu
des p
otentially arou
nd
460 sp
ecies in total from
the W
IO (A
llen 2
005, D
avidson
et al. 2
006, O
bu
ra 2004, Sam
oilys 1988), an
d rep
resents 5
1%
of th
e total nu
mber of coral reef sp
ecies from 9
2 fam
ilies rep
orted from
Mad
agascar (Allen
2005). Th
is w
as consid
ered
to be b
road en
ough
and
diverse en
ough
to captu
re pattern
s in
diversity of fi sh
es across Mad
agascar’s north
eastern reefs as
well as b
roadly w
ithin
the W
IO region
. T
o sup
plem
ent an
d exp
and
the sp
ecies checklist collated
from
surveys of th
e 19 fam
ilies detailed
above, on
e of us
(BR
) also recorded
species p
resence of an
extra 16 reef asso-
ciated fi sh
families (T
able 2
.2).
Th
e diversity of coral reef fi sh
es was m
easured
from p
re-sen
ce/absen
ce of species ob
tained
from SC
UB
A an
d sn
orkel u
nd
erwater visu
al censu
s (UV
C) su
rveys at the selected
sites of th
e RA
P exp
edition
over a range of coral reef h
abitats an
d
dep
th zon
es. Ou
ter reef slope, b
ack reef and
bay h
abitats
were p
rioritized for th
e survey sites to m
aximize d
iversity, th
ough
surveys on
outer slop
es were lim
ited d
ue to stron
g w
ind
s. A
t each site, each
auth
or recorded
species p
resence/
absen
ce, with
the fi rst au
thor (M
S) recordin
g all species
with
in th
e 19 “core” fam
ilies and
the secon
d au
thor (B
R)
recordin
g species w
ithin
the “extra” 1
6 fam
ilies. After each
d
ive the tw
o observers com
pared
notes an
d sp
ecies observed
u
sing p
hotograp
hs, taxon
omic referen
ces and
ph
otograph
ic gu
ides (see referen
ce list). Species n
ames w
ere later verifi ed
Table 2.1. Study sites for 16 fi sh diversity and 8 abundance surveys, the latter marked A. Num
bers correspond to RAP sites listed and described in Table 1 on page 30.
LocationFish survey sites
RAP site No.Description
Am
bod
ivahib
eA
mbod
ivahib
e Inn
er A
01
Margin
al shallow
reef off man
groves.
Am
bod
ivahib
e S AA
02, A
03
Ou
ter reef
Am
bod
ivahib
e S, chan
nel
A04
In b
ay
Am
bod
ivahib
e N A
A05
Ou
ter reef off island
Loky
Loky S A
A06, A
07
Ou
ter reef south
of Bay en
trance
Loky In
ner
A19
Inn
er bay
Loky N
orth W
est AA
08, A
09
Ou
ter reef off island
Loky N
orthA
10
North
ern ou
ter reef slope
Loky N
N W
A12, A
11B
Back reef off islan
d
Voh
emar
Voh
emar N
orth A
A13, A
15
Ou
ter reef north
of town
Voh
emar In
ner
A14
Insid
e chan
nel on
north
side
An
dravin
aA
nd
ravina B
ay S pt
A17
Narrow
steep ed
ge to rocky shore
An
kaoA
nkao N
orth A
A20, A
23
Reef ed
ge off island
An
kao North
-East
AA
22
Ou
ter reef slope off frin
ging fl at
An
kao South
AA
21A
, A21B
Back reef in
major ch
ann
el betw
een islan
ds
An
kao Bay
A24
Inn
er fringin
g reef on m
ainlan
d
Reef fishes of northeast Madagascar
31 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
from th
e onlin
e Catalog of F
ishes (C
alifornia A
cadem
y of Scien
ces). Tw
o dives resu
lted in
species ob
servations over an
average p
eriod of 7
0 m
ins p
er site, thou
gh at th
ree sites only
30 m
in of ob
servations w
ere possib
le becau
se only on
e dive
was d
one: A
mbod
ivahib
e North
, Am
bod
ivahib
e Ch
ann
el an
d A
nkao N
orth-E
ast. In total a m
inim
um
of 140 m
in
(Voh
emar) an
d a m
aximu
m of 3
95 m
in (L
oky) per b
road
location w
as achieved
for species record
ing.
Reef fi sh abundanceSep
arate surveys of fi sh
abu
nd
ance of selected
taxa were d
one
to contrib
ute to th
e RA
P Su
rvey’s assessmen
t of the h
ealth of
these coral reefs, w
here h
ealth can
be exp
lained
in term
s of th
e reef’s ecological resilience –
its ability to resist th
reats and
to recover to a h
ealthy state w
hen
an im
pact d
oes occur (see
Ch
apter 6
). A
broad
range of taxa w
ere selected for su
rveys that w
ere th
en assign
ed to sp
ecifi c troph
ic group
s relevant in
assessing
resilience (T
able 2
.3). T
axa were categorised
into seven
fun
c-tion
al troph
ic group
s: piscivores, om
nivores, corallivores,
invertivores, p
lanktivores, d
etritivores, and
herb
ivores (Green
an
d B
ellwood
2009, O
bu
ra and
Grim
sditch
2009, L
ieske an
d M
yers 1996, Sam
oilys and
Carlos 2
000). Th
e h
erbivores
were fu
rther b
roken d
own
into six fu
nction
al group
s (Green
an
d B
ellwood
2009): large excavators, sm
all excavators, scrap
ers, grazers, brow
sers and
grazers/detritivores; each
of th
ese group
s are thou
ght to p
lay a diff eren
t ecological role in
coral reef resilience to clim
ate chan
ge. A
n u
nd
erwater visu
al censu
s (UV
C) b
elt transect of
50 ×
5m, con
sidered
one of th
e stand
ards for coral reef
fi shes (B
rock 1982, H
armelin
-Vivien
et al 1985, E
nglish
et al 1
994, Sam
oilys 1997, Sam
oilys and
Carlos 2
000) w
as u
sed to m
easure fi sh
abu
nd
ance. A
t each site (T
able 2
.1) a
combin
ation of tw
o “long sw
ims” (L
S n=2, ap
prox. 4
000m
2) an
d fi ve 5
0 ×
5 m
transects (T
S n=5, 2
50m
2) were cen
sused
, w
here p
ossible. Th
is w
as generally ach
ieved from
two sites
on th
e reef with
in arou
nd
500 –
1000m
of each oth
er. O
ccasionally on
ly one d
ive was p
ossible givin
g LS=
1, T
S=3.
Th
e long sw
im in
volved a tim
ed sw
im of 1
0 m
in coverin
g ap
proxim
ately 200m
along th
e reef parallel to th
e reef crest ran
ging over th
e dep
ths of th
e reef slope b
ut n
o deep
er than
20m
, coun
ting fi sh
with
in a 2
0m
swath
e, dep
end
ing on
visib
ility. Th
is meth
od w
as used
to maxim
ize samp
ling of th
e larger m
obile an
d d
iver – sh
y fi sh (e.g. b
um
ph
ead p
arrotfi sh,
large group
ers, Nap
olean w
rasse and
sharks). A
ny d
epartu
re from
the 2
00 ×
20m
long sw
im area, eg d
ue to stron
g curren
t, w
as recorded
and
the d
ensity estim
ation corrected
accor-d
ingly. O
n th
is basis th
e abu
nd
ance of all sp
ecies/species
group
s listed in
Tab
le 2.3
were cen
sused
.
RESULTS
Diversity
We record
ed a total of 2
71 sp
ecies from 9
6 gen
era based
on
the su
bset of 1
9 core fam
ilies surveyed
, pooled
across the fi ve
locations. Th
is rep
resents 7
4%
of all species rep
orted (3
67)
from th
ese 19 fam
ilies for Mad
agascar based
on record
s da-
ting b
ack to 1891 (A
llen 2
005).
A fu
rther 2
5 sp
ecies from 1
8 gen
era from th
e extra 16 fam
ilies (Tab
le 2.2
) were also ob
served, givin
g a total spe-
cies coun
t of 296 from
114 gen
era observed
in N
ortheastern
M
adagascar in
March
-Ap
ril 2010 (A
pp
end
ix 6).
Th
e Coral R
eef Fish
Diversity In
dex (C
FD
I), develop
ed
by A
llen an
d W
erner (2
002) for estim
ating reef fi sh
diversity
Table 2.2. Families selected for coral reef fi sh diversity surveys for
biogeographic analyses. Top 19 families used for regional biogeographic
analyses (MS); additional 16 fam
ilies used to supplement species
checklist (BR).
Family
Notes
Core Families for Biogeographic Analyses
Lab
ridae (w
rasse)*Serran
idae (grou
pers)
Pom
acentrid
ae (dam
selfi shes)*
Largest an
d m
ost diverse
families of all sh
ore fi shes
Ch
aetodon
tidae (b
utterfl yfi sh
es)* Scarid
ae (parrotfi sh
es)*A
canth
urid
ae (surgeon
fi shes)*
Lu
tjanid
ae (snap
pers)
Pom
acanth
idae (an
gelfi shes)*
Kn
own
ind
icator families
of biogeograp
hical p
atterns
and
/or coral reef health
Leth
rinid
ae (emp
erors)H
aemu
lidae (gru
nts)
Mu
llidae (goatfi sh
es)Sigan
idae (rab
bitfi sh
)N
emip
teridae (b
ream)
Caran
gidae (trevally)
Ad
dition
al families in
clud
ed
for fi shery im
portan
ce
Caesion
idae (fu
siliers)B
alistidae (triggerfi sh
)M
onacan
thid
ae (fi lefi sh)
Ostraciid
ae (boxfi sh
)T
etraodon
tidae (p
uff erfi sh
)
Ad
dition
al families in
clud
ed
to broad
en taxon
omic ran
ge
Supplementary Fam
ilies for Species Inventory
Au
lostomid
ae (trum
petfi sh
)C
irrhitid
ae (haw
kfi sh)
Ep
hip
pid
ae (batfi sh
)H
olocentrid
ae (squ
irrelfi shes)
Kyp
hosid
ae (dru
mm
ers)M
icrodesm
idae (d
artfi shes)
Mu
raenid
ae (eels)Pem
ph
eridae (sw
eepers)
Platycep
halid
ae (fl athead
s)P
riacanth
idae (b
igeyes)Scom
brid
ae (tun
as) Scorp
aenid
ae (scorpion
fi sh)
Sph
yraenid
ae (barracu
da)
Elasm
obranchs:
Dasyatid
ae (stingrays)
Torp
edin
idae (torp
edo rays)
Myliob
atidae (eagle rays)
*= C
FD
I taxa (Allen
and
Wern
er 2002, see text)
Chapter 2
32Rapid Assessm
ent Program
Table 2.3. Taxa surveyed for fi sh abundance with their assigned trophic functional group (after Green and Bellwood 2009, Obura and Grimsditch 2009).
Functional GroupGroup/fam
ilyEnglish nam
e or speciesNotes on feeding habits and selection of species
PiscivoresCarangidae
TrevallyT
op level p
redators, exert top
-dow
n con
trol on low
er troph
ic levels of fi sh
, are very vuln
erable to overfi sh
ing, an
d good
in
dicators of th
e level of anth
ropogen
ic distu
rban
ce (fi shin
g) on
a reef.
Serranid
aeG
roup
ers
Lu
tjanid
aeA
prion viriscen
s
Elasm
obran
chii
Sharks an
d rays
Scombrid
aeT
un
as
Om
nivoresH
aemu
lidae
Sweetlip
sSecon
d-level p
redators w
ith h
ighly m
ixed d
iets inclu
din
g sm
all fi sh, in
vertebrates an
d d
ead an
imals, th
eir presen
ce/ab
sence is also a good
ind
icator of anth
oropogen
ic d
isturb
ance (fi sh
ing).
Leth
rinid
aeE
mp
erors
Lu
tjanid
aeSn
app
ers except A
prion viriscen
s
Corallivores
Ch
aetodon
tidae
Bu
tterfl yfi shO
bligate an
d facu
ltative corallivores are a second
ary in
dicator of coral com
mu
nity h
ealth. 8
species: C
. benn
etti, C
. lineolatu
s, C. m
elann
otus, C
. meyeri, C
. ornatissim
us,
C. trifascialis, C
. trifasciatus, C
. zanzibaren
sis
InvertivoresPom
acanth
idae
An
gelfi shF
eed on
coral comp
etitors such
as soft corals and
spon
ges, th
eir relative abu
nd
ance m
ay be a secon
dary in
dicator of
abu
nd
ance/stab
ility of these grou
ps an
d of a p
hase sh
ift. E
xcept C
entropyge sp
p. wh
ich are grazer-d
etrivores
Balistid
aeT
riggerfi shB
enth
ic triggerfi sh (e.g. Su
ffl amen
spp.)
Ch
aetodon
tidae
Bu
tterfl yfi shN
on-corallivore sp
ecies - all others excep
t the 8
above an
d
H. zoster an
d H
. diphreutes w
hich
are plan
ktivorous
Lab
ridae
Cheilin
us u
ndu
latus
On
ly this lab
rid su
rveyed
PlanktivoresB
alistidae
Triggerfi sh
Resid
ent on
reef surfaces, b
ut feed
in th
e water colu
mn
. Th
eir p
resence/ab
sence m
ay be related
to hab
itat for shelter
and
water colu
mn
cond
itions
Trigger fi sh
in th
e water colu
mn
(eg. Melichthys sp
p., Odon
us
niger)
Ch
aetodon
tidae
Hem
itaurichthys zoster
Hen
iochus diphreu
tes
Acan
thu
ridae
Naso sp
p. >20cm
Large N
aso in w
ater colum
n, excep
t un
icornis an
d tu
berosus
(below
) wh
ich are alw
ays Brow
sers
A
. mata
A
. nubilu
s
A
. thompson
i
Paracan
thuru
s
Caesion
idae
Fu
siliers
Detritivores
Acan
thu
ridae
Cten
ochaetus
Feed
on organ
ic matter in
sedim
ent an
d on
reef su
rfaces, their relative ab
un
dan
ce may b
e an in
dicator of
eutrop
hication
and
cond
itions u
nsu
itable for corals.
Herbivores
Exert th
e prim
ary control on
coral-algal dyn
amics an
d are
imp
licated in
determ
inin
g ph
ase shifts from
coral to algal d
omin
ance esp
ecially in resp
onse to oth
er pressu
res such
as eu
troph
ication, m
ass coral mortality
Large excavators Scarid
aeB
olbometopon
Take few
, large, deep
bites, rem
ove mu
ch su
bstratu
m an
d
play a key role in
bioerosion
C
hloruru
s spp. >
35cm
C
etoscarus bicolor
Small excavators
C
hloruru
s spp. <
35cm
Rem
ove substrate - p
lay a smaller role in
bioerosion
table contin
ued on
next page
Reef fishes of northeast Madagascar
33 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
based
on th
e six most ab
un
dan
t, speciose an
d ch
aracteris-tic fam
ilies of coral reef fi shes (m
arked * in
Tab
le 2.2
) was
calculated
to assess the overall d
iversity of coral reef fi shes
in N
ortheast M
adagascar (T
able 2
.4) an
d to com
pare w
ith
other location
s in M
adagascar an
d n
eighbou
ring cou
ntries.
Th
e CF
DI valu
e of 172 record
ed h
ere for North
east Ma-
dagascar is sim
ilar to that rep
orted for N
orthw
est Mad
agas-car by A
llen from
the R
AP
in 2
002 (A
llen 2
005), su
ggesting
our su
rveys provid
e a good rep
resentation
of the d
iversity of coral reef fi sh
es for the N
ortheast coast, an
d th
at the
diversity of fi sh
es of the tw
o coasts is similar. Th
is d
iversity rep
resents 8
4%
of the total n
um
ber of sp
ecies (204) rep
orted
from M
adagascar from
these 6
families (A
llen 2
005). Th
is
relatively high
species d
iversity is high
er than
that record
ed
in n
eighbou
ring C
omoros (T
able 2
.4), d
espite th
e strong
win
ds th
at limited
our ab
ility to survey th
e outerm
ost SE
facing reef slop
es, and
initial p
erception
s that th
e reefs were
imp
overished
du
e to high
turb
idity. Th
e valu
e for north
ern
Mozam
biq
ue (T
able 2
.4) is b
ased on
a restricted an
d sh
ort su
rvey of only six d
ays on reefs arou
nd
Metu
nd
o Island
(Sam
oilys et al in p
ress) and
therefore rep
resents on
ly a p
artial assessmen
t, bu
t is non
etheless p
rovided
for comp
ari-son
. Data on
coral reef fi sh d
iversity with
in a core region
of
high
biod
iversity in th
e WIO
centered
aroun
d latitu
de 1
0o
S contin
ue to b
e collected as p
art of an on
going C
OR
DIO
E
ast Africa p
roject (Obu
ra et al 2008).
Tab
le 2.5
show
s clearly the sign
ifi cant d
iff erences in
sp
ecies diversity b
etween
sites. Th
e total nu
mber of sp
ecies seen
per site ran
ged from
60 to 1
27 w
ith an
average of 91,
slightly less th
an A
llen’s average of 117 for N
orthw
est Mad
a-gascar (in
2002). Th
e rich
est sites (117-1
27 sp
ecies) were
Loky Sou
th, A
mbod
ivahib
e Inn
er and
South
, and
An
kao
Functional GroupGroup/fam
ilyEnglish nam
e or speciesNotes on feeding habits and selection of species
Scrapers
Scarus sp
p.R
emove algae, sed
imen
t and
other m
aterial by closely
cropp
ing or scrap
ing th
e substrate
H
ipposcarus sp
p.
Browsers
Scaridae
Calotom
us sp
p.F
eed on
large macro-algae
L
eptoscarus sp
p.
Acan
thu
ridae
Naso u
nicorn
is
N
aso tuberosu
s
N
aso spp. <
21cm
Ep
hip
pid
aeB
at fi sh –
Platax sp
p.
Siganid
aeS. can
aliculatu
s
Kyp
hosid
aeR
ud
der fi sh
Grazers
Acan
thu
ridae
Zebrasom
a spp.
graze epilith
ic algal turfs, w
hich
can also lim
it growth
of m
acroalgae
A
. nigrofu
scus
A
canthu
rus sp
p. Sm
all surgeon
species, in
cl. lineatu
s
Siganid
aeSigan
us sp
p.E
xcept Sigan
us can
aliculatu
s
Grazer-detritivores
Acan
thu
ridae
A. blochii
Rin
g tails. Feed
on algal tu
rf, sedim
ent an
d som
e anim
al m
aterial. Similar role to grazers - rem
ove macroalgae
A
. dussu
mieri
A
. leucocheilu
s
A
. nigricau
da
A
. xanthopteru
s
A
. tenn
enti
Pom
acanth
idae
Cen
tropyge spp.
Table 2.3. continued
Table 2.4. Coral reef fi sh diversity index (CFDI) values from M
adagascar and neighbouring countries. CFDI =
total no. species in 6 families:
Chaetodontidae, Pomacanthidae, Pom
acentridae, Labridae, Scaridae and Acanthuridae.
LocationCFDI
Source/date of survey
North
eastern M
adagascar
172
Th
is report / 2
010
North
western
Mad
agascar176
Allen
2005 / 2
002
Com
oros - Gran
de C
omore
and
Moh
eli147
Samoilys u
np
ubl., 2
010
Mozam
biq
ue –
N
. Qu
irimbas
137
Samoilys in
press, 2
010
Chapter 2
34Rapid Assessm
ent Program
Table 2.5. Total number of species per fam
ily per site, sites listed in decreasing order of total species number. Allen’s (2005) num
ber of species for M
adagascar is also shown for comparison – note these values refer to all species records for M
adagascar collated by Allen. Shaded families represent those
used for the CFDI. * = incom
plete surveys (only 30 min).
Family
All Mada-
gascar (Allen)All sites RAP 3
(2010)Loky
SAm
bodi-vahibe Inner
Ankao North
Ambodi-
vahibe SLoky NW
Ankao South
Vohemar
OuterLoky North
Pom
acentrid
ae49
38
18
22
22
14
14
21
14
14
Pom
acanth
idae
96
42
33
63
16
Lab
ridae
73
57
22
25
30
23
21
25
19
23
Ch
aetodon
tidae
21
22
16
11
15
14
12
812
12
Scaridae
22
20
12
11
13
12
10
11
46
Acan
thu
ridae
26
28
18
16
15
16
13
12
16
14
Serranid
ae41
14
45
210
41
31
Leth
rinid
ae18
10
35
24
37
21
Lu
tjanid
ae16
11
76
54
25
22
Caesion
idae
77
44
15
21
32
Haem
ulid
ae9
64
21
11
21
0
Nem
ipterid
ae3
21
10
01
10
0
Mu
llidae
98
24
33
23
30
Siganid
ae4
62
12
22
22
2
Balistid
ae14
12
42
32
50
75
Mon
acanth
idae
97
32
31
30
21
Ostraciid
ae5
21
10
11
02
0
Tetraod
ontid
ae9
81
42
23
12
2
Caran
gidae
23
71
10
01
13
0
TO
TA
L3
67
27
11
27
12
51
22
11
71
06
10
49
89
1
Table 2.5. continued.
Family
Ankao Bay
Ankao North-East*
Loky Inner
Ambodivahibe
S – channel*Loky
North WVohem
ar Inner
Ambodi-
vahibe N*Andravina
Bay*
Pom
acentrid
ae17
13
14
715
13
911
Pom
acanth
idae
23
23
22
21
Lab
ridae
19
18
15
11
14
16
17
9
Ch
aetodon
tidae
13
11
89
86
84
Scaridae
12
88
97
65
2
Acan
thu
ridae
10
912
10
811
716
Serranid
ae2
13
53
01
2
Leth
rinid
ae1
22
21
10
1
Lu
tjanid
ae3
21
33
01
4
Caesion
idae
01
22
00
21
Haem
ulid
ae2
01
01
21
1
Nem
ipterid
ae1
00
00
10
0
Mu
llidae
13
12
12
12
Siganid
ae1
11
01
21
3
Balistid
ae0
21
61
04
0
Mon
acanth
idae
11
00
11
11
Ostraciid
ae0
00
00
11
0
Tetraod
ontid
ae1
12
13
12
1
Caran
gidae
20
10
00
01
TO
TA
L8
87
67
47
06
96
56
36
0
Reef fishes of northeast Madagascar
35 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
North
. Th
e richest location
s were th
e two d
eep b
ay systems
of Loky an
d A
mbod
ivahib
e (Tab
le 2.5
). Th
e lowest d
iversi-ties w
ere recorded
at sites at Voh
emar an
d som
e inn
er sites w
ithin
the b
ays of Am
bod
ivahib
e and
Loky.
Th
e more sp
eciose families ch
aracteristic of coral reefs su
ch as th
e pom
acentrid
s (dam
selfi shes), lab
rids (w
rasses) an
d ch
aetodon
tids (b
utterfl yfi sh
) were m
ore diverse at sites
in L
oky Bay, A
mbod
ivahib
e and
Nosy A
nkao. Th
e d
iver-sity of b
alistids (triggerfi sh
), serranid
s (group
ers), lutjan
ids
(snap
pers) an
d leth
rinid
s (emp
erors) were con
sistently low
an
d th
e carangid
s (trevally) almost ab
sent (T
able 2
.5). Th
ese
latter four fam
ilies are targeted fi sh
ery species, th
erefore the
low d
iversity may refl ect overfi sh
ing. Th
e excep
tions w
ere at N
osy An
kao (wh
ere the d
iversity of lutjan
ids an
d leth
ri-n
ids w
as the h
ighest on
the en
tire survey) an
d at th
e outer
(more exp
osed an
d less tu
rbid
) south
ern site of A
mbod
i-vah
ibe, w
here th
e diversity of serran
ids w
as relatively high
(T
able 2
.5).
TAXO
NO
MIC O
BSERVATIO
NS O
F PARTICU
LAR IN
TEREST
No n
ew sp
ecies were fou
nd
du
ring th
e course of th
is survey.
How
ever, 23 sp
ecies not p
reviously record
ed for M
adagascar
were sigh
ted. Th
ese n
ew record
s may refl ect th
at the reefs
of North
east Mad
agascar provid
e some d
iff erent h
abitat
and
structu
re comp
ared w
ith th
ose of other region
s in th
e cou
ntry.
Notew
orthy sp
ecies observed
inclu
de th
e dam
selfi sh
Pom
acentru
s caeruleopu
nctatu
s, end
emic to M
adagascar an
d
describ
ed by A
llen in
2002 from
North
west M
adagascar. It
was on
ly seen at on
e site in A
mbod
ivahib
e. In ad
dition
, at tw
o sites in L
oky Bay th
e Ind
ian O
cean p
uff erfi sh
Can
thi-gaster petersii (B
iancon
i, 1854) w
hich
Allen
(pers com
m.)
believes sh
ould
be rem
oved from
synon
ymisation
with
C.
solandri, P
late. 1), w
as sighted
. Oth
er interestin
g species of
note in
clud
e:
• C
haetodon dolosu
s, a south
west In
dian
Ocean
chae-
todon
tid, u
sually fou
nd
deep
er than
40m
. Rare - on
e in
divid
ual w
as seen at arou
nd
15m
at Am
bod
ivahib
e.•
Chaetodon
blackburn
ii, end
emic to E
ast Africa, M
ada-
gascar, Mau
ritius an
d R
eun
ion. R
are – on
e seen at
An
dravin
a Bay (by B
R).
• C
haetodon leu
copleura –
“Somali b
utterfl yfi sh”, sou
thern
R
ed Sea an
d W
IO, n
ot previou
sly recorded
south
of Seych
elles. Rare, on
ly one in
divid
ual seen
at Am
bod
iva-h
ibe, site A
02.
• A
mphiprion
latifasciatus –
anem
one fi sh
end
emic to
Mad
agascar and
Com
oros. Com
mon
. •
Zebrasom
a gemm
atum
– a restricted
range en
dem
ic – from
East A
frica (Mozam
biq
ue, Sou
th A
frica,) to C
omoros, M
adagascar, M
auritiu
s and
Reu
nion
. Usu
ally at >
20m
. Rare, on
e observed
in L
oky Bay arou
nd
12m
d
epth
.
• A
canthu
rus gu
ttatus –
wid
espread
Ind
o-Pacifi c sp
ecies bu
t in su
rge zones, n
ot previou
sly recorded
in M
adagas-
car. On
ly seen in
An
dravin
a Bay in
surge at en
trance to
bay in
<1m
, a school of arou
nd
8 in
divid
uals.
• C
oris freiri – a W
IO an
d R
ed Sea sp
ecies. Rare, on
ly one
seen at V
ohem
ar, comm
on in
Com
oros.
Oth
er fi sh sp
ecies generally com
mon
on coral reefs in
the
region w
ere rare. For exam
ple, th
e snap
per L
utjan
us kasm
ira, w
as only seen
on th
ree dives in
Am
bod
ivahib
e and
one in
N
osy An
kao, wh
ile Scarus fren
atus, w
hich
is norm
ally one
of the m
ost comm
on reef scarid
s, was on
ly seen at N
osy A
nkao. A
llen (2
005) fou
nd
similar rarity of L
. kasmira in
N
orthw
est Mad
agascar. We also saw
no P
ygoplites diacanthu
s, a w
idesp
read coral reef an
gel fi sh th
at has b
een record
ed
elsewh
ere in M
adagascar. N
or were th
e parrotfi sh
Calotom
us
spiniden
s or C. carolin
us seen
at any sites, d
espite th
eir ge-n
eral abu
nd
ance on
coral reefs in th
is region (th
ough
notin
g A
llen (2
005) also on
ly saw on
e ind
ividu
al of C. carolin
us in
N
orthw
est Mad
agascar).
Reef fi sh abundance – resilience indicatorsTh
e ab
un
dan
ce of the d
iff erent trop
hic grou
ps varied
con-
siderab
ly betw
een location
s (Figu
re 2.1
) and
was relatively
low. Th
e h
ighest overall ab
un
dan
ce was fou
nd
at Am
bod
i-vah
ibe Sou
th (site A
02, A
03); h
ere the h
ighest d
ensities
were record
ed for th
e generally sm
all species of in
vertivores (ch
aetodon
tids an
d p
omacan
thid
s), 106 fi sh
/1000 m
2, and
p
lanktivores (e.g. b
alistids, acan
thu
rids), 1
69 fi sh
/1000 m
2. Th
is location
also had
the h
ighest d
ensities of sm
all excava-tors (4
3 fi sh
/1000m
2). Overall ab
un
dan
ce was also relatively
high
at An
kao South
, bu
t with
a very diff eren
t represen
tation
of troph
ic group
s. Here th
e herb
ivores were m
ost abu
nd
ant
- the scrap
ers (scarids), 1
14 fi sh
/1000 m
2, and
grazer-detriti-
vores (acanth
urid
s), 94 fi sh
/1000 m
2. At oth
er sites scrapers
were n
ot abu
nd
ant, ran
ging from
17-6
8 fi sh
/1000 m
2. Th
e ab
un
dan
ce of grazers was also gen
erally low across m
ost location
s (Figu
re 2.1
). Th
ere were n
otably very low
den
sities (3
-8 fi sh
/1000 m
2) or a comp
lete absen
ce of the top
troph
ic level grou
p, the p
iscivores (serranid
s, carangid
s, scombrid
s), at all location
s. Th
ere were also very low
den
sities of omn
i-vores (lu
tjanid
s, lethrin
ids an
d h
aemu
lids), ran
ging from
1.6
-44.8
fi sh/1
000 m
2, except at A
nkao Sou
th (F
igure 2
.1).
It is likely that th
ese pattern
s of abu
nd
ance refl ect fi sh
ing
pressu
re from resid
ent fi sh
ers and
migran
t fi shers, an
d
that th
e two b
ays Am
bod
ivahib
e and
Loky w
ere heavily
sedim
ent-aff ected
bays w
here reefs are lim
ited in
growth
and
stru
cture. In
contrast, N
osy An
kao’s reefs mostly lie ou
tside
major in
pu
ts of terrestrial sedim
ent, th
ough
there is a size-
able h
um
an p
opu
lation su
pp
orted b
y the large com
mercial
algal farm. A
pp
arently th
e farm restricts fi sh
ing on
its neigh
-bou
ring reefs an
d m
any local fi sh
ers have sw
itched
to algae farm
ing, so fi sh
ing p
ressure is low
er than
at the oth
er loca-tion
s surveyed
. Neverth
eless, piscivore d
ensities w
ere low an
d
ranged
from 0
- 3 fi sh
/1000m
2 at Nosy A
nkao, su
ggesting
fi shin
g pressu
re is high
. Voh
emar is a region
al capital w
ith
Chapter 2
36Rapid Assessm
ent Program
Figure 2.1. Density of fi sh (mean num
ber per 1000 m2) by trophic group for eight sites. See M
ethods for description of trophic groups.The six herbivore trophic groups are indicated by arrow.
Reef fishes of northeast Madagascar
37 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
noticeab
ly high
anth
ropogen
ic pressu
re from b
oth p
ollution
an
d fi sh
ing, as seen
in its low
diversity an
d low
abu
nd
ance,
especially in
large piscivorou
s fi sh.
Com
parison
s with
other sites su
rveyed by on
e of us (su
r-veys by B
R in
2009, u
np
ubl. d
ata) in M
adagascar (M
asoala, T
amp
olo and
Tan
jona arou
nd
Masoala M
arine P
ark in
North
east Mad
agascar, furth
er south
of Voh
emar, an
d in
Salary in
south
western
Mad
agascar), are usefu
l in in
terpre-
ting th
e levels of abu
nd
ance record
ed d
urin
g the R
AP. F
or exam
ple p
iscivore den
sity in M
asaola Marin
e Park w
as mu
ch
high
er, rangin
g from 1
6 to 3
8 fi sh
/1000m
2. Om
nivore d
en-
sities were also h
igher, ran
ging from
66 –
137 fi sh
/1000m
2, excep
t at Tan
jona (2
7 fi sh
/1000m
2). Th
ese comp
arisons
sup
port th
e observation
that th
e abu
nd
ance of fi sh
ery target grou
ps w
as generally very low
in th
e North
east Mad
agascar R
AP.
Pattern
s in fi sh
abu
nd
ance w
ill also refl ect the h
ealth of
the corals p
articularly levels of m
ortality from b
leachin
g and
stru
ctural com
plexity. Stru
ctural com
plexity is closely related
to reef erosion
wh
ich occu
rs wh
en corals d
ie, therefore coral
mortality an
d loss of com
plexity are stron
gly correlated an
d
together aff ect th
e fi sh com
mu
nity (G
raham
, et al. 2008).
Low
er coral cover/comp
lexity has b
een sh
own
to be closely
correlated w
ith low
er abu
nd
ance of corallivores, p
lanktivores
and
small sized
(<20cm
) fi shes across th
e WIO
(Grah
am, et
al. 2008). Th
e h
ighest corallivore ab
un
dan
ce in th
e RA
P su
r-vey w
as recorded
at Voh
emar (F
igure 2
.1) w
here coral cover
and
structu
ral comp
lexity were also h
ighest (see C
hap
ter 6).
Fu
rther su
pp
ort for correlations b
etween
fi sh ab
un
dan
ce and
reef h
ealth is seen
in th
e high
est den
sities of invertivores an
d
detritivores at V
ohem
ar; these grou
ps are gen
erally <20cm
in
size. Mod
erately high
den
sities of corallivores and
high
coral cover w
ere also recorded
at Nosy A
nkao. H
owever,
the p
lanktivores en
um
erated by G
raham
et al (2008) refer
largely (>90%
) to pom
acentrid
s wh
ich w
ere not cou
nted
h
ere; the p
lanktivores of th
e presen
t survey com
prise acan
-th
urid
s, balistid
s and
caesionid
s (Tab
le 2.3
) wh
ich varied
by
orders of m
agnitu
de b
etween
sites with
high
est den
sities at A
mbod
ivahib
e South
(Figu
re 2.1
). We are yet to u
nd
erstand
clearly h
ow th
ese plan
ktivore taxa respon
d to coral cover/
reef structu
re declin
e, and
therefore th
eir role as ind
icators of clim
ate chan
ge imp
acts. Th
e ab
un
dan
ce of herb
ivorous trop
hic grou
ps su
ch as th
e scrap
ers and
grazers of the scarid
s and
acanth
urid
s (Tab
le 2.3
), that are con
sidered
to play an
imp
ortant role in
main
-tain
ing reefs th
at will b
e more resilien
t to coral bleach
ing,
was n
otably low
. Th
ese den
sities were stark w
hen
comp
ared
with
locations elsew
here in
Mad
agascar: scraper ab
un
dan
ce at M
asaola Marin
e Park ran
ging from
60 –
117 fi sh
/1000m
2, d
ensities th
at were n
ever recorded
du
ring th
e curren
t RA
P
except at A
nkao Sou
th. G
razer abu
nd
ance in
Masaola
Marin
e Park an
d at Salary in
the sou
thw
est of the cou
ntry,
was 3
9 –
123 fi sh
/1000m
2 an
d 1
30 fi sh
/1000m
2, respectively,
comp
ared w
ith th
e curren
t surveys of <
11 fi sh
/1000m
2 at several sites, an
d th
e high
est den
sity recorded
, at Voh
e-m
ar, of only 5
4 fi sh
/1000 m
2 (Figu
re 2.1
). Man
y of the
herb
ivore troph
ic group
taxa fall with
in th
e <20cm
bod
y size class (T
able 2
.3) an
d th
eir den
sities are therefore likely
to be correlated
with
coral cover and
structu
ral comp
lexity. Sin
ce these w
ere mod
erately high
on th
is stretch of coastlin
e (C
hap
ter 6), th
e low ab
un
dan
ces of herb
ivores is surp
rising
and
worryin
g consid
ering th
eir role in facilitatin
g a reef’s recovery from
bleach
ing.
Oth
er reef health
ind
icators relevant to th
e fi sh com
mu
-n
ity are coral colony size an
d algae (m
acro and
turf) cover.
Coral colon
y sizes suggested
a relatively low level of reef d
is-tu
rban
ce over the p
ast few d
ecades at A
mbod
ivahib
e, Loky
Bay, an
d N
osy An
kao (Ch
apter 6
). Th
ese three location
s also con
tained
the m
ost health
y fi sh com
mu
nities as m
easured
by h
igher d
ensities of h
erbivorou
s troph
ic group
s and
all trop
hic grou
ps b
eing relatively w
ell represen
ted. H
owever,
there w
ere strong d
iff erences in
fi sh p
opu
lations am
ong sites
with
in th
e three b
ays, with
the m
ost health
y fi sh com
mu
ni-
ties at Am
bod
ivahib
e South
, Loky Sou
th an
d A
nkao Sou
th
(see Tab
le 1 in
“Rep
ort at a Glan
ce,” on p
age 13 for site
nu
mbers). Th
ese sites also h
ad gen
erally low levels of m
acro an
d tu
rf algae (Ch
apter 6
). W
e observed
the N
apoleon
wrasse C
heilinus u
ndu
latus at
six sites, rangin
g from ju
veniles (on
e 16-2
0 cm
TL
size class, on
e 46-5
0 cm
TL
) to adu
lts (81-8
6 cm
TL
), thou
gh large
termin
al ph
ase males w
ere not seen
. Wh
ile this freq
uen
cy is greater th
an in
man
y other sites su
rveyed in
the W
IO
(Samoilys u
np
ubl.) an
d is h
igher th
an th
e nu
mber ob
served
by A
llen in
2002 (on
e small ad
ult an
d tw
o juven
iles), the
lack of the largest term
inal p
hase m
ales also ind
icates a likely im
pact of fi sh
ing.
Th
rough
out th
e survey, n
ot a single sh
ark, nor th
e large excavatin
g Bolbom
etapon m
uricatu
m (B
um
ph
ead p
arrotfi sh)
were record
ed on
our fi sh
surveys. O
nly tw
o med
ium
-sized
grey reef sharks (C
archarhinus am
blyrhynchos) w
ere observed
in
deep
water off A
mbod
ivahib
e (by D
avid O
bu
ra) wh
ile sn
orkeling p
rior to a dive.
CON
CLUSIO
NS A
ND
CON
SERVATION
RECOM
MEN
DATIO
NS
Th
e diversity of reef fi sh
species m
easured
on th
is relatively rem
ote part of th
e coastline (b
ased on
a diversity in
dex of 1
9
families) w
as 74%
of all species kn
own
to occur in
the w
hole
of Mad
agascar. Certain
families w
ere very poorly rep
re-sen
ted, b
ut oth
ers were w
ell represen
ted an
d 2
3 n
ew sp
ecies record
s for Mad
agascar were ob
tained
. Th
e CF
DI in
dex of
172 w
as similar to th
at measu
red b
y Allen
for North
west
Mad
agascar of 176. C
onsid
ering th
e relatively low relief
reefs of simp
le structu
re (presu
mab
ly du
e to the exp
osure of
the coastlin
e to oceanic seas an
d trad
e win
ds an
d th
e high
tu
rbid
ity), plu
s the lim
ited su
rveys on th
e outer SE
slopes,
this d
iversity is consid
ered to b
e high
and
comp
arable to th
e N
orthw
est coast.Th
e d
eep b
ays of Loky B
ay and
Am
bod
ivahib
e had
the
high
est fi sh d
iversity, particu
larly of the m
ost speciose fam
i-lies of p
omacen
trids an
d lab
rids an
d also th
e chaetod
ontid
s,
Chapter 2
38Rapid Assessm
ent Program
and
this is likely d
ue to greater h
abitat d
iversity in th
ese bays. N
otably, th
e serranid
s, a targeted fi sh
ery group, w
ere relatively d
iverse at the ou
ter (more exp
osed an
d less tu
rbid
) sou
thern
site of Am
bod
ivahib
e. Mu
ltiple-zon
e conservation
areas in
these b
ays off er a good op
portu
nity for m
aximizin
g con
servation of fi sh
species (see also b
elow). Th
e d
iversity of lu
tjanid
s and
lethrin
ids w
as also high
est at Nosy A
nkao.
Th
e absen
ce, or low d
ensities, of h
ighly vu
lnerab
le species
such
as sharks, b
um
ph
ead p
arrotfi sh (B
olbometapon
murica-
tum
) and
Nap
oleon w
rasse (Cheilin
us u
ndu
latus) w
as notice-
able, an
d sp
ecies specifi c m
anagem
ent p
lans are stron
gly recom
men
ded
to facilitate their recovery.
Th
e low ab
un
dan
ce or absen
ce of the larger p
redators su
ch
as sharks, grou
pers (serran
ids) an
d trevally (caran
gids), an
d
also low ab
un
dan
ces of omn
ivores and
herb
ivores is high
ly likely to b
e ind
icative of fi shin
g imp
acts. Th
e latter two tro-
ph
ic group
s may refl ect fi sh
ing by locally d
eployed
gears that
captu
re mixed
species, typ
ical of artisanal fi sh
eries. Th
e pre-
sence of sm
all villages, the region
al centers at V
ohem
ar and
D
iego, and
activity by seasonal m
igrant fi sh
ers are all likely to im
pose m
oderate to h
igh levels of p
ressure even
on th
ese seem
ingly rem
ote locations. Th
e su
bseq
uen
t socio-econom
ic su
rvey to be con
du
cted b
y CI as p
art of this R
AP
shou
ld
provid
e the n
ecessary data to p
roperly d
etermin
e the lin
ks betw
een th
e reef fi sh fau
na an
d local com
mu
nities, m
igrant
fi shers an
d th
e fi shin
g pressu
re they exert, to p
rovide gu
i-d
ance on
village level man
agemen
t plan
nin
g. Th
e presen
ce of a com
mercial algae farm
at Nosy A
nkao, w
hich
discou
-rages fi sh
ing in
the area, w
ith n
oticeably h
igher ab
un
dan
ce of som
e fi shery target sp
ecies (omn
ivores) than
seen in
the
other location
s, sup
ports th
e conclu
sion th
at heavy fi sh
ing
pressu
re elsewh
ere on th
is coastline h
as driven
pop
ulation
d
ensities d
own
. In
terms of con
servation p
lann
ing for en
han
cing th
e resilien
ce of the coral reefs of n
ortheastern
Mad
agascar to clim
ate chan
ge, a mu
lti-zone ap
proach
to protect th
e fi sh
pop
ulation
s is recomm
end
ed. Th
e ab
un
dan
ce surveys su
ggest th
at conservation
eff orts shou
ld focu
s on p
rotecting th
e most
health
y fi sh com
mu
nities w
hich
were fou
nd
at Am
bod
i-vah
ibe Sou
th, L
oky South
and
Nosy A
nkao Sou
th. Th
is
inclu
des th
e most h
ealthy h
erbivore p
opu
lations (T
able 2
.6)
both
in term
s of a) overall abu
nd
ance (A
nkao Sou
th)
and
b) b
road rep
resentation
of troph
ic group
s (grazers/scrap
ers etc.) (Am
bod
ivahib
e South
, Figu
re 2.1
), and
these
sites had
correspon
din
gly low algal cover. Th
ese sites also
had
the h
ighest (th
ough
still low) p
iscivore den
sities. Reef-
associated fi sh
ery species fi sh
ed th
rough
artisanal fi sh
eries are best m
anaged
throu
gh p
erman
ent n
o-take zones in
order to
protect sp
awn
ing stocks, m
aximise rep
rodu
ctive outp
ut an
d
thereb
y facilitate recruitm
ent to n
eighbou
ring reefs (R
uss
2002). It is th
erefore recomm
end
ed th
at no- take zon
es be
establish
ed w
ithin
these th
ree bay location
s. Th
is will p
rotect h
erbivores w
hich
shou
ld en
han
ce recovery of corals, and
also h
elp p
opu
lations of th
e fi shery target sp
ecies (piscivores
and
omn
ivores) recover. Fortu
nately, th
e most d
iverse fi sh
comm
un
ities on th
is stretch of coastlin
e were also fou
nd
in
two of th
ese bays (A
mbod
ivahib
e and
Loky) th
erefore the
no-take zon
es wou
ld also p
rotect the m
ost signifi can
t sites in
terms of fi sh
diversity.
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atic Biology 6
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Table 2.6. Total herbivore abundance (all trophic groups combined). Sites
recomm
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SiteDensity (fi sh/1000m
2)
Am
bod
ivahib
e N96.0
Am
bo
divah
ibe S
19
7.6
Loky N
orth W
est77.6
Lo
ky S
ou
th1
40
.8
An
kao North
132.8
An
kao North
east121.3
An
kao
So
uth
25
0.4
Voh
emar N
orth67.2
Reef fishes of northeast Madagascar
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arine E
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uloch
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olun
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limate W
ar-m
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arine P
rotected A
reas and
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ood. 2
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onitorin
g fun
c-tion
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icators of coral reef resilien
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ide for coral reef
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agers in th
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g grou
p on
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ivien, M
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ily Ep
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and
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olke, R.S. Sten
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ilson, 2
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id M
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iodiversity A
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ulletin
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iological Assessm
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an, T
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d d
iversity of fi sh an
d fi sh
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res and
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agemen
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6.
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ss, G.R
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ishery M
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ools. In: Sale P.F. (ed
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ishes: D
ynam
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a C
omp
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ress. Pp. 4
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43.
Obu
ra, D. 2
004. B
iodiversity Su
rveys of the C
oral Reefs
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azi Bay - R
uvu
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stuary M
arine P
ark. IUC
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ra, D.O
. 2008. Scleractin
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e Western
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ra, D.O
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en years after b
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uen
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ian
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ombasa. K
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47.
Obu
ra, D.O
., M.A
. Samoilys, J. L
utjeh
arms an
d J. H
ermes.
2008. Is th
ere a Western
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ian O
cean “C
oral Tri-
angle”? M
arine Scien
ce for Man
agemen
t/WIO
MSA
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roject docu
men
t. Web
site:http
://ww
w.cord
ioea.org/w
io-core/O
bu
ra, D.O
. and
G.G
rimsd
ith. 2
009. R
esilience A
ssess-m
ent of coral reefs –
Assessm
ent p
rotocol for coral reefs, focu
sing on
coral bleach
ing an
d th
ermal stress. IU
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p on
Clim
ate Ch
ange an
d C
oral Reefs.
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N, G
land
, Switzerlan
d.
Samoilys, M
.A. 1
988. A
bu
nd
ance an
d sp
ecies richn
ess of coral reef fi sh
on th
e Ken
yan coast: th
e eff ects of protec-
tive man
agemen
t and
fi shin
g. Proc. 6
th In
t. Coral R
eef Sym
p. 2: 2
61-2
66.
Samoilys, M
.A. 1
997. U
nd
erwater visu
al censu
s surveys. In
: Sam
oilys, M. A
. (ed.) M
anu
al for assessing fi sh
stocks on
Pacifi c coral reefs. Q
ueen
sland
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artmen
t of Pri-
mary In
du
stries, Brisb
ane. P
p. 16-2
9.
Samoilys, M
.A. an
d G
. Carlos. 2
000. D
etermin
ing m
ethod
s of u
nd
erwater visu
al censu
s for estimatin
g the ab
un
-d
ance of coral reef fi sh
es. En
v. Biol. F
ish. 5
7: 2
89-3
04.
Samoilys, M
.A., I. Silva, J. N
dagala, S. M
uciva an
d D
. M
acharia. In
press. A
rapid
assessmen
t of coral reefs in
Cab
o Delgad
o, north
ern M
ozambiq
ue. C
OR
DIO
R
eport.
Smith
, M.M
. and
P.C. H
eemstra. 2
003. Sm
ith’s Sea Fish
es. Stru
ik Pu
blish
ers.
40Rapid Assessm
ent Program
INTRO
DU
CTION
Th
is stud
y was realized
un
der th
e Rap
id A
ssessmen
t Program
(Marin
e RA
P) of m
arine b
iodi-
versity by C
onservation
Intern
ational M
adagascar. In
total, 26 sites w
ere surveyed
over 20 d
ays, on
the n
ortheast coast of M
adagascar from
Am
bod
ivahib
e Bay to V
ohem
ar. Th
ere are few
pu
blication
s or inven
tories of Ech
inod
erms in
Mad
agascar notab
ly Ch
erbon
nier (1
988), C
her-
bon
nier &
Gu
ille (1978), M
arshall &
Row
e (1981). O
nly h
olothu
rians, w
hich
are extensively
exploited
for the p
reparation
of trepan
g, have b
een stu
died
in d
etail.
Th
e main
objectives of th
e stud
y were to:
• in
ventory th
e echin
oderm
s of north
east Mad
agascar; •
test hyp
otheses ab
out th
e ecological roles of echin
oderm
s related coastal ecosystem
s and
in
particu
lar the coral reef ecosystem
;•
determ
ine th
e existence or d
egree of overexploitation
of sea cucu
mbers as on
e of the m
ain
fi sheries in
Mad
agascar.
METH
OD
S
In th
is survey, on
ly the fou
r most com
mon
classes nam
ely echin
oids, asteroid
s, holoth
uroid
s an
d op
hiu
roids w
ere stud
ied, an
d som
e crinoid
s were also ob
served. T
wen
ty six sites were su
r-veyed
in a total of 3
0 h
ours of ob
servation, b
y walkin
g on th
e reef fl at, snorkelin
g in sh
allow
dep
ths an
d b
y SCU
BA
divin
g (up
to 25 m
eters) on ou
ter reef slopes.
Species id
entifi cation
was p
rimarily m
ade from
in-situ
observation
based
on p
ersonal
experien
ce and
guid
es on ech
inod
erms (G
uille et al. 1
986). M
any sp
ecies are easily iden
tifi ed
by eye, how
ever man
y aspects of extern
al app
earance are variab
le, and
therefore u
nreliab
le for id
entifi cation
for some sp
ecies. Ad
dition
ally, there is often
very similar ap
pearan
ce of two
diff eren
t species. U
nd
erwater p
hotograp
hy w
as used
to docu
men
t and
check on
iden
tifi cations,
and
to avoid collectin
g man
y specim
ens, p
articularly for rare or over-u
tilized sp
ecies such
as h
olothu
rians. O
ne n
ight d
ive was u
nd
ertaken to ch
eck on d
iversity pattern
s at nigh
t, in L
oky B
ay. Specim
ens th
at were collected
were p
reserved in
70° eth
anol, ren
ewed
after several days as
the colou
r from th
e anim
als is leached
into th
e alcohol. In
itial preservation
in d
ilute (b
uff ered
) form
alin (1
0%
in seaw
ater) for several days h
elps to p
reserve colour.
RESULTS
Ech
inod
erms w
ere foun
d across a ran
ge of hab
itats, from reef fl ats an
d seagrass b
eds to d
eep
water on
outer slop
es. Most op
hiu
roids live eith
er in th
e bran
ches of live corals like A
cropora or P
orites, or un
der coral d
ebris or b
locks of dead
coral. Man
y juven
ile sea cucu
mbers are also
un
der coral d
ebris an
d d
ead b
locks sometim
es covered w
ith algae. M
any ech
inod
erms are
Chapter 3
Echinoderms from
the north-east coast of M
adagascar
Jean Maharavo
Echinoderms from
the north-east coast of Madagascar
41 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
active only at n
ight, in
particu
lar sea cucu
mbers, crin
oids
and
gorgonocep
halid
s.In
total, 68 sp
ecies belon
ging to th
e fi ve classes were
recorded
. Th
ese are 3 sp
ecies of crinoid
, 18 h
olothu
rians,
27 op
hiu
roids, 10
echin
oids an
d 10
asteroids. T
wo oth
er
species w
ere not fou
nd
du
ring th
is survey, b
ut w
ere foun
d
by th
e auth
or in A
mbod
ivahib
e in 2
007. Th
is b
rings th
e total n
um
ber of in
ventoried
Ech
inod
erms in
the area to
70 sp
ecies.Th
e m
ost diverse sites w
ere site 10 in
Loky B
ay, site 14 in
V
ohem
ar Bay an
d site 2
in A
mbod
ivahib
e Bay. It sh
ould
be
noted
that th
e large nu
mber of ech
inod
erm sp
ecies recorded
in
Voh
emar B
ay is main
ly du
e to the ab
un
dan
ce of opp
or-tu
nistic sp
ecies that b
enefi t from
the in
pu
t of organic m
atter in
the vicin
ity of the tow
n. A
mon
g these sp
ecies are the
echin
idea T
ripneu
tes gratilla, Diadem
a setosum
, Toxopn
eutes
pileolus; th
e oph
iuroid
s Macrophiotrix cf. ru
gosa, Ophioplocu
s im
bricatus, O
phiocoma erin
aceus, O
. scolopendrin
a and
holo-
thu
rians Syn
apta macu
lata and E
uapta godeff royi.
Acan
thaster planci, a seastar p
redator of h
ard corals, w
as n
ot encou
ntered
. Th
e scarcity of comm
ercial holoth
urian
s su
ch as H
olothuria scabra, Th
elen
ota anan
as, Microthele
fuscogilva an
d Microthele n
obilis was ob
served, an
d M
icrothele fu
scopun
ctata was en
coutered
only on
ce in th
e Loky
Bay. A
single sp
ecimen
of Holothu
ria scabra versicolor was
encou
tered on
ce in th
e An
dravin
a Bay on
mu
dd
y bottom
at 1
2 m
eters in d
epth
. On
ly holoth
urian
species of in
ferior qu
ality, or un
exploited
species, w
ere encou
ntered
. Am
ong
these are H
olothuria atra, H
olothuria verru
cosa, H. leu
cospi-lota, Syn
apta macu
lata, Euapta godeff royi etc.
Figure 3.1. Distribution of echinderms by class, northeast M
adagascar.
Figure 3.2. Number of species of echinoderm
s by site, with the location of the site also indicated.
Ophiuroidea40%
Holothuroidea26%
Echinoidea15%
Crinoidea, 4%
Asteroidea15%
0 2 4 6 8 10 12 14 16 18 20
Ankao
Ambo
Ankao
Andr
Andr
Voh
Loky
Loky
Ankao
Ankao
Voh
Loky
Ambo
Ankao
Loky
Andr
Loky
Loky
Ambo
Loky
Voh
Loky
Ambo
Ambo
Voh
Loky
10 14 2 4 8 16 20 1 7 9 18 21 26 3 12 15 22 23 6 11 13 17 19 25 5 24
# of species per site
Chapter 3
42Rapid Assessm
ent Program
DISCU
SSION
Echinoderm diversity
Seventy sp
ecies of echin
oderm
s are reported
here from
n
ortheast M
adagascar, a low
nu
mber com
pared
to the few
ech
inod
erm in
ventories from
other p
arts of Mad
agascar, and
com
pared
to other p
arts of the w
orld. Th
is n
um
ber com
-p
ares with
125 sp
ecies (Mah
aravo 1990, u
np
ublish
ed) from
T
ulear in
south
west M
adagascar w
here coral reefs are m
uch
m
ore develop
ed geom
orph
ologically, 90 sp
ecies in R
odrigu
es lagoon
(Mau
ritius) an
d 2
40 sp
ecies in N
ew C
aledon
ia (G
uille et al. 1
986).
Several factors may cau
se this low
nu
mber of sp
ecies in
the n
ortheast, su
ch as th
e low d
iversity of reef hab
itats in
the region
, low reef area an
d ecological con
dition
s. Hab
itats occu
pied
by the ech
inod
erms in
clud
ed reef fl ats (seagrass
bed
s, detritu
s reef fl ats, basin
reef, and
comp
act reef fl at) and
ou
ter slopes. B
ut th
e narrow
coastline p
revents th
e develop
-m
ent of large lagoon
al and
island
systems w
ith com
plex reef
topograp
hy, th
us lim
iting d
iversifi cation of h
abitats th
at m
ight favou
r a high
er diversity of ech
inod
erms. Th
e am
oun
t of reef h
abitat in
the area is lim
ited b
y the n
arrow an
d steep
con
tinen
tal coast, restricting reefs to a n
arrow frin
ge along
the coastlin
e. Th
e ph
ysico-ecological cond
ition of th
e reefs w
as relatively severe – w
ith h
igh w
ave energy an
d exp
osure
aff ecting all ou
ter reefs, and
high
turb
idity an
d sed
imen
ta-tion
from ru
noff aff ectin
g all reefs and
especially th
ose in th
e larger b
ay systems. F
inally, th
ough
man
y echin
oderm
s are n
octural, a n
ight d
ive did
not en
coun
ter a large nu
mber of
new
species oth
er than
a basket star (E
uryale aspera).
In q
uan
titative terms, th
e 26 sites su
rveyed ap
pears su
f-fi cien
t to represen
t the en
tire survey area, sh
own
by the
plateau
in sp
ecies nu
mber again
st nu
mber of sam
plin
g sites (fi g. 3
.3). In
fact, no m
ore new
species w
ere discovered
after site 2
1. W
e therefore b
elieve that th
e samp
ling w
as suffi cien
t to get a rep
resentative sam
ple of ech
inod
erms in
the area,
and
the d
iversity of echin
oderm
s here is low
.
Echinoderms as environm
ental indicatorsTh
e d
istribu
tion of sp
ecies in th
e diff eren
t classes of echin
o-d
erms refl ects en
vironm
ental con
dition
s, sum
marized
below
for op
hiu
roids, ech
inoid
s and
holoth
urian
s.Som
e Op
hiu
rids (b
rittle stars) live in d
egraded
areas u
nd
er rocks, or in crevices in
the com
pact reef fl ats. Th
ey
extend
their arm
s out at h
igh tid
e to trap su
spen
ded
solids
on w
hich
they feed
. Am
ong th
ese species associated
with
d
egraded
areas, we fou
nd
Ophiocom
a scolopendrin
a, Ophio-
coma erin
aceus, O
phioplocus im
bricatus an
d O
phiarthrum
elegan
s. Macrophiothrix cf.ru
gosa hid
es ben
eath d
ead b
locks w
hich
mu
st be reversed
to discover it. O
ther op
hu
iroids live
in association
with
over living organ
isms su
ch as b
ranch
ing
corals, antip
atharian
s or spon
ges, and
feed b
y grazing settled
p
articulate m
atter on th
e surface of th
ese organism
s. Th
ese sp
ecies are often associated
with
outer slop
es, and
we fou
nd
O
phiothela tigris and
Ophiom
astix caryophillata belon
g to this
group. O
phionereis porecta seem
s to prefer h
igh san
dy areas
subject to stron
g tidal cu
rrents. F
inally, u
biq
uitou
s species
were fou
nd
wh
ich occu
py d
iff erent typ
es of environ
men
t at th
e reef fl at and
deep
er sedim
ent slop
es or live ben
eath
blocks of d
ead coral.
Five fam
ilies of echin
oids w
ere represen
ted: C
idarid
ae (E
ucidaris m
etularia), T
oxopn
eustid
ae (Tripn
eustes gratilla
and
Toxopn
eustes pileolu
s), Diad
ematid
ae (Diadem
a setosum
, E
chinothrix diadem
a, Echin
othrix calamaris, D
iadema savign
yi an
d Astropyga radiata), Stom
opn
eustid
ae (Stomopn
eustes
variolaris) and
Ech
inom
etridae (E
chinom
etra mathaei).
Echin
ostrephus m
olaris lives in h
oles in coral rock an
d w
as the
most w
idely d
istribu
ted sea u
rchin
, encou
ntered
on 1
2 sites
du
ring th
is exped
ition, an
d is typ
ical of health
y reef environ
-m
ents. M
ost other sp
ecies were n
early or totally absen
t from
un
distu
rbed
sites such
as Loky B
ay, Am
bod
ivahib
e Bay an
d
An
dravin
a Bay.
Voh
emar, b
y contrast, su
pp
orted a large an
d d
iverse ech
inoid
comm
un
ity. Diadem
a setosum
is a susp
ension
feeder,
usin
g its spin
es to trap su
spen
ded
material su
ch as algal
Figure 3.3. Accumulation of species with subsequent sam
ples, northeast Madagascar.
0 10 20 30 40 50 60 70 80
2625
2423
2221
2019
1817
1615
1413
1211
109
87
65
43
21
Number of species
Site #
Echinoderms from
the north-east coast of Madagascar
43 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
frond
s to ingest (R
égis and
Th
omassin
, 1982). C
onseq
uen
tly, it can
often b
e an in
dicator sp
ecies of degrad
ation. It w
as on
ly encou
ntered
at the reef fl at at V
ohem
ar. Echin
ometra
mathaei w
as foun
d in
crevices in th
e comp
act reef fl at in
Voh
emar, th
us con
tribu
ting to b
ioerosion of th
e reef, and
is also in
dicative of reef alteration
by overfi shin
g (McC
lanah
an
and
Mu
thiga, 1
989). T
oxopneu
stes pileolus an
d T
ripneu
stes gratilla w
ere foun
d on
seagrass bed
s and
areas of deb
ris in
Voh
emar. T
ripneu
stes gratilla is an h
erbivore as w
ell as fee-d
ing on
organic w
aste. Toxopn
eustes pileolu
s is similar, b
ut is
a more cryp
tic species th
at covers itself with
deb
ris (Mah
a-ravo, 1
993). It is n
ormal to en
coun
ter these tw
o species on
reef d
ebris an
d seagrass b
eds n
ear town
s wh
ere the in
pu
t of organ
ic matter is h
igh. Th
is ech
inoid
pop
ulation
at Voh
emar
is an in
dicator of en
vironm
ent d
egradation
. By con
trast, the
near-ab
sence of th
ese species from
the oth
er locations (L
oky B
ay, Am
bod
ivahib
e Bay an
d A
nd
ravina B
ay), testifi es to th
eir ecological health
. M
any ech
inoid
s comm
on to oth
er parts of M
adagsacar,
particu
larly the w
est coast, were ab
sent h
ere. Th
is inclu
des
the great C
idarid
ae (Prion
ocidaris and
Phylacan
thus),
Oreasterid
ae (Protoresater, P
entaceraster an
d C
horiaster), the
Lu
idid
ae and
Astrop
ectenid
ae. Colobocen
trotus atratu
s, an
urch
in typ
ically foun
d in
very high
energy con
dition
s was
also not fou
nd
. Th
ese species are also n
ot comm
ercially exp
loited, an
d th
e reefs otherw
ise app
ear to be in
an u
nd
is-tu
rbed
cond
ition. It is n
ot clear wh
y they are n
ot presen
t on
the n
ortheast coast.
Variou
s species of exp
loited an
d n
on exp
loited h
olothu
-rian
s were record
ed. Th
e m
ost comm
on w
as Holothu
ria atra, at fi ve sites. Th
is sp
ecies is very comm
on in
the w
estern
Ind
ian O
cean, occu
ring b
etween
the su
rface and
30m
dep
th,
and
seems to p
refer substrates of coral san
d w
ith or w
ithou
t seagrass. Its b
ody is alw
ays covered w
ith th
in layer of san
d,
bu
t once th
is layer is rubbed
with
fi ngers, it gives off a red
-d
ish color. Th
e m
ost distin
ctive fi nd
ing of th
is inven
tory was
the extrem
e scarcity of holoth
urian
s, especially th
e target sp
ecies of exploitation
: Holothu
ria scabra, Microthele n
obilis, M
icrothele fuscogilva, an
d Th
elen
ota anan
as. In m
ost cases, on
ly one sp
ecimen
of these exp
loited sp
ecies was seen
.Th
is scarcity is p
robab
ly the resu
lt of overexploitation
of sea cu
cum
bers, as in
other region
s of Mad
agascar, for the
prep
aration of trep
ang (b
êche-d
e-mer) for exp
ort, main
ly to A
sia. Du
ring th
e survey, en
qu
iries in th
e villages and
discu
s-sion
s with
fi sherm
en con
fi rmed
the in
tensive exp
loitation
of holoth
urian
s. For tw
enty years n
ow, sea cu
cum
bers h
ave been
extensively exp
loited alon
g every coast of Mad
agascar resu
lting in
the q
uasi-ab
sence of target sp
ecies in th
e dep
th
range from
0 to 2
0 m
eters.
CON
CLUSIO
N
Th
is survey id
entifi ed
70 sp
ecies of echin
oderm
s from th
e n
orth-east coast of M
adagascar, fi n
din
g lower levels of sp
e-cies d
iversity than
the w
est coast. Un
usu
ally for Mad
agascar,
there w
ere no large sp
ecies of seastars (asteroids) from
the
Oreasterid
ae family, su
ch as P
rotoreaster or Pen
taceraster, nor
urch
ins (ech
inoid
s) from th
e family C
idarid
ae, such
as Prio-
nocidaris, P
hylacanthu
s, Heterocen
trotus etc. Th
e ab
sence of
the coral p
redator starfi sh
, Acan
thaster planci, w
as also noted
.In
general, th
ere was a scarcity of ech
inoid
s and
asteroids,
a scarcity that seem
s abn
ormal given
that th
e reefs are fairly w
ell develop
ed. It w
ould
be in
teresting to stu
dy th
e relation-
ship
betw
een th
e distrib
ution
of larvae and
the cu
rrents of
the n
ortheast coast in
order to exp
lain th
is situation
. Over-
fi shin
g of holoth
urian
s, already ob
served in
other p
arts of M
adagascar, w
as confi rm
ed d
urin
g this exp
edition
. Ind
eed,
there is an
extreme scarcity of target sp
ecies. Th
is result
emp
hasizes th
e need
for fi sheries m
anagem
ent an
d th
e need
for regen
eration of h
olothu
rian stocks.
Ech
inod
erm sp
ecies are also ind
icators of environ
men
-tal con
dition
. Ind
icator species of d
egradation
inclu
ded
D
iadema setosu
m, T
ripneu
stes gratilla and
Echin
ometra
mathaei, w
hich
were on
ly encou
ntered
in th
e imm
ediate
vicinity of th
e town
and
port of V
ohem
ar. Th
is is ind
ica-tive of th
e signifi can
t reef pollu
tion from
port activities an
d
organic p
ollution
related to th
e proxim
ity of the tow
n.
REFERENCES
Ch
erbon
nier, G
. 1988. E
chin
oderm
es : Holoth
urid
es. Faun
e d
e Mad
agascar 70.
Ch
erbon
nier, G
. and
A. G
uille. 1
978. E
chin
oderm
es : O
ph
iurid
es. Faun
e de M
adagascar 4
8.
Gu
ille, A., P. L
abou
te and
J.L. M
enou
. 1986. G
uid
e des
étoiles de m
er, oursin
s et autres E
chin
oderm
es de
Nou
velle Caléd
onie. C
ollection fau
ne trop
icale n°X
XV
. O
RST
OM
. Paris. F
rance
Mah
aravo, J. 1993.-E
tud
e de l’ou
rsin com
estible T
ripn
eus-
tes gratilla (L.1
758) d
ans la région
de N
osy-Be (côte
nord
-ouest d
e Mad
agascar): den
sité, morp
hom
étrie, n
utrition
, croissance, p
rocessus rep
rodu
cteurs, im
pact
de l’exp
loitation su
r les pop
ulation
s. Ph
.D Th
esis. U
niv.
Aix-M
arseille III. Fran
ce.M
arshall, J.I. an
d F.W
.E. R
owe. 1
981. Th
e crin
oids of
Mad
agascar. Bu
ll. Mu
s. Nat. Ser. 4
, 3A
(2):3
79-4
13.
McC
lanah
an, T
.R. an
d N
.A. M
uth
iga. 1989. P
atterns of
pred
ation on
a sea urch
in, E
chin
ometra m
athaei (d
e B
lainville), on
Ken
yan coral reefs. J. E
xp. Mar. B
iol. E
col: 1-1
8.
O’L
ough
lin, P.M
. and
F.W.E
. Row
e. 2006. A
systematic
revision of th
e asterinid
genu
s Aqu
ilonastra O
.Lou
ghlin
, 2004 (E
chin
oderm
ata: Asteroid
ea). Mem
. Mu
s. Victoria
63 (2
): 257-2
87.
Régis, M
.B. an
d B
.A. Th
om
assin. 1
982. E
cologie des E
chi-
noid
es réguliers d
ans les récifs corallien
s de T
oliara (S.O.
of Mad
agascar). Ad
aptation
de la m
icrostructu
re des
piq
uan
ts. An
n. In
st. Océan
ogr. 58 (2
): 117-1
58.
44Rapid Assessm
ent Program
SUM
MA
RY
Th
e north
-eastern coast of M
adagascar p
resents exten
sive and
high
ly diverse algal an
d seagrass
assemblages. A
s this p
art of the coastlin
e is almost com
pletely u
nd
eveloped
, these h
abitats are
in a p
ristine state, on
ly limited
by en
vironm
ental con
dition
s, such
as wave action
, sedim
ent
run
off and
nu
trient con
centration
s. Ten
species of seagrass w
ere comm
on, com
parab
le to th
e species d
iversity foun
d elsew
here in
the W
estern In
dian
Ocean
: Th alassoden
dron ciliatu
m,
Th alassia hem
prichii, Syringodiu
m isoetifoliu
m, C
ymodocea rotu
ndata C
. serrulata, H
alodule
un
inervis, H
. wrightii, H
alophila ovalis, H. stipu
lacea and Z
ostera capensis. L
arger species
(e.g. T. ciliatu
m an
d T
. hemprichii) w
ere main
ly foun
d on
stable su
bstrates, m
ostly in coastal
lagoons or on
the sh
allow, in
ner ed
ge of coral reef fl ats. In areas w
here sed
imen
t cond
itions
were p
articularly d
ynam
ic and
hyd
rodyn
amic forces p
lay a major role, sm
aller, fast growin
g sp
ecies (e.g. H. u
nin
ervis) were d
omin
ant, w
ith con
siderab
le spatial an
d tem
poral variation
. In
the few
urb
an areas p
resent alon
g this coastlin
e, seagrass distrib
ution
was lim
ited b
y pollu
-tion
and
eutrop
hication
resultin
g from sew
age outfalls an
d farm
ing activities.
To cap
ture th
e diversity an
d ab
un
dan
ce of algae and
seagrass bed
s in th
e stud
y area, four
hab
itat categories are describ
ed h
ere: coastal, inclu
din
g both
exposed
areas and
lagoons;
reef fl ats; riverine system
s and
deep
/open
water. Th
e list of sp
ecies, frequ
ency of occu
rrence
and
shoot d
ensity (th
e latter for T. hem
prichii and
T. ciliatu
m) are rep
orted for each
hab
itat, in
clud
ing key ecological p
rocesses such
as sedim
ent typ
e, ph
ysical distu
rban
ce, light lim
itation
and
hu
man
-ind
uced
threats. Th
e u
nd
erstand
ing of d
iff erences in
the h
abitats, th
eir key eco-logical fu
nction
s and
poten
tial threats d
escribed
here p
rovide recom
men
dation
s for adap
tively m
anagin
g algal bed
s and
seagrass mead
ows an
d th
eir ecosystem services.
INTRO
DU
CTION
Seagrasses are fl owerin
g plan
ts that h
ave adap
ted to exist fu
lly subm
ersed in
the sea. A
lthou
gh
they are d
istribu
ted glob
ally, along tem
peratu
re and
tropical coastlin
e of the w
orld, th
ey dis-
play h
ighest d
iversity of species in
the trop
ical Ind
o-Pacifi c region
with
24 sp
ecies presen
t(Sh
ort et al., 2008). Seagrasses form
extensive m
eadow
s that su
pp
ort biologically d
iverse ecosystem
s and
play key ecological fu
nction
s in th
e coastal marin
e environ
men
t (Orth
et al., 2006). Seagrasses are typ
ically foun
d in
shallow
coastal areas, conn
ecting estu
arine, m
angrove
and
coral reef hab
itats, hen
ce the ecological services th
ey provid
e are consid
ered to h
ave a h
igh glob
al value (C
ostanza et al., 1
997). In
deep
er water (i.e. on
the ou
ter edge of coral reefs),
seagrasses tend
to be ligh
t limited
with
only som
e species b
eing fou
nd
. Seagrasses colonize
coastal and
riverine lagoon
s and
often act as a b
uff er zon
e for nu
trient recyclin
g, imp
roving
water q
uality, alterin
g water fl ow
and
stabilizin
g sedim
ents, w
ith con
sequ
ent b
enefi cial eff ects
for nearb
y coral hab
itats and
min
imizin
g beach
erosion an
d fl ood
s. In ad
dition
, seagrass bed
s p
rovide h
abitats for a w
ide variety of p
lants an
d an
imals, in
clud
ing food
for micro (i.e. sea
Chapter 4
Seagrasses and algae of north-eastern M
adagascar
Giuseppe Di Carlo & M
onica Tombolahy
Seagrasses and algae of north-eastern Madagascar
45 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
urch
ins) an
d m
acrograzers (sea turtles an
d d
ugon
gs) as well
as nu
rsery and
feedin
g groun
ds for a ran
ge of anim
als and
com
mercially im
portan
t fi sh sp
ecies. A
lgae are comm
only fou
nd
on rocky sh
ores, amon
g m
angrove roots, seagrass b
eds, coral reefs, on
mu
d fl ats
and
coastal lagoons. A
ll three m
ajor algal group
s, the
Rh
odop
hyta or red
algae, Ch
loroph
yta or green algae, an
d
Ph
eoph
yta or brow
n algae, are com
mon
in th
ese hab
itats. Th
ey p
lay a crucial role in
main
tainin
g ecological balan
ce of th
e aqu
atic environ
men
t. Algae are p
rimary p
rodu
cers wh
ich
provid
e imp
ortant food
sources for a d
iversity of marin
e w
ildlife. A
s a result of p
hotosyn
thesis, algae also release oxy-
gen in
to the w
ater colum
n, b
enefi tin
g man
y fi sh, m
ollusks,
crabs an
d oth
er marin
e species. Th
erefore, algae, togeth
er w
ith seagrasses form
the b
ase of the m
arine food
web
that
sup
ports th
e life of herb
ivores and
prim
ary consu
mers.
With
in coral h
abitats, algae,su
ch as th
e green alga H
alimeda
and
red corallin
e algae (i.e. Am
phiroa), contrib
ute to reef
formation
and
develop
men
t throu
gh th
e secretion of a cal-
cium
carbon
ate skeleton. In
Mad
agascar, 20 sp
ecies of algae in
clud
ing eigh
t species of red
algae, 10 sp
ecies of brow
n an
d
green algae are com
mercially exp
loited (w
ww
.meeft.gov.m
g). Th
e m
ost comm
ercially exploited
are the red
algae Eucheu
ma,
Gelidiu
m an
d G
racilaria.Th
e n
orth-eastern
coast of Mad
agascar presen
ts several d
iverse assemblages of algae an
d seagrass h
abitats, ran
ging
from p
atchy seagrass b
eds to m
eadow
s extend
ing over several
kilometers. Th
ose m
eadow
s harb
our an
incred
ibly h
igh
diversity of associated
macroalgae an
d ep
iph
ytes as well as
a diversity of b
enth
ic inverteb
rates (i.e. sea cucu
mber an
d
sea urch
ins) h
ence th
ey represen
t areas of high
biod
iversity im
portan
ce. In ad
dition
, prim
ary prod
uction
and
hab
itat con
struction
in seagrass b
eds m
ake them
imp
ortant n
ur-
sery areas for shrim
p, lobster, an
d several sp
ecies of fi sh (i.e.
siganid
s and
scarids) th
at later live in coral reef h
abitats.
Seagrasses are also imp
ortant for th
e main
tenan
ce of sea tu
rtles (Chelon
ia mydas), an
d h
istorically they su
pp
orted
du
gong p
opu
lations (D
ugon
g dugon
g). Seagrasses and
algae are a m
ajor comp
onen
t of the rich
and
prod
uctive coastal
and
marin
e ecosystem of E
ast Africa (ref).
Here w
e report on
the algae an
d seagrass assem
blages
foun
d alon
g the n
ortheastern
coastline of M
adagascar,
inclu
din
g comm
un
ity comp
osition, d
istribu
tion an
d sp
ecies d
iversity. In ad
dition
, we attem
pt to cap
ture th
e key features
of these assem
blages, com
mu
nity com
position
, inclu
din
g ecological an
d p
hysical p
rocesses and
threats to th
ese ecosystem
s.
METH
OD
S
Five location
s were sam
pled
betw
een th
e town
of Diego
Suarez an
d V
ohem
ar, nam
ely Am
bod
ivahib
e Bay, L
oky Bay,
Voh
emar, A
nd
ravina B
ay and
Nosy A
nkao (T
able 4
.4). Th
e
hab
itats foun
d w
ithin
these location
s were classifi ed
into
four b
asic types:
Riverin
e seagrass/algae hab
itats were fou
nd
in sh
allow,
intertid
al areas, in th
e proxim
ity of large freshw
ater inp
uts,
wh
ere sedim
ents w
ere mostly m
ud
dy san
d to m
ud
.
Coastal h
abitats w
ere iden
tifi ed as in
tertidal to su
btid
al lagoon
s located b
etween
the sh
oreline an
d th
e reef fl ats, with
a m
ixture of con
solidated
(sand
) and
un
consolid
ated (coral
rubble) su
bstrates an
d w
ith a m
ax dep
th of 2
-3m
.
Reef fl ats gen
erally covered th
e entire exten
t of the reef,
with
a mixtu
re of sand
and
coral rubble an
d w
ith seagrass
and
algae growin
g in th
e proxim
ity of coral. Reef fl ats are
exposed
at low tid
e, hen
ce seagrasses are mostly fou
nd
in
intertid
al pools.
Deep/open
water seagrass an
d algae h
abitats w
ere foun
d
at the ou
ter edge of th
e reef below
a dep
th of 8
m, w
ith
Halophila sp
p b
eing th
e only seagrass sp
ecies foun
d in
these
hab
itats.
Algae an
d seagrass h
abitats, com
mu
nity com
position
an
d ab
un
dan
ce were assessed
usin
g a rapid
, visual assess-
men
t techn
iqu
e know
n as th
e Brau
n-B
lanqu
et meth
od
(Brau
n-B
lanqu
et, 1972). Th
is m
ethod
is qu
ick, requ
iring
only m
inu
tes at each sam
plin
g site, bu
t at the sam
e time is
consid
ered rob
ust an
d h
ighly rep
eatable, th
ereby m
inim
i-zin
g amon
g observer d
iff erences. A
t each of th
e fi ve locations
samp
led, a n
um
ber of station
s were ch
osen to rep
resent th
e h
abitat d
iff erences an
d sp
atial variability of th
e area. At each
station
, three rep
licate 50m
-long tran
sects were estab
lished
at ran
dom
GP
S poin
ts. For each
transect, a m
eter tape w
as exten
ded
along th
e bottom
and
secured
with
metal rod
s. Ten
qu
adrats (0
.25m
2) were th
en p
laced alon
g the lin
e at pred
e-term
ined
rand
om p
ositions, re-calcu
lated for each
transect.
Each
qu
adrat w
as examin
ed u
sing SC
UB
A, listin
g all algae
Table 4.1. The Braun-Blanquet abundance score. Each algae and seagrass species was scored in each quadrat according to this scale.
Cover ClassDescription
0Sp
ecies absen
t
0.1
Species fou
nd
in solitary sh
oots, < 5
% cover
0.5
Species fou
nd
in few
shoots, <
5%
cover
1Sp
ecies foun
d in
man
y shoots, <
5%
cover
2Sp
ecies foun
d in
5%
- 25%
cover
3Sp
ecies foun
d in
25%
- 50%
cover
4Sp
ecies foun
d in
50%
- 75%
cover
5Sp
ecies foun
d in
75%
- 100%
cover
Chapter 4
46Rapid Assessm
ent Program
and
seagrass species p
resent an
d scorin
g their p
ercent cover
based
on th
e fraction of th
e total area of the q
uad
rat that
was ob
scured
by a particu
lar species (T
able 4
.1). A
dd
itio-n
ally, shoots of large, clim
ax seagrass species w
ere coun
ted to
establish
their d
ensity. O
ther featu
res (ie fl owers, seed
s, etc), n
ew sp
ecies or anom
alies were n
oted as w
ell as sedim
ent
type an
d evid
ence of grazin
g and
canop
y heigh
t.A
t each site, key p
rocesses and
threats w
ere noted
such
as fi sh
ing activities, h
igh sed
imen
t dep
osition, p
resence of
river systems, p
ollution
and
/or sewage ou
tfalls, etc. Wh
ere taxon
omic d
oubts existed
(i.e. for Halodu
le wrightii), sp
e-cies w
ere ph
otograph
ed for fu
rther id
entifi cation
in th
e lab
oratory.
RESULTS A
ND
DISCU
SSION
Algae
Nin
ety one sp
ecies of algae were record
ed in
the su
rveys, in
the th
ree major taxon
omic grou
ps R
hod
oph
yta or red
algae, Ch
loroph
yta or green algae, an
d P
heop
hyta or b
rown
algae.Th
e largest grou
p w
as the red
algae with
44 sp
ecies in
12 fam
ilies, followed
by 32 sp
ecies of green algae in
10 fam
i-lies an
d b
rown
algae were least rep
resented
with
11 sp
ecies in
2 fam
ilies (Dictyotacea an
d Sargassacea) (fi g.4
.1). A
ll group
s w
ere comm
on in
all the fou
r hab
itat types.
Th
e dom
inan
t green algae fam
ilies were V
aloniacae,
Ud
oteacae and
Cau
lerpacae (T
able 4
.2), w
hile w
ithin
the red
algae th
e dom
inan
t families w
ere Rh
odom
elacae and
Hyp
-n
eacae. Th
e calcareous red
algae Coralin
acae and
Galaxau
ra-cae w
ere in ab
un
dan
ce in A
nd
ravina an
d V
ohem
ar.Som
e algal species are on
ly foun
d season
ally at the loca-
tions sam
pled
, or in d
iff erent p
hases of th
eir life cycle. For
examp
le, some green
algae species in
An
dravin
a and
Loky
bay w
ere encou
ntered
at low ab
un
dan
ce in th
eir growin
g
ph
ase (e.g. Acetabu
laria pavula, M
icrodyction aghardian
um
), an
d sim
ilarly for the red
algae such
as Gelidiela acerosa,
Lau
rencia sp.
Algae sp
ecies that are in
dicative of ecological im
balan
ce w
ere recorded
at several locations, in
particu
lar Tydem
ania
expeditionis an
d M
icrodyctionsp in
Am
bod
ivahib
e. Cau
lerpa racem
osa and
C. brachypu
s were ab
un
dan
t in N
osy An
kao d
ue to th
e seaweed
farm p
resent at th
is location. A
t Nosy
An
kao and
Am
bod
ivahib
e blu
e green algae (i.e. C
yanop
hy-
cees, Lyn
gbia sp.) were record
ed coverin
g a large prop
ortion
of corals and
spon
ges. D
iff erent alga sp
ecies were fou
nd
across the fi ve loca-
tions, refl ectin
g local ecological cond
itions. C
alcareous
Figure 4.1. Distribution in percentage of algal groups across the fi ve locations.
Table 4.2. Number of species of algae in each Fam
ily, by survey location (AM
BDV = Am
bodivahibe Bay)
AMBDV
LOKYANKAO
ANDRAVINA VOHEMAR
CHLOROPHYTA
Ulvacae
11
11
2
Clad
oph
oracae0
00
02
Siph
onoclad
acae1
00
12
An
adyom
enacae
10
11
0
Valon
iacae4
02
05
Cau
lerpacae
36
56
1
Ud
oteacae6
65
43
Cau
diacae
01
11
0
Dasyclad
acae1
01
01
Polyp
hisacae
10
00
0
TO
TA
L1
81
41
61
41
6
RHODOPHYTA
Rh
odom
elacae5
01
411
Delesseriacae
11
10
0
Ch
amp
iacae0
01
12
Soleriacae0
00
03
Schizym
eniacae
10
00
1
Hyp
neacae
42
11
4
Corallin
acae0
22
55
Peysson
neliacae
02
12
0
Halym
eniacae
10
21
2
Gracillariacae
10
01
3
Gelid
iacae1
10
01
Galaxau
racae0
30
50
TO
TA
L1
41
19
20
32
PHEOPHYTA
Dictyotacae
43
45
2
Sargassacae0
30
00
Sub
total4
64
52
TO
TA
L3
63
12
93
95
0
Phaeo
phyta, 13%
Chlo
rop
hyta, 37%
Rho
do
phyta, 50%
Seagrasses and algae of north-eastern Madagascar
47 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Table 4.3. Summ
ary of algal comm
unity in the four main habitats (riverine, coastal, reef fl at and deep reefs), and the locations in which these were found.
LKB (Loky Bay), VOHM (Vohem
ar), ANDR (Andravina Bay), AMBD (Am
bodivahibe Bay). In each habitat, algal species are listed by their prime habitat, with
blanks in the substrate column indicating the sam
e substrate as the species above.
Habitat%
speciesKey algae species
SubstrateLocation occurrence
Riverin
e7
Avrain
villea erectaH
ypnea sp1
Hypn
ea cornuata
mu
dL
KB
, VO
HM
, AN
DR
Coastal
36
Cladophora sp
.coral ru
bble/h
ard b
ottoms
AM
BD
, LK
B
Chaetom
orpha indica
Cau
lerpa sertularoides
En
teromorpha com
pressaN
eomeris van
bosseaeD
ictyopteris versluysii
sand
/coral rubble
Cham
pia sp.
Chon
drophycus papillosu
sG
racilaria salicornia
Acan
thophora spicifera H
ypnea corn
uata
Tolypiocladia glom
erulata
Dictyota dichotom
aPadin
a gymn
ospora Padin
a boryana
Avrain
villea erecta
sand
/coral rubble
Reef fl at
30
Cladophora sp
.D
ictyosphaeria cavernosa
Ulva lactu
caE
nterom
orpha compressa
Sarconem
a fi liforme
Digen
ia simplex
Acan
thophora spiciferaG
racilaria salicornia
Gracilaria acru
ataJan
ia adhaerens
coral rubble/h
ard b
ottoms
VO
HM
Halim
eda opun
tiasan
d
Boergen
senia forbesii
sand
/coral rubble
Ven
tricaria ventricosa
sand
/mu
dd
y sand
Deep
/open
water
27
Neu
rymen
ia fraxinifolia
Peysson
nelia capen
sisP
eyssonn
elia simulan
s
coralV
OH
M, A
ND
R
Udotea in
dicaC
aulerpa serru
latasan
d
Halim
eda opun
tiaA
man
cia rhodanta
Galaxau
ra fasciculata
Galaxau
ra elongata
sand
/coral
Galaxau
ra sp2.
Coralin
a sp.
Halim
enia du
rvillei
sand
/coral
sedim
ent-form
ing algae (e.g. H
alimeda m
acroloba) are tole-ran
t of low ligh
t levels and
were com
mon
ly foun
d at d
epth
s greater th
an 2
5 m
(Wein
berg, 1
976). A
vrainvillea erecta
and
some red
algae such
as Hypn
ea sp. and
H. cu
neata w
ere com
mon
in sh
allow an
d in
tertidal seagrass b
eds aff ected
by
terrigenou
s run
off and
sedim
entation
, wh
ere high
levels of an
oxia on th
e mu
dd
y substratu
m occu
r.Seaw
eed p
opu
lations on
intertid
al rocky shores ten
d
to be m
ore prod
uctive an
d exten
sive than
those on
sand
y beach
es or mu
d fl ats (B
ranch
et al., 1994). Som
e areas p
resented
abu
nd
ant sp
ecies in each
hab
itat. In p
articular,
Chapter 4
48Rapid Assessm
ent Program
in A
mbod
ivahib
e Bay, w
e noted
thrivin
g red algae A
can-
thophora spicifera and
the b
rown
algae Dictyota dichotom
a. Th
e fi rst sp
ecies could
likely be an
invasive sp
ecies that
becam
e abu
nd
ant as en
vironm
ental factors are favorab
le (e.g h
um
an-in
du
ced organ
ic matter in
crease). In
areas frequ
ently exp
osed to d
isturb
ances, op
portu
nistic
species, su
ch as U
lva lactuca an
d E
nterom
orpha compressa
were fou
nd
. A few
rare species w
ere foun
d at h
igh ab
un
-d
ance at in
divid
ual sites, som
etimes in
dicative of d
istur-
ban
ce or invasion
, such
as Cau
lerpa and
Halim
eda spp. A
n
un
know
n sp
ecies of blu
e-green algae w
as foun
d at m
ost sites, bu
t was p
articularly ab
un
dan
t at Nosy A
nkao. Th
is fi n
din
g, togeth
er with
the p
resence of C
aulerpa at th
is site, suggests
enrich
men
t in n
utrien
ts from th
e seaweed
farm (K
raufvelin
et al. 2
010). Sp
ecies of Udotea w
ere comm
on, an
d alw
ays fou
nd
small p
atches in
the san
d p
atches b
etween
reefs, m
ostly at An
kao, Loky b
ay, and
An
dravin
a.C
oastal hab
itats had
the h
ighest n
um
ber of algal sp
ecies (3
6%
, Tab
le 4.3
), dom
inated
by green
algae, in som
e cases by sp
ecies ind
icative of eutrop
hic in
pu
t (e.g. En
teromorpha
compressa an
d U
lva lactuca). Th
ese w
ere followed
in im
por-
tance b
y brow
n algae, su
ch as P
adina gym
nospora an
d P. bory-
ana (m
ore abu
nd
ant in
the L
oky Bay). W
ithin
seagrass bed
s, p
articularly in
Am
bod
ivahib
e bay, A
nd
ravina b
ay and
Loky
Bay, D
ictyota dichotoma w
as abu
nd
ant. O
n reef fl ats, w
here
aerial exposu
re is comm
on at low
tide (esp
ecially at Voh
e-m
ar) the red
algae Gracilaria salicorn
ia and
Dygen
ia simplex
were ab
un
dan
t. In sh
allow p
ools on th
e reef fl ats that retain
w
ater du
ring low
tide, toleran
t species su
ch as G
racilaria acru
ata, Acan
thophora spicifera and
Boergen
senia forbesii w
ere com
mon
. Fin
ally, deep
er areas (below
2 m
) were occu
pied
by a d
iversity of larger red algae.
SeagrassesT
en seagrass sp
ecies were com
mon
, with
the h
ighest n
um
ber
of species (7
) in riverin
e hab
itats, followed
by th
e shallow
p
latform of reef fl ats an
d coastal areas (6
species each
, both
exp
osed an
d sh
eltered) an
d low
est in d
eep h
abitats (2
spe-
cies) (Tab
le 4.4
). Alth
ough
seagrass bed
s were n
ot map
ped
, ou
r observation
s suggest th
at seagrasses form exten
sive m
eadow
s along th
is coastline coverin
g in som
e areas (i.e. L
oky Bay an
d V
ohem
ar) tens of kilom
eters of coastline,
generally on
shallow
sheltered
coastal lagoons. A
nn
ual b
eds
formed
mostly b
y smaller, fast-grow
ing sp
ecies (i.e. Halodu
le un
inervis, H
alophila ovalis) were often
foun
d in
exposed
and
tran
sient en
vironm
ents, su
ch as sh
allow san
d b
anks an
d reef
fl ats (Tab
le 4.5
). Th
ose mead
ows are largely aff ected
by sed
i-m
ent m
ovemen
t du
ring storm
s and
cyclones.
Alon
g this coastlin
e, seagrasses typically form
mixed
sp
ecies assemblages th
ough
with
dom
inan
ce by on
e species,
controlled
prin
cipally b
y dep
th (su
btid
al vs intertid
al) and
sed
imen
t cond
itions (T
able 4
.5). In
the in
tertidal, Z
ostera capen
sis, H. u
nin
ervis and
Cym
odocea rotun
data were com
-m
on. Z
. capensis w
as most com
mon
ly presen
t in exp
osed
areas, wh
ile H. u
nin
ervis and
C. rotu
ndata colon
ise tidal
pools on
mu
dfl ats (e.g. in
Voh
emar, T
able 4
.4). In
the
subtid
al, T. hem
prichii and
S. isoetifolium
formed
extensive
mead
ows on
sand
or mu
dd
y sand
(e.g. in V
ohem
ar, Am
bod
i-vah
ibe B
ay), largely infl u
enced
by sed
imen
t comp
osition. A
ll sp
ecies were associated
with
mu
d, san
d, m
ud
dy san
d, an
d
coarse sand
substrates, w
hile on
ly T. hem
prichii, S. isoetifo-liu
m an
d Th
alassoden
dron ciliatu
m colon
ised ru
bble su
bstrate
wh
ere the sed
imen
t layer was m
inim
al (Am
bod
ivahib
e Bay
and
Voh
emar). A
t mid
dep
ths on
the ed
ges of reefs (8-1
2 m
), or am
ong coral p
atches on
sand
, Halophila ovalis form
ed
extensive p
atches often
mixed
with
H. stipu
lacea. No sea-
grasses were fou
nd
below
12 m
on th
e deep
er reefs.F
igures 4
.2 an
d 4
.3 sh
ow th
e relative abu
nd
ance an
d sh
oot d
ensity p
arameters, resp
ectively, for seagrass species at th
e fi ve location
s. Com
parin
g the fi ve location
s samp
led, L
oky B
ay had
the h
ighest d
iversity of species (9
species) follow
ed
by N
osy An
kao (8 sp
ecies) (Fig. 4
.2). Th
is w
as du
e to their
diversity of h
abitats, from
reef fl ats to riverine system
s. In
Loky B
ay, T. ciliatu
m w
as comm
only fou
nd
on coral ru
bble
with
the h
ighest cover an
d sh
oot den
sity (mean
260±95
and
212.5
±89 sh
oot m-2 for coastal lagoon
and
reef fl at resp
ectively, Fig. 4
.3). O
n san
d an
d m
ud
dy san
d, seagrasses
formed
mixed
comm
un
ities of T. hem
prichii, C. rotu
ndata
and
H. u
nin
ervis. In th
is area, T. hem
prichii shoot d
ensity
was th
e lowest record
ed, ran
ging from
14.2
±40.1
in th
e coastal lagoon
s to 31.2
±60.4
shoot m
-2 on th
e reef fl at and
zero in
riverine h
abitats. C
. serrulata an
d Z
. capensis occu
rred
in sh
allow, sh
eltered lagoon
s adjacen
t to river deltas or fresh
-w
ater run
off s.
Table 4.4. Seagrass species recorded at each habitat, with site occurrence. LKB (Loky Bay), VOHM (Vohem
ar), ANDR (Andravina Bay), AMBD (Am
bodivahibe Bay).
Habitat
T. hemprichii
T. ciliatum
C. serrulata
C. rotundata
H. ovalis
H. stipulacea
Z. capensis
H. uninervis
H. wrightii
S. isoetifolium
Site occurrence
Riverin
ex
xx
xx
xx
LK
B, V
OH
M, A
ND
R
Coastal
xx
xx
xx
AM
BD
, LK
B, L
KB
Reef fl at
xx
xx
xx
VO
HM
, LK
B
Deep
xx
LK
B A
ND
R
Seagrasses and algae of north-eastern Madagascar
49 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Figure 4.2. Frequency of occurrence according to the Braun-Blanquet assessment of seagrass percent cover at the fi ve locations in northeast M
adagascar: Am
bodivahibe Bay, Loky Bay, Vohemar, Andravina Bay and Nosy Ankao. The legend for Braun-Blanquet scores is shown in the upper fi gure (and see Table 1).
Greater dominance by a single species is shown by a larger proportion of the higher density classes (ie. shorter length of the dark black, low-density class).
Chapter 4
50Rapid Assessm
ent Program
Table 4.5. Summ
ary of the seagrass comm
unity in the three main habitats (riverine, coastal, reef fl at) in which seagrasses were found.
HabitatLim
iting factorsSeagrass species
SubstrateR
iverine
Terrigen
ous ru
noff
Cym
odocea serrulata
mu
d
salinity
Cym
odocea rotun
datam
ud
sedim
ent an
oxiaH
alophila ovalism
ud
biotu
rbation
Halophila stipu
laceam
ud
Zostera capen
sism
ud
Halodu
le un
inervis
mu
d
Halodu
le wrightii
mu
d
Coastal
ph
ysical distu
rban
ceTh alassia hem
prichiisan
d/m
ud
dy san
d
biotu
rbation
Th alassoden
dron ciliatu
mcoral ru
bble/h
ard b
ottoms
Syringodiu
m isoetifoliu
msan
d/coral ru
bble
Cym
odocea rotun
datasan
d/m
ud
dy san
d
Halophila ovalis
sand
/mu
dd
y sand
Halophila stipu
laceasan
d/m
ud
dy san
d
Reef fl at
ph
ysical distu
rban
ceTh alassia hem
prichiisan
d/m
ud
dy san
d
Th alassoden
dron ciliatu
mcoral ru
bble/h
ard b
ottoms
Syringodiu
m isoetifoliu
msan
d/coral ru
bble
Cym
odocea rotun
datasan
d
Halophila ovalis
sand
Halophila ovalis
sand
Halophila stipu
laceasan
d
Figure 4.3. Thalassia hemprichii and Thalassondendron ciliatum
(m ±
stdev) in locations and habitat of occurrence. For T. hemprichii on reef fl ats in
Vohemar, the stdev goes off the scale, sd =
340.
Seagrasses and algae of north-eastern Madagascar
51 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
In N
osy An
kao, T. hem
prichii dom
inated
sand
ban
ks, fol-low
ed b
y C. rotu
ndata an
d T
. ciliatum
(average shoot d
ensity
87.5
±62 sh
oot m-2) th
at were fou
nd
on th
e inn
er reef fl at on
coral rubble. In
this area, T
. hemprichii sh
oot den
sity ranged
from
116.4
±65.3
in th
e coastal lagoons to 1
64.5
±118.9
sh
oot m-2 on
the reef fl at. C
. serrulata w
as also presen
t in th
is area close to a fresh
water ru
noff .
In A
mbod
ivahib
e Bay seven
species of seagrass w
ere foun
d,
wh
ere T. hem
prichii and
C. rotu
ndata w
ere largely presen
t on
the san
d fl at b
etween
the m
angrove frin
ge and
the reef fl at.
On
the sou
thern
side of th
e bay, seagrasses form
ed m
ixed
intertid
al and
subtid
al comm
un
ities wh
ere T. hem
prichii (m
ean sh
oot den
sity 158.7
±140 sh
oot m-2), C
. rotun
data and
S. isoetifoliu
m w
ere most ab
un
dan
t. Near fresh
water ru
noff
also on th
e south
ern en
d of th
e bay, Z
. capensis w
as com-
mon
ly distrib
uted
on th
e intertid
al. T. ciliatu
m w
as sparsely
distrib
uted
on coral ru
bble on
the in
ner p
art of the reef fl at,
with
an average sh
oot den
sity of 235±100 sh
oot m-2.
In V
ohem
ar and
An
dravin
a Bay fi ve sp
ecies of seagrass w
ere foun
d. In
Voh
emar, th
e extensive reef fl at an
d associ-
ated san
d b
ars were m
ostly dom
inated
by mixed
assemblages
of T. hem
prichii, C. rotu
ndata,H
. un
ivervis and
S. isoetifolium
. In
areas of the fl at th
at are more exp
osed to sed
imen
t dis-
turb
ance, H
. ovalis is comm
only fou
nd
. On
the N
orthw
est sid
e of the sh
ore, toward
s the river d
elta, T. hem
prichii and
H
. un
inervis w
ere the on
ly species p
resent. In
this area, T
. hem
prichii show
ed th
e high
est shoot d
ensity of all sites, w
ith
den
sities of 386.9
±337.8
on th
e reef fl at and
72.9
±113.4
sh
oot m-2.
In A
nd
ravina B
ay, the sou
thern
side of th
e bay is exp
osed
to the w
ind
fetch, w
here T
. ciliatum
is foun
d on
hard
sub-
strate on th
e inn
er edge of th
e reef and
sparse S. isoetifoliu
m
can b
e foun
d on
sand
patch
es. In th
is area, T. ciliatu
m sh
oot d
ensity averaged
200±75 sh
oot m-2. In
the sh
allow, q
uiescen
t lagoon
, near th
e river mou
th, H
alophilasp an
d H
. wrightii
were th
e only sp
ecies presen
t.
CON
CLUSIO
NS A
ND
RECOM
MEN
DATIO
NS
Seagrass bed
s are in p
ristine con
dition
along th
is coastline,
except in
proxim
ity of sewage ou
tfalls or larger hu
man
settlem
ents. In
these areas (i.e. V
ohem
ar), the h
igh organ
ic m
atter conten
t in th
e sedim
ent an
d w
ater turb
idity in
crease ep
iph
yte growth
. T
en sp
ecies of seagrass werefou
nd
along th
e coast of n
ortheastern
Mad
agascar: Th alassoden
dron ciliatu
m, Th
alassia
hemprichii, Syrin
godium
isoetifolium
, Cym
odocea rotun
data an
d C
. serrulata, H
alodule u
nin
ervis and
H. w
rightii, Halo-
phila ovalis and
H. stipu
lacea, Zostera capen
sis. In d
eeper
water (i.e. b
elow 1
0 m
) (i.e. on th
e outer ed
ge of coral reefs) in
light lim
ited con
dition
s, only H
alophila species w
ere fou
nd
.N
inety on
e species of algae w
ere recorded
in th
e surveys,
dom
inated
by the red
algae (45 sp
p), follow
ed by th
e green
then
brow
n algae.
As coastal d
evelopm
ent an
d p
opu
lation grow
th in
creases in
the areas su
rveyed, seagrasses an
d algae w
ill face growin
g p
ressures su
ch as m
echan
ical removal for in
frastructu
re d
evelopm
ent (h
otels, marin
as, etc), increasin
g sedim
ent ru
n-
off du
e to deforestation
, and
prop
eller and
anch
or dam
age as p
ower en
gines b
ecome m
ore readily availab
le. Th
e imp
acts of clim
ate chan
ge will also likely aff ect seagrass ecosystem
s, th
rough
increase in
sea surface tem
peratu
re (expected
to rise u
p to 0
.6 °C
in th
e Region
) and
sea level rise (pred
icted u
p
to 50cm
by 2
100) an
d ch
anges in
storms/cyclon
e pattern
s, freq
uen
cy and
inten
sity.T
o main
tain th
e fun
ctions an
d ecosystem
services of algal an
d seagrass ecosystem
s the follow
ing recom
men
dation
s sh
ould
be im
plem
ented
:
1.
Estab
lish m
onitorin
g program
s to assess chan
ges in
algae and
seagrass bed
s over time; Th
is w
ould
be p
ar-ticu
larly imp
ortant in
areas wh
ere Con
servation A
reas m
ay be estab
lished
(e.g. Am
bod
ivahib
e Bay) an
d in
p
roximity to villags an
d u
rban
areas (e.g. Voh
emar).
2.
Create m
arine p
rotected areas (M
PA
) that w
ill redu
ce or avoid
altogether fu
ture im
pacts on
coastal ecosystems,
inclu
din
g man
groves, seagrasses and
coral reefs;
3.
En
force existing an
d estab
lish n
ew req
uirem
ents an
d
zonin
g codes for su
stainab
le coastal develop
men
t that
min
imizes im
pacts on
shallow
coastal ecosystems;
4.
Estab
lish m
arine p
rotected areas th
at limit or b
an
destru
ctive fi shin
g practices th
at up
root and
remove sea-
grass to rebu
ild h
ealthy m
arine ecosystem
s, fi sh larvae
and
pop
ulation
s;
5.
Prom
ote sustain
able aq
uacu
lture p
ractices (eg. low d
en-
sity and
chem
ical-free) that m
inim
ize waste an
d ru
noff
that cau
se algal bloom
s redu
cing ligh
t availability;
6.
En
courage w
aste man
agemen
t solution
s to add
ress im
prop
er disp
osal and
critically redu
ce eutrop
hication
, organ
ic matter p
ollution
and
plastic p
ollution
into th
e coastal zon
e;
7.
Restore m
angroves an
d forests to p
revent siltation
and
ru
noff th
at inh
ibits seagrass grow
th an
d can
cause
seagrass bu
rial;
8.
En
han
ce awaren
ess camp
aigns on
the im
portan
ce of coastal ecosystem
s, particu
larly man
groves and
seagrasses to en
sure th
ey are specifi cally ad
dressed
in
coastal man
agemen
t plan
s and
comm
un
ity engagem
ent
in coastal p
rotection is en
han
ced;
9.
Hold
trainin
g sessions for coastal m
anagers to im
prove
mon
itoring skills an
d d
evelop eff ective con
servation
measu
res;
Chapter 4
52Rapid Assessm
ent Program
10.
Con
du
ct outreach
program
s for comm
un
ities, govern-
men
ts, and
tourists to h
elp th
em get in
volved in
the
conservation
of ecosystems an
d th
e services they
provid
e.
Given
their im
portan
ce for in th
e marin
e environ
men
t an
d th
eir ecosystem services, th
e full assessm
ent of algae
assemblages is critical to d
efi ne m
arine con
servation sites
and
establish
new
marin
e protected
areas (MP
A). A
lgal d
istribu
tion can
be u
sed as an
ecological ind
icator and
their
presen
ce entails im
portan
t nu
tritional elem
ents th
at are ben
efi cial to fi sh com
mu
nities an
d oth
er marin
e organism
s. Th
erefore, in
clud
ing algal su
rveys in ecological m
onitorin
gof M
PA
s provid
es inform
ation on
site prod
uctivity an
d
ecosystem h
ealth. A
s the m
arine algae of M
adagascar are
little know
n, it is im
portan
t to cond
uct an
exhau
stive inven
-tory to w
iden
man
agemen
t interven
tions an
d ad
opt m
ore d
etailed m
anagem
ent p
lans.
REFERENCES
Bran
ch, G
.M. an
d C
.A. M
oreno. 1
994. In
tertidal an
d
subtid
al grazers. In: Sigfried
W.R
. (ed.). R
ocky shores:
exploitation
in C
hile an
d Sou
th A
frica. Sprin
ger-Verlag,
Berlin
. Germ
any. P
p 7
5–100.
Brau
n-B
lanqu
et, J. 1972. P
lant sociology: th
e stud
y of plan
com
mu
nities. H
afner, N
ew York.
Costan
za, R., R
. d’A
rge, R. d
e Groot, S. Farb
er, M. G
rasso, B
. Han
non
, K. L
imbu
rg, S. Naeem
, R.V
. O’N
eill, J. P
aruelo, R
.G. R
askin, P. Su
tton an
d M
. van d
en B
elt. 1997. Th
e valu
e of the w
orld’s ecosystem
services and
n
atural cap
ital. Natu
re 387: 2
53-2
60.
Krau
fvelin, P., A
. Lin
dh
olm, M
. F. Ped
ersen, L
.A. K
irkerud
, an
d E
. Bon
sdorff . 2
010. B
iomass, d
iversity and
prod
uc-
tion of rocky sh
ore macroalgae at tw
o nu
trient en
rich-
men
t and
wave action
levels. Marin
e Biology 1
57(1
): 29-4
7.
Orth
, R.J., T
.J.B. C
arruth
ers, W.C
. Den
nison
, C.M
. Du
arte, J.W
. Fou
rqu
rean, K
.L. H
eck, A.R
. Hu
ghes, G
.A. K
end
-rick, W
.J. Ken
worth
y, S. Olyarn
ik, F.T. Sh
ort, M. W
ay-cott an
d S.L
. William
. 2006. A
global crisis for seagrass
ecosystems. B
ioScience 5
6(1
2): 9
87-9
96.
Short ,F.T
., T.J.B
. Carru
thers, W
.C. D
enn
ison an
d M
. Way-
cott. 2007. G
lobal seagrass d
istribu
tion an
d d
iversity: A
bioregion
al mod
el. Exp
erimen
tal Marin
e Biology an
d
Ecology 3
50: 3
-20.
Wein
berg, S. 1
976. Su
bm
arine d
aylight an
d ecology, M
ar. B
iol.37, 2
91–304.
53A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
INTRO
DU
CTION
Ou
r best estim
ates of global b
iodiversity are p
oor. Even
getting w
ithin
the correct ord
er of m
agnitu
de h
as proven
to be a d
iffi cult goal. E
ntom
ologists workin
g in rain
forests have
provid
ed th
e best w
orking estim
ates of terrestrial diversity. B
ased on
the correlation
betw
een
structu
ral species (trees) an
d h
ost specifi c sym
bion
ts (herb
ivorous in
sects), they h
ave estimated
glob
al insect ab
un
dan
ce at 2-5
0 m
illion sp
ecies. Symbioses are u
biq
uitou
s on coral reefs, yet
a similar ap
proach
has n
ot been
taken w
ith regard
to estimatin
g the n
um
bers of h
ost-specifi c
reef associates, and
the fu
rther extrap
olation to estim
ates of reef diversity. Th
is w
ork will b
uild
u
p a w
orking collection
of marin
e structu
ral species (h
exacorals, octocorals, etc.) and
their
associated sp
ecies (families of crab
s, shrim
p, barn
acles, gastropod
s, bivalves, am
ph
ipod
s etc.) th
at in th
e futu
re may con
tribu
te to derivin
g a total biod
iversity estimate for th
e region.
METH
OD
S
Stand
ard collectin
g meth
ods w
ere used
- searchin
g structu
ral species in
the fi eld
for externally
visible exosym
bion
ts or anom
alies in th
e surface textu
re. Samp
ling sites are sh
own
on T
able 1
in
the “R
eport at a G
lance” section
on p
age 13. If visib
le exosymbion
ts or anom
alies were
seen, sam
ples of th
e host w
ere collected an
d later rin
sed in
ethan
ol and
preserved
for mu
seum
collection
s. Coral gen
etic samp
les were p
reserved in
95%
ethan
ol and
dim
ethyl su
lph
oxide
(DM
SO). Skeletal sp
ecimen
s were d
ried in
the su
n. O
ctocorals were p
reserved in
ethan
ol. Th
e associated
faun
a separated
from th
e skeletons w
as preserved
usin
g ethan
ol. Samp
ling w
as op
portu
nistic th
rough
out th
e RA
P su
rvey, creating on
e datab
ase, not site-sp
ecifi c inven
tories.
RESULTS
In all, 1
05 sam
ples w
ere taken, com
prisin
g app
roximately 5
1 stru
ctural sp
ecies (i.e. Op
era-tion
al Taxon
omic U
nits, O
TU
s), of wh
ich 6
were A
ntip
atharia (b
lack corals), 16 w
ere O
ctocorallia (soft corals) and
83 w
ere Hexacoralia (h
ard corals) (see ap
pen
dices A
, B an
d C
). A
mon
g these, a p
relimin
ary list of 66 d
iff erent exosym
bion
ts was d
istingu
ished
. Prelim
inary
sorting in
the fi eld
show
ed a h
igh d
egree of specifi city of h
osts and
symbion
ts (fi gure 5
.1),
with
a small n
um
ber of h
osts and
symbion
ts bein
g high
ly polyvalen
t. Of th
e 66 exosym
bion
ts, 51 (7
7 %
) were fou
nd
in a sin
gle host sp
ecies (fi g. 5.1
, left), with
only 2
taxa bein
g foun
d in
m
ore than
5 h
ost taxa. Of th
e 51 h
ost taxa, 30 (5
9 %
) were fou
nd
with
a single sym
bion
t spe-
cies (fi g. 5.1
, right), w
ith on
ly 5 h
ost taxa havin
g more th
an 5
symbion
t taxa. More d
etailed
taxonom
ic sorting w
ill requ
ire several years of work to ach
ieve satisfactory assignm
ent to
species.Several of th
e most w
ell know
n fam
ilies of exosymbion
ts proved
to be of in
terest. Both
T
rapezia (Pocillop
orid associated
crabs) an
d T
etralia (Acropora associated
crabs) w
ere abu
nd
ant,
Chapter 5
Coral-associated exosymbionts of
northeast Madagascar
Sea McKeon
Chapter 5
54Rapid Assessm
ent Program
and
revealed u
nexp
ected d
iversity in color an
d p
attern.
Gen
etic meth
ods w
ill be u
sed to u
ntan
gle the taxon
omic
issues, an
d verify th
e presen
ce of un
describ
ed taxa. A
s these
crabs p
rovide essen
tial services for their coral h
osts, an
un
derstan
din
g as to the p
atterns p
resent in
the W
estern
Ind
ian O
cean is of im
portan
ce in th
e conservation
of the
area’s reef systems. C
ryptoch
irid crab
s were also ab
un
dan
t, an
d exceed
ed th
e nu
mbers of sp
ecies previou
sly know
n for
the region
.
CON
SERVATION
RECOM
MEN
DATIO
NS
At p
resent n
o specifi c recom
men
dation
s can b
e given for
exosymbion
ts – as th
ey are high
ly specifi c to th
eir host sp
e-cies, con
servation action
s for those w
ill be ap
plicab
le to the
symbion
ts as well.
As h
as been
foun
d in
other region
s, explorin
g a new
area for coral exosym
bion
ts has yield
ed a large n
um
ber of u
nd
es-crib
ed form
s and
likely new
species. P
rediction
s of total biod
iversity based
on th
ese new
forms are h
igh, em
ph
asizing
the n
eed for con
servation m
easures to p
revent loss of h
abitat
and
host sp
ecies before th
ese un
know
n sp
ecies are describ
ed.
Th
ey provid
e a poten
t ind
icator of the loss of u
nkn
own
spe-
cies that occu
rs wh
en h
abitat loss occu
rs.
Figure 5.1. Association of exosymbionts with host taxa.
# hosts with 'x' symbiont taxa
# symbtions on 'x' host taxa
# symbiont taxa per host
# host taxa per symbiont
0 3 6 9 12 15 18 21 24 27 30
87
65
43
21
0 5 10 15 20 25 30 35 40 45 50 55
1110
98
76
54
32
1
Table 5.1. List of host and symbiont taxa.
Host TaxonSym
biont Taxa
Antipatharia
Cirrip
athes A
Bryan
iopsis sp.
Cirrip
athes A
Pon
toniin
ae sp.
Cirrip
athes B
Pon
toniin
ae sp.
Cirrip
athes C
Pon
toniin
ae sp.
Cirrip
athes D
Bryan
iopsis sp.
Cirrip
athes D
Pon
toniin
ae sp.
Hexacorals
Acropora abrotan
oidesT
etralia “blu
e eyes” *
Acropora abrotan
oidesT
etralia glabberima
Acropora cf. apressa
Tetralia glabberim
a
Acropora hyacin
thus
Coralliocaris cf. su
perba
Acropora hyacin
thus
Cym
o sp.
Acropora hyacin
thus
Jocaste sp.
Acropora hyacin
thus
Tetralia “b
lue eyes” *
Acropora hyacin
thus
Tetralia “ob
scura”*
Acropora hyacin
thus
Tetralia glabberim
a
Acropora hyacin
thus
Tetralia ru
bridactyla
Acropora m
uricata
Pyrgom
atidae
Acropora palifera
Lith
oph
agidae sp. A
Acropora palifera
Pon
toniin
e sp. A.
Acropora palifera
Tetralia n
igrolineata
Acropora sp. ‘m
ayote’T
etralia “blu
e eyes” *
Acropora su
mm
erensis
Tetralia “ob
scura”
Acropora ten
uis
Coralliocaris cf. su
perba
Acropora ten
uis
Coralliocaris cf. viridis
Acropora ten
uis
Reliqu
icava sp.
table contin
ued on
next page
Coral-associated exosymbionts of northeast M
adagascar
55 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Host TaxonSym
biont Taxa
Hexacorals (continued)
Acropora ten
uis
Tetralia “b
lue eyes” *
Acropora ten
uis
Tetralia “ob
scura”*
Acropora ten
uis
Tetraloides “gogators”
Alveopora
Coralliophila n
eritoides
Coscin
aria crassaC
ryptoch
iridae sp.
Coscin
aria crassaP
yrgomatid
ae sp.
Coscin
aria mon
ileC
ryptoch
iridae sp.
Echin
opora gemm
aceaC
ryptoch
iridae sp.
Favites A
Cryp
tochirid
ae sp.
Galaxea astreata
Pon
toniin
e sp.
Galaxea fascicu
larisIschn
oponton
ia lophos
Galaxea fascicu
larisR
acilius com
pressus
Gon
iastrea pectinata
Cryp
tochirid
ae sp.
Gyrosm
ilia interru
ptaSyn
alpheus sp.
Leptoria phrygia
Cryp
tochirid
ae sp.
Meru
lina am
pliataC
ryptoch
iridae sp.
Oxypora ‘lacera’
Cryp
tochirid
ae sp.
Pavon
a clavus
Opercarcin
us sp. F
Pavon
a duerden
iO
percarcinus sp. E
Pavon
a maldiven
sisO
percarcinus sp. D
Playtgyra daedelae
Cryp
tochirid
ae sp.
Pleu
rogyra sinuosa
Vir phillipen
sis
Pocillopora dam
icornis
Alpheu
s lottini ‘sp
ots’
Pocillopora dam
icornis
Alpheu
s lottini “strip
e”
Pocillopora dam
icornis
Hapalocarcin
us sp. A
Pocillopora dam
icornis
Paragobiodon
cf. lacun
icolus
Pocillopora dam
icornis
Synalpheu
s charon
Pocillopora dam
icornis
Trapezia gu
ttata
Pocillopora eydou
xiH
apalocarcinus sp. C
*
Pocillopora eydou
xiH
arpiliopsis beaupressi
Pocillopora eydou
xiT
rapezia digitalis
Pocillopora eydou
xiT
rapezia rufopu
nctata
Pocillopora eydou
xiU
tinom
iella sp. B
Pocillopora verru
cosaA
lpheus lottin
i ‘spots’
Pocillopora verru
cosaParagobiodon
echinocephalu
s
Pocillopora verru
cosaSyn
alpheus charon
Pocillopora verru
cosaT
rapezia cf. bidentata*
Pocillopora verru
cosaT
rapezia cf. lutea
Pocillopora verru
cosaT
rapezia cf. speciosa
Pocillopora verru
cosaT
rapezia cymodoce
Host TaxonSym
biont Taxa
Pocillopora verru
cosaT
rapezia richtersi
Pocillopora w
oodjonesi
Hapalocarcin
us sp. B
*
Pocillopora w
oodjonesi
Quoyu
la sp. A
Pocillopora w
oodjonesi
Trapezia tigrin
a
Pocillopora w
oodjonesi
Utin
omiella sp. A
Porites lobata
Lith
oph
agidae sp. B
Porites lobata
Pagu
ritta sp.
Porites lobata
Pedu
m spon
dyloideum
Porites ru
sC
oralliophila sp.
Seriatopora hysterixA
lpheus lottin
i “stripe”
Seriatopora hysterixH
apalocarcinus sp. D
*
Seriatopora hysterixT
rapezia guttata
Stylophora pistillata A
lpheus lottin
i “stripe”
Stylophora pistillata H
apalocarcinus sp. E
*
Stylophora pistillata T
rapezia cf. lutea
Stylophora pistillata T
rapezia cymodoce
Stylophora pistillata “little”A
lpheus lottin
i ‘spots’
Stylophora pistillata “little”H
apalocarcinus sp. F
*
Stylophora pistillata “little”T
rapezia guttata
Tubastrea m
icrantha
Xan
thoid
ea sp.
Turbin
aria BO
percarcinus sp. B
.*
Turbin
aria CO
percarcinus sp. C
.*
Turbin
aria mesen
terina
Opercarcin
us sp
A.*
Wan
derin
g Coral
Wan
derin
g Coral W
orm
Octocorals
Gorgon
ia sp. AG
alathea sp.
Gorgon
ia sp. AM
ysida sp.
Gorgon
ia sp. AP
onton
iinae sp.
Gorgon
ia sp. BP
onton
iinae sp.
Lobophyton
sp.O
vulidae sp.
Sarcophyton sp.
Pon
toniin
ae sp.
Scleronephthea 1
Galathea sp.
Scleronephthea 1
Mysida sp.
Scleronephthea 1
Ovu
lid A
Scleronephthea 1
Pon
toniin
ae sp.
Scleronephthea 2
Mysida sp.
Scleronephthea 2
Pon
toniin
ae sp.
Sinularia
Mysida sp.
Sinularia
Pon
toniin
ae sp.
Tubipora m
usica
Alpheidae sp.
Tubipora m
usica
Pon
toniin
ae sp.
Table 5.1. continued
56Rapid Assessm
ent Program
Chapter 6
Coral reef health and status
David Obura and Tom Oliver
SUM
MA
RY
To p
rioritize ecologically resilient reefs for con
servation action
, we evalu
ated th
e sites surveyed
in
the N
ortheast M
adagascar M
arine R
AP
usin
g resilience assessm
ent m
ethod
s develop
ed b
y IU
CN
and
CO
RD
IO E
ast Africa. O
verall the region
show
ed h
igh coral cover (m
ean of 4
8%
), an
d coral p
opu
lations th
at span
ned
the exp
ected size ran
ge for the gen
us in
qu
estion, in
clu-
din
g large, matu
re colonies. Th
e com
mon
occurren
ce of large colonies su
ggests that th
ere has
been
no catastrop
hic m
ortality in th
e past few
decad
es. Coral recru
itmen
t was p
resent, b
ut
relatively low.
Even
after an exten
ded
heatin
g event th
at end
ed im
med
iately preced
ing th
e survey, coral
bleach
ing w
as relatively low, sh
owin
g a mean
~5%
of colonies aff ected
. Th
is, and
the low
im
pact d
etected from
the 1
998 m
ass bleach
ing even
t, suggests th
at the corals in
this region
h
ave largely resisted th
e negative eff ects of h
eating even
ts, and
sup
ports th
e hyp
othesis th
at th
ese reefs are therm
ally resistant. In
general, th
e north
ern th
ree locations (A
mbod
ivahib
e, N
osy An
kao, Loky B
ay) show
ed greater coral cover, larger corals an
d few
er bleach
ed colon
ies th
an th
e south
ern tw
o sites (An
dravin
a, and
Voh
emar). C
oral recruitm
ent w
as high
est on th
e n
orthern
and
south
ern extrem
es, in A
mbod
ivahib
e and
Voh
emar, th
ough
overall was com
para-
tively low.
Th
e generally in
tact structu
re of the reefs an
d ap
paren
t resistance to coral b
leachin
g suggests
that th
e region is a p
riority for conservation
. We recom
men
d con
servation action
s to enh
ance
coral recruitm
ent, a key factor for ecological resilien
ce, throu
gh th
e targeted restriction
of fi -sh
ing p
ressure an
d w
atershed
man
agemen
t to limit/red
uce sed
imen
tation.
INTRO
DU
CTION
Coral reefs an
d th
eir associated seagrass b
eds an
d m
angrove h
abitats su
pp
ort the h
ighest
marin
e biod
iversity in th
e world
(Kn
owlton
et al. 2010) as w
ell as the livelih
oods an
d econ
o-m
ies of million
s of coastal peop
le (Mob
erg 2009). Th
e coral reefs of M
adagascar h
ave long
been
recognized
as a key asset in su
pp
orting th
e du
al aims of b
iodiversity con
servation an
d
poverty alleviation
. A
n issu
e of prim
ary concern
for coral reefs is climate ch
ange, n
ow recogn
ized as on
e of the
greatest threats to coral reefs w
orldw
ide (H
oegh-G
uld
berg et al. 2
007). M
ass coral bleach
ing
remain
s one of th
e most im
med
iate imp
acts of climate ch
ange on
corals reefs, as abn
ormally
high
water tem
peratu
res trigger the b
reakdow
n of th
e coral-algal symbiosis an
d can
lead to
mass coral m
ortality (Coles an
d B
rown
2003). O
ther factors th
at aff ect reefs in th
e region
inclu
de cyclon
es, terrestrial sedim
ent ru
n-off , p
redator ou
tbreaks su
ch as crow
n of th
orns sea-
stars, and
anth
ropogen
ic threats su
ch as fi sh
ing, p
ollution
, and
nu
trient ad
dition
s (Wilkin
son
2004).
Each
of these factors aff ects th
e ecological state of reefs, and
alone or in
concert th
ey can act
to drive th
e reef from a h
ighly d
iverse system cap
able of p
rovidin
g susten
ance for m
any p
eople
Coral reef health and status
57 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
to a degrad
ed state th
at sup
ports few
species an
d su
stains
few p
eople. Th
e likelih
ood th
at a given reef w
ill succu
mb to
these factors an
d slid
e dow
n th
is scale of “reef health” can
be
explain
ed in
terms of th
e reef’s ecological resilience –
i.e. its ab
ility to resist threats an
d to recover to a h
ealthy state w
hen
an
imp
act does occu
r (Obu
ra and
Grim
sditch
2009).
Th
e natu
ral resilience of reefs is b
eing u
nd
ermin
ed b
y stresses associated
with
hu
man
activities and
these local
pressu
res redu
ce the resilien
ce of the system
by redu
cing
its ability to cop
e with
add
itional stresses, su
ch as th
ose p
resented
by clim
ate chan
ge (Hoegh
-Gu
ldberg et al. 2
007).
Increasin
gly, policy-m
akers, conservation
ists, scientists an
d
the b
roader com
mu
nity are callin
g for man
agemen
t actions
to restore and
main
tain th
e resilience of th
e coral reefs to clim
ate chan
ge, and
thu
s avoid w
orst-case scenarios. T
o assist m
anagem
ent au
thorities in
focusin
g man
agemen
t eff orts on
priority areas, th
e IUC
N C
limate C
han
ge and
Coral R
eefs w
orking grou
p (h
ttp://cm
s.iucn
.org/cccr), led by C
OR
DIO
E
ast Africa, h
as outlin
ed a series of p
rotocols to qu
antify
basic resistan
ce and
resilience in
dicators for coral reefs.
We ap
plied
this resilien
ce assessmen
t meth
od to th
e reefs su
rveyed in
the N
ortheast M
adagascar M
arine R
AP. Th
is
comp
onen
t of the R
AP
focuses on
reef health
and
resilience,
to more d
irectly sup
port con
servation ou
tcomes recom
-m
end
ed b
y the sp
ecies-oriented
comp
onen
ts.N
ortheast M
adagascar is a p
articularly in
teresting region
for th
ose interested
in reef resilien
ce in th
e face of climate
chan
ge. To d
ate, man
y regions in
Mad
agascar have su
ff ered
from n
otable b
leachin
g-ind
uced
mortalities (Q
uod
et al. 2002, N
OA
A 2
007, M
aharavo et al. in
press, O
bu
ra 2009),
how
ever, reefs in N
ortheast M
adagascar h
ave reported
ly escap
ed m
ajor bleach
ing im
pacts (W
ebster an
d M
cMah
on
2002, M
cKen
na et al. 2
005). O
ur su
rvey aimed
to critically evalu
ate these fi n
din
gs, and
bu
ild a b
roader p
erspective on
reef resilien
ce in th
e region.
METH
OD
S
Th
e meth
ods th
at we ap
plied
in th
is stud
y were d
eveloped
by th
e IUC
N w
orking grou
p on
Clim
ate Ch
ange an
d C
oral R
eefs, as a rapid
assessmen
t of the resilien
ce of coral reefs to clim
ate chan
ge and
its most im
med
iate conseq
uen
ce, h
igh seaw
ater temp
erature. C
onseq
uen
tly, its objectives
and
meth
odology are con
sistent w
ith th
ose of the m
arine
RA
Ps, ad
din
g a comp
onen
t on reef h
ealth to th
e biod
iversity assessm
ent. Several com
pon
ents of th
e reef ecosystem w
ere m
easured
at varying levels of d
etail, as follows:
1.
Ben
thic cover –
provid
es the m
ain overall in
dicators
of reef state, and
particu
larly the b
alance b
etween
corals an
d algae. B
enth
ic cover was estim
ated by eye,
and
reported
here. B
enth
ic ph
oto qu
adrats w
ere also collected
for futu
re analysis an
d for verifi cation
of visual
estimates if n
ecessary.
2.
Flesh
y algae – p
rovides in
formation
on th
e main
com-
petitors to corals on
degrad
ing reefs. F
leshy algae cover
(%) an
d h
eight (cm
) were estim
ated.
3.
Coral com
mu
nity stru
cture –
provid
es an overview
of th
e relative abu
nd
ance of coral gen
era, and
that are
suscep
tible or resistan
t to coral bleach
ing. Th
e ab
un
-d
ance of all coral gen
era was estim
ated d
urin
g fi eld visits
along a fi ve-p
oint scale (0
-absen
t, 1-rare, 2
-un
comm
on,
3-com
mon
, 4-ab
un
dan
t, 5-d
omin
at).
4.
Coral p
opu
lation stru
cture –
the size class d
istribu
tion
of selected corals p
rovides d
etailed in
formation
on th
eir d
emograp
hy (in
clud
ing recru
itmen
t, growth
and
mor-
tality). It inclu
des sam
plin
g of small corals (<
= 1
0cm
) in
1 m
2 qu
adrats, an
d larger corals (>
10 cm
) in b
elt tran
sects of 1m
wid
th (th
e length
of transect sam
pled
is m
aximized
, with
in th
e constrain
ts of time an
d op
portu
-n
ity, with
> 5
0m
bein
g desirab
le).
5.
Coral th
reats – gives an
ind
ication of th
e curren
t health
of th
e coral comm
un
ity, and
inclu
des ob
servations on
coral b
leachin
g, disease, an
d m
ortality, and
presen
ce of p
redators an
d th
reats such
as crown
of thorn
s stars.
6.
Fish
herb
ivores and
other fu
nction
al group
s – fi sh
exert p
rimary con
trol on th
e reef comm
un
ity, and
on
algae throu
gh h
erbivory, th
us con
trolling com
petition
betw
een algae an
d corals. Th
is d
ata was collected
and
rep
orted u
nd
er the sep
arate RA
P com
pon
ent on
fi sh
biod
iversity and
biom
ass.
7.
Resilien
ce ind
icators – variou
s add
itional factors th
at aff ect th
e resistance of corals to b
leachin
g and
the
resilience or recovery p
otential of th
e reef comm
u-
nity. A
broad
range of in
dicators in
diff eren
t classes is m
easured
, inclu
din
g of aspects in
1-6
above, b
ut at less
qu
antitative levels. A
detailed
descrip
tion of th
e me-
thod
ology is available in
Obu
ra and
Grim
sditch
(2009).
Th
e main
classes of ind
icators are listed in
Tab
le 6.1
.
RESULTS
Resilience IndicatorsIn
dicators are su
mm
arized in
Tab
le 6.2
for ben
thic cover
(%), site d
epth
(m) an
d in
dicators of p
hysical an
d coral com
-m
un
ity characteristics (1
-5 scale). D
etailed resu
lts follow.
Hard
corals were th
e dom
inan
t cover at all the areas
visited, con
fi rmin
g the h
ealthy state of th
e coral reefs of th
e region. C
oral cover averaged 4
8%
throu
ghou
t the
area (Fig. 6
.1), w
ith h
ighest cover in
Voh
emar (6
0%
) and
d
ecreasing to 2
1%
in A
nd
ravina.
Sites with
the low
est coral cover (A17, A
05, A
20, etc)
all show
ed h
igh in
fl uen
ce of sedim
entation
and
/or wave
energy, th
e low coral cover b
eing a n
atural resp
onse to th
ese
Chapter 6
58Rapid Assessm
ent Program
factors. Soft corals were also ab
un
dan
t at ≈ 2
0%
cover, at a sim
ilar level to algal turf. Th
e h
igh ab
un
dan
ce of soft corals is con
sistent w
ith th
e app
arently h
igh levels of n
utrien
ts in
the area, as a resu
lt of freshw
ater run
off from lan
d, an
d h
igh
levels of wave en
ergy along th
e exposed
coast. Tu
rfs were
the d
omin
ant algal form
foun
d on
the reefs at an
average of 20%
, with
macroalgae averagin
g < 5
%. V
ohem
ar had
high
er m
acroalgal abu
nd
ance th
an th
e other areas, likely a resu
lt of eu
troph
ication an
d h
eavy fi shin
g pressu
re.W
hile sam
plin
g for resilience in
dicators focu
sed on
shal-
low sites ab
out 1
0 m
, we fou
nd
that reef p
rofi les were very
shallow
, with
man
y sites havin
g the m
ain zon
e of coral deve-
lopm
ent at <
10 m
(Tab
le 6.2
). In m
ost cases, the reef m
axi-m
um
reef dep
th w
as at 12-1
8m
, wh
ere the rocky su
bstrate
gave way to san
d an
d ru
bble slop
es. Given
the accu
mu
lation
of fi ne sed
imen
t and
the relatively low
light levels at th
ese d
epth
s, it app
ears that coral grow
th m
ay have b
een lim
ited
by sed
imen
tation an
d/or h
igh tu
rbid
ity.P
hysical ch
aracteristics of the areas are su
mm
arized in
F
ig. 6.2
. Exp
osure of reefs to w
ave energy w
as generally
mod
erate to high
, thou
gh th
e most h
ighly exp
osed sites in
th
e region w
ere not sam
pled
du
e to accessibility p
roblem
s. Th
e fore reef at V
ohem
ar was th
e most exp
osed site sam
pled
an
d sh
owed
characteristics of th
is exposu
re, with
strong
develop
men
t of reef spu
rs and
pillars sim
ilar to structu
res fou
nd
farther sou
th in
the M
asoala pen
insu
la. Th
e most
sheltered
sites were w
ithin
Loky B
ay and
show
ed ch
aracteris-tically h
igh levels of fi n
e terrigenou
s silt. Ch
aracteristics that
contrib
ute to coolin
g of surface w
aters (up
wellin
g, proxim
i-ty to d
eep w
ater, mixin
g by w
ave energy) w
ere foun
d to b
e h
ighest at A
mbod
ivahib
e and
Voh
emar. Th
ey w
ere lower at
the oth
er three location
s as these w
ere either w
ithin
bays, or
on large p
latforms d
istancin
g them
from cooler d
eep w
aters.A
spects of coral com
mu
nity stru
cture are su
mm
arized in
F
ig. 6.3
. Bleach
ing w
as at mod
erately low levels th
rough
out
the region
, refl ecting a region
al bleach
ing alert d
ue to th
e p
resence of a m
oderate h
otspot in
the region
(see NO
AA
an
d C
OR
DIO
alerts). Bleach
ing levels in
creased sou
th-
ward
s, with
high
est levels recorded
at the sou
thern
sites of A
nd
ravina an
d V
ohem
ar, correspon
din
g to up
to 20%
of corals sh
owin
g some level of b
leachin
g or palin
g. Mortality
as a result of b
leachin
g was n
ot app
arent, h
owever. Th
e lack
of mortality ob
served so far m
ay ind
icate that m
ortality from
the m
oderate b
leachin
g event m
ay be lim
ited, w
ith n
o m
ajor imp
act on th
e coral comm
un
ities in th
e short term
. R
epeat ob
servations of som
e of the stu
dy sites, p
articularly
at Am
bod
ivahib
e and
Voh
emar in
mid
-2010 cou
ld test th
is. Q
uan
titative bleach
ing tran
sect data is p
resented
below
(See C
oral Bleachin
g Data).
Disease an
d m
ortality levels for corals were low
throu
gh-
out th
e region, con
fi rmin
g the low
imp
act of anth
ropogen
ic an
d d
isease-causin
g agents. C
oral recruitm
ent levels w
ere
Table 6.1. Resilience Indicators
GroupFactor
VariableGroup
FactorVariable
1-C
overC
oralC
oral cover3-C
oral comm
un
itySize/age
Largest corals (3
)
Algae
Flesh
y Algae
Con
dition
Coral b
leachin
g
CC
AM
ortality-new
Substrate
Ru
bble
Mortality-old
2-P
hysical
Substrate
Top
ogr. Com
plex. - m
icroR
ecovery-old
Top
ogr. Com
pl. - m
acroC
oral disease
Sedim
ent textu
re4-C
oral associatesO
bligate feed
ers
Sedim
ent layer
Bran
chin
g residen
ts
Coolin
g & fl u
shin
gw
ater movem
ent
Com
petitors
deep
water (3
0-5
0m
)B
ioeroders (u
rchin
s, non
fi sh)
dep
th of reef b
aseB
ioeroders (in
ternal, sp
o)
wave en
ergy/ exposu
reC
orallivores (negative)
Tem
peratu
reT
emp
erature (oC
)5-A
nth
ropogen
icW
aterN
utrien
t inp
ut
Shad
ing &
screenin
gd
epth
(m)
Pollu
tion (ch
emical)
aspect
Substrate
Pollu
tion (solid
)
slope (d
egrees)T
urb
idity/ sed
imen
tation
ph
ys. shad
ing
Ph
ysical dam
age
canop
y coralsF
ishin
gD
estructive fi sh
ing
Visib
ility (m)/ tu
rbid
ityF
ishin
g pressu
re
Acclim
atizationE
xposed
low tid
e
Pon
din
g/poolin
g
Coral reef health and status
59 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Table 6.2. Selected resilience indicators (Obura & Grimsditch 2009) for sites surveyed in Northeast M
adagascar. Variable type is shown in the table, including percent cover (%
),depth (m) and indicators on a 1-5 scale, where 1 is low and 5 is high for the individual indicator.
Location/Site
Variable
Benthic comm
unityPhysical characeristics
Coral comm
unity characteristics
Hard Coral
Soft Coral
CCA
Macro algae
Turf Algae
Rubble
Depth
Exposure
Topographic Complexity
Estuarine infl uence
Cooling
Bleaching
Disease/mortality
Recruitment
Fragmentation
Mature corals
Type%
%%
%%
%m
1-51-5
1-51-5
1-51-5
1-51-5
1-5
Am
bodivah
ibe
A01
3010
030
105
102.0
2.53.3
3.12
2.52
25.0
A01B
/C60
202
510
810
3.03.0
2.23.6
22.5
22
5.0
A02/3
2520
153
2010
82.0
2.53.3
3.42
3.02
23.5
A04
600
155
205
154.0
3.02.4
4.52
2.03
34.5
A05
2025
215
255
105.0
1.52.0
3.12
1.02
23.0
Loky
A06
6020
51
130
72.0
4.02.1
3.82
1.03
24.5
A07
405
02
305
81.0
2.54.6
3.72
1.52
25.0
A08/9
2530
25
305
103.0
1.53.3
3.33
1.02
24.0
A10
3020
02
405
104.0
2.02.4
3.12
1.03
24.5
A11
852
30
50
11.0
3.52.2
2.14
1.02
35.0
A12
801
50
102
53.0
3.53.6
2.43
1.03
25.0
An
kao
A20
7020
00
53
62.0
2.02.8
2.32
1.52
25.0
A21
4010
52
205
52.0
3.02.0
2.72
2.03
24.5
A22
2050
05
202
103.0
2.52.8
3.12
2.01
24.5
A23
4535
22
105
52.0
3.03.0
2.43
1.02
24.5
A24
5520
00
205
53.0
3.04.6
2.43
2.52
25.0
An
dravin
a
A17
1510
03
605
62.0
2.54.5
2.83
1.01
14.5
A18
3015
05
305
64.0
3.53.7
2.03
1.52
23.5
Voh
emar
A13
700
153
100
105.0
3.52.7
3.64
2.02
24.0
A14
605
05
255
52.0
3.04.4
2.94
2.03
24.5
A15
4040
105
50
105.0
3.03.0
3.64
2.52
24.0
A16
7010
30
150
105.0
4.02.2
3.83
1.02
24.5
Figure 6.1. Benthic cover for Northeast Madagascar, and by sam
pling locations.
Chapter 6
60Rapid Assessm
ent Program
relatively low; fu
rther resu
lts are presen
ted b
elow from
the
size class transects. F
ragmen
tation of corals w
as relatively low
, and
mostly from
natu
ral causes su
ch as w
ave energy.
Th
e reefs show
ed little to n
o eviden
ce of anch
or scars or boat strikes, an
d d
irect dam
age by p
eople is lim
ited as fi sh
ers op
erate from b
oats and
there is n
o regular d
iving tou
rism.
Matu
re corals dom
inated
the su
rvey sites, even w
here coral
cover was relatively low
, ind
icating n
o major m
ortality of corals in
the recen
t past (1
-2 d
ecades) th
at wou
ld h
ave elimi-
nated
large matu
re colonies an
d w
ithou
t suffi cien
t time for
them
to grow b
ack.
Coral Generic A
bundanceTh
e N
ortheast M
adagascar R
AP
recorded
fi fty-eight coral
genera on
reefs (Fig. 6
.4), ran
ging from
the stron
gly dom
i-n
ant gen
era Acropora an
d P
orites, to those p
resent in
only
one or few
instan
ces – C
aulastrea, M
icromussa, San
dalolitha an
d Scolym
ia. In b
etween
these extrem
es, genera typ
ical of E
ast African
reefs were fou
nd
. Most in
teresting for region
al biogeograp
hy, th
e mon
otypic gen
era that are en
dem
ic to the
Western
Ind
ian O
cean, G
yrosmilia in
terrupta (3
6), H
orastrea in
dica (45) an
d A
nom
astrea irregularis (5
2) w
ere foun
d at
high
er abu
nd
ances th
an occu
r elsewh
ere in E
ast Africa. Th
e
most sign
ifi cant sites for th
ese genera w
ere sites A8, 9
and
10 in
Loky B
ay and
site A22 in
Nosy A
nkao for G
yrosmilia,
and
site A17 in
An
dravin
a for Horastrea an
d A
nom
astrea. Th
e region
al end
emic C
raterastrea laevis was n
ot recorded
du
ring
the su
rveys as it was con
fused
with
Leptoseris sp., th
ough
was
Figure 6.2. Physical indicators for reefs in Northeast Madagascar, on a
scale of 1 (low) to 5 (high).
Figure 6. 3. Coral comm
unity characteristics for reefs in Northeast M
adagascar, on a scale of 1 (low) to 5 (high).
Figure 6.4. Ranked abundance of coral genera in Northeast Madagascar. Indices are scaled from
5 (highest) to lowest (0), and are the following: #sites - proportion of sites at which the genus was found; ave-presence - average of abundance index at sites at which the genus was present; ave-overall - average of abundance index for each genus, including absences; m
ax - maxim
um abundance index across all sites; RA - average of all the preceding indices.
Coral reef health and status
61 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
iden
tifi ed su
bseq
uen
tly from p
hotograp
hs an
d w
as collected
at An
dravin
a.P
atterns of gen
eric comp
osition varied
across surveyed
location
s. Exam
inin
g only com
mon
genera in
qu
antitative
measu
res of colony size an
d total area (see n
ext section),
the com
petitively d
omin
ant gen
era Acropora an
d P
orites (record
ed in
both
massive an
d b
ranch
ing form
s) were
comm
on everyw
here, b
ut location
s diff ered
greatly in th
e p
resence of th
e genera ch
aracterized b
y smaller colon
y size, like Stylophora an
d Seriatopora. G
alaxea was com
mon
only at
Am
bod
ivahib
e and
Nosy A
nkao, w
here G
alaxea occurred
in
large stand
s, exclud
ing oth
er corals.
Coral Size StructureO
f these 5
8 gen
era, 23 rep
resentative an
d com
mon
genera
were su
rveyed for coral p
opu
lation size stru
cture (see IU
CN
R
esilience assessm
ent m
ethod
man
ual). H
ere we p
resent
these size stru
cture d
ata two w
ays: (1) b
y geograph
ic area for all gen
era pooled
(Figs. 6
.5 &
6.6
), and
(2) for th
e top
9 gen
era (with
Porites b
eing p
resented
as bran
chin
g and
m
assive forms sep
arately), pooled
across the w
hole stu
dy area
(Figs. 6
.7 &
6.8
). In b
oth cases w
e presen
t the size stru
cture
as (A) n
um
ber of colon
ies coun
ted in
each size class (F
igs. 6.5
& 6
.7), an
d as (B
) the total area of colon
ies in each
size class (F
igs. 6.6
& 6
.7). In
all cases, nu
mbers are given
per
stand
ard area of 1
00 m
2 to stand
ardize for d
iff erent levels of
samp
ling at each
dive site d
epen
din
g on tim
e and
logistics.Size class d
ata allow on
e to better u
nd
erstand
both
coral recru
itmen
t and
coral matu
rity pattern
s. We d
id see evid
ence
of recruitm
ent, b
ut region
al recruitm
ent levels ap
peared
gen
erally low. R
ecruitm
ent size classes (coral d
iameter <
=
10 cm
) usu
ally dom
inate size class d
istribu
tions by colon
y n
um
ber (F
ig. 6.5
, Obu
ra in p
rep), b
ut h
ere we often
foun
d
that th
e smallest size classes w
ere a small fraction
of colonies
in m
ost sites. Am
ong geograp
hic areas, th
e high
est absolu
te recru
itmen
t levels were seen
in A
mbod
ivahib
e and
Voh
e-m
ar. Am
ong gen
era, we cou
nted
the h
ighest n
um
ber of th
e sm
allest size classes in th
e most com
mon
genera, i.e. A
cro-pora, m
assive Porites, G
alaxea, Seriatopora, and
Pocillopora
(Fig. 6
.8). H
owever, w
hen
norm
alized by relative ab
un
dan
ce, P
ocillopora show
s the stron
gest recruitm
ent sign
al, with
Seriatopora, F
un
gia, and
both
massive an
d b
ranch
ing P
orites close b
ehin
d (F
ig 6.7
).N
oting th
e pattern
amon
g sites and
genera in
the largest
size classes also allows u
s to discu
ss the p
otential d
isturb
ance
dyn
amics in
the region
. Gen
erally speakin
g, corals in N
orth-
east Mad
agascar span
ned
their exp
ected size class d
istribu
-tion
, inclu
din
g the largest colon
ies expected
for a given
genu
s. Th
e presen
ce of all size classes, especially th
e largest, in
dicates a relatively low
level of catastroph
ic distu
rban
ce over th
e past few
decad
es. Am
ong th
e locations, A
mbod
ivahib
e, L
oky Bay, an
d N
osy An
kao all show
ed d
omin
ance by area of
the largest size classes (n
ote y-axis break in
Fig. 6
.6). V
ohem
ar an
d A
nd
ravina on
the oth
er han
d sh
owed
either low
er dom
i-n
ance or ab
sence of th
e larger size classes (respectively), su
g-gestin
g a pattern
of either on
going d
isturb
ance or a p
revious
severe distu
rban
ce to these coral com
mu
nities. In
the case of
An
dravin
a, this is m
ost likely the h
eavy sedim
ent load
s tole-rated
by these reefs, an
d in
the case of V
ohem
ar, a combin
a-tion
of wave stress from
extreme exp
osure on
the reef crest,
and
anth
ropogen
ic stresses in th
e harb
or. Am
ong gen
era, the
largest colonies w
ere represen
ted by stagh
orn A
cropora thic-
kets, large Acropora tab
les, large stand
s of Galaxea, an
d grou
ps
of platin
g Mon
tipora. Un
like man
y other region
s in th
e Ind
o-P
acifi c, very large massive P
orites “bom
mies” w
ere rare.
Coral Bleaching D
ataTh
e n
ortheast m
onsoon
season of 2
010, from
Janu
ary to M
ay, is the local su
mm
er season w
hen
bleach
ing risk is
high
est. In th
e mon
ths d
irectly prior to ou
r survey, th
e region
of North
east Mad
agascar show
ed a h
igh tem
peratu
re an
omaly, lead
ing to alerts th
rough
the N
OA
A “H
otspot”
program
, and
a “High
Risk W
arnin
g” from C
OR
DIO
/IU
CN
Western
Ind
ian O
cean R
egional B
leachin
g Warn
ing
Netw
ork (CO
RD
IO/IU
CN
March
29, 2
010). Th
e latter
warn
ings, u
pd
ated every 2
weeks, b
egan w
ith a “m
oderate”
warn
ing for N
ortheast M
adagascar from
Janu
ary 15, u
ntil
Feb
ruary 12
, at wh
ich p
oint it w
as up
graded
to a “high
risk” w
arnin
g that ran
un
til March
29. Th
e w
armin
g event ab
ated
as the su
mm
er cond
itions cooled
rough
ly 3-4
weeks early,
and
the stron
g SE m
onsoon
/trade w
ind
s that sign
al the sh
ift in
seasons w
ere already in
force by th
e exped
ition’s start in
the last d
ays of March
. Th
is resulted
in an
early cooling an
d
dissip
ation of th
e regional w
arm p
ool of water.
Over th
e heatin
g period
, sites across the region
accum
u-
lated 4
-7 D
egree Heatin
g Weeks (D
HW
), defi n
ed as th
e n
um
ber of w
eeks over a 12-w
eek win
dow
in w
hich
tem-
peratu
res exceed lon
g-term su
mm
er maxim
a by 1
° C (L
iu
et al. 2005, Stron
g et al. 2002, Skirvin
g et al. 2006). A
s a ru
le of thu
mb, corals are at risk for b
leachin
g with
as little as 1 D
HW
, and
with
mod
erate levels of bleach
ing at 4
DH
W,
and
wid
espread
bleach
ing w
ith m
ortality by 8
DH
W
(Skirving et al. 2
006). O
ur su
rvey began
imm
ediately after
the h
otspot d
issipated
, maxim
izing ou
r chan
ces of fi nd
ing
maxim
al bleach
ing eff ects. A
s our stu
dy region
reached
a m
aximu
m of 7
DH
W im
med
iately before ou
r survey, an
d
previou
s work on
east-coast Malagasy reefs in
2005 h
as sh
own
that con
dition
s of 6 D
HW
correlated w
ith m
ean
bleach
ing rates of 3
8%
(+/- 3
.0 SE
) (McC
lanah
an et al.
2007), w
e expected
wid
espread
bleach
ing, an
d th
e possib
ility of m
ortality.W
e did
observe b
leachin
g throu
ghou
t the ran
ge of the
survey, b
ut th
e overall extent of b
leachin
g was low
(Fig. 6
.9).
Th
e mean
bleach
ing exten
t across all transects w
as 5.1
%
(+/- 1
.0%
se) of colonies aff ected
, with
maxim
a in A
nd
ravina
and
Voh
emar (9
.7%
+/- 4
.0%
, 9.5
% +
/- 2.1
%, resp
ectively, m
ean ±
se). If we con
sider th
e eff ect by colon
y area, the
overall mean
estimate rises to 7
.54%
(+/- 2
.3%
se) of colony
area per 1
00 m
2 prim
arily du
e to the com
mon
occurren
ce of large, p
artially bleach
ed A
cropora thickets (F
ig. 6.10). Th
ese
thickets w
ere particu
larly comm
on in
Loky B
ay, wh
ere there
Chapter 6
62Rapid Assessm
ent Program
Figure 6.5. Coral size distribution by number of colonies counted per transect, grouped by area. In all graphs, size classes are shown in the x axis, the dark
gray for recruit and juvenile sizes (0 - 10 cm diam
eter) and light gray for adult sizes (> 10 cm
diameter). The x axis represents the size classes in cm
- 001-002 is <
2.5 cm; 011-020 is 11-20 cm
, etc.
Figure 6.6. Coral size distribution by colony area surveyed per transect, grouped by area. Note break in y-axis for Ambodivahibe, Loky Bay, and Nosy Ankao,
due to domination in area of vary large colonies.
Coral reef health and status
63 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
is a clear distin
ction am
ong p
er colony an
d p
er area esti-m
ates (Fig. 6
.9, F
ig. 6.1
0).
Wh
ile bleach
ing w
as presen
t throu
ghou
t the area, w
e wit-
nessed
little to no b
leachin
g-ind
uced
mortality, w
ith overall
mean
s of aff ected colon
ies at <1%
in a valu
e statistically in
distin
guish
able from
zero (0.1
6%
+/- 0
.13%
se). Du
ring
our su
rveys in early-m
id A
pril, m
any of th
e colonies n
oted
as “bleach
ed” still retain
ed p
ortions of u
naff ected
tissue, an
d
were likely to recover w
ithou
t mortality. U
pon
return
ing to
Ivovna in
early Jun
e, 2-3
km n
orth of A
mbod
ivahib
e, only
3 of 1
1 tran
sects show
ed an
y bleach
ing at all, th
e overall bleach
ing exten
t was 0
.6%
of colonies aff ected
(+/- 0
.5%
se), an
d n
o bleach
ing-in
du
ced m
ortality was ap
paren
t.Th
e low
level of bleach
ing w
e witn
essed d
urin
g the R
AP
su
rvey occurred
across the w
hole size class d
istribu
tion,
wh
ich d
oes not su
ggest that an
y particu
lar life stage/size class w
as more or less su
sceptib
le to bleach
ing eff ects d
urin
g this
event (F
ig. 6.1
1).
Th
ere were, h
owever, m
ajor distin
ctions in
bleach
-in
g extent am
ong coral gen
era (Fig. 6
.12). M
any of th
ese gen
eric-level pattern
s followed
expectation
s based
on p
re-viou
s suscep
tibility m
easures, w
ith “su
sceptib
le” genera like
Stylophora, Mon
tipora, and
Seriatopora show
ing m
ajor eff ects, an
d m
ore interm
ediate/resistan
t genera like P
avona, G
alaxea, an
d H
ydnophora sh
owin
g more m
inor eff ects. H
owever, cer-
tain p
atterns w
ere more u
nexp
ected. Th
e “m
assive” growth
form
s in gen
era like massive P
orites, Favia, F
avites, Gon
ias-trea, an
d P
latygyra, show
ed greater eff ects th
an th
e bran
ch-
ing form
s of Porites, A
cropora and
Pocillopora, even
thou
gh
these b
ranch
ing gen
era are generally con
sidered
to be m
ore su
sceptib
le to bleach
ing an
d b
leachin
g-ind
uced
mortality. It
is neverth
eless comm
on in
East A
frican reefs to see relatively
high
levels of bleach
ing in
massive P
orites, thou
gh w
ith low
m
ortality, as foun
d h
ere.Th
ese gen
eric-level pattern
s can also h
elp exp
lain th
e dis-
tinction
s in b
leachin
g extent am
ong th
e samp
ling location
s. W
e recorded
the greatest b
leachin
g extent in
An
dravin
a an
d V
ohem
ar, wh
ere 9.5
-9.7
% of colon
ies were aff ected
, w
hile th
e other th
ree sites show
ed 1
.6-5
.0%
of colonies
aff ected (F
igure 6
.9). E
xamin
ing th
e relative abu
nd
ance of
coral genera across location
s (Figu
re 6.5
), we can
see that
both
An
dravin
a and
Voh
emar are d
omin
ated by gen
era th
at show
ed th
e greatest eff ects of bleach
ing. In
An
dravin
a, over 2
0%
of the corals sam
pled
were Stylophora, w
hile in
V
ohem
ar, over 20%
were m
assive Porites. Th
ese tw
o genera
show
ed th
e greatest eff ects of bleach
ing in
our su
rvey, and
th
eir abu
nd
ance in
these p
opu
lations con
tribu
tes to the
bleach
ing p
atterns w
e saw am
ong location
s.
CON
SERVATION
RECOM
MEN
DATIO
NS
Th
e reefs in N
ortheast M
adagascar gen
erally show
very good
coral cover for East A
frica (average 48%
) and
colonies th
at sp
an th
e expected
size distrib
ution
s, inclu
din
g large, old
colonies. Th
e com
mon
occurren
ce of these large colon
ies
suggests th
at this region
has n
ot suff ered
a catastroph
ic mor-
tality in th
e recent p
ast (1-2
decad
es). Th
is gives the area a
high
priority for con
servation, as reef h
ealth, at least in
terms
of the b
enth
ic comm
un
ity) has n
ot yet shifted
far from b
ase-lin
e cond
itions b
efore hu
man
distu
rban
ces becam
e comm
on.
Of th
e fi ve locations, th
ree are heavily sed
imen
t-aff ected
bay system
s: Am
bod
ivahib
e, Loky B
ay and
An
dravin
a. Of
these, th
e fi rst two are large an
d d
eep en
ough
to have areas
with
in th
em th
at are less sedim
ent-aff ected
and
allow for
comp
lex reef structu
re (i.e. A02/A
04 in
Am
bod
ivahib
e, A
06/A
11A
in L
oky Bay) w
hile th
e small size of A
nd
ravina
resulted
in a very h
eavily sedim
ent-aff ected
system w
ith low
coral ab
un
dan
ce and
small corals. Th
e sou
thern
end
of our
survey, V
ohem
ar, has a very exp
osed fore-reef an
d h
eavily sed
imen
t- and
hu
man
-infl u
enced
back-reef. F
inally N
osy A
nkao, on
a seaward
ban
k bath
ed b
y cleaner w
ater and
less in
fl uen
ced b
y sedim
entation
, is also the site of an
inten
sive algal farm
ing op
eration.
Th
e second
hottest th
ermal stress even
t in th
e region, after
the E
l Niñ
o of 1997-9
8 occu
rred d
urin
g the tim
e of the
RA
P su
rveys. Ou
r fi nd
ings of low
bleach
ing levels, <
5%
, su
ggest that th
e corals and
reefs of North
east Mad
agascar are su
rprisin
gly resistant to tem
peratu
re stress. Total aff ected
colon
ies (pale &
bleach
ed) w
ere also low, 9
.9 %
(1.6
SE),
especially as com
pared
to estimates of 4
4.2
% (2
.9 SE
)%
aff ected corals in
the sou
theast, at A
nd
avadoaka an
d 6
2.9
%
(7.4
SE) at B
elo Sur M
er du
ring th
e same p
eriod (Sop
hie
Ben
bow
, Blu
e Ven
tures). W
e also foun
d evid
ence of low
im
pact of th
e 1997-9
8 an
d 2
005 even
ts by th
e abu
nd
ance
of large colonies several d
ecades old
. Th
is pattern
app
ears at sites across th
e region, in
clud
ing A
mbod
ivahib
e, Loky B
ay, an
d N
osy An
kao. W
hile th
e mech
anism
for this ap
paren
t bleach
ing resis-
tance req
uires fu
rther research
, both
the u
pw
elling of cool
water cau
sed b
y fl ow of th
e South
Equ
atorial Cu
rrent on
to th
e coastline an
d its d
efl ection n
orth over th
e tip of M
ada-
gascar, and
of turb
idity cau
sed b
y terrestrial water, are likely
to be sign
ifi cant factors. Th
is fi n
din
g sup
ports th
e notion
th
at the corals of N
ortheast M
adagascar are m
ore resistant
to bleach
ing eff ects th
an oth
er reefs in th
e region. B
oth th
e lack of recen
t bleach
ing in
the face of w
armin
g and
eviden
ce of low
historical levels of m
ajor distu
rban
ce suggest th
at the
reefs of North
east Mad
agascar shou
ld b
e high
priority for
conservation
action.
Alth
ough
the size stru
cture d
ata show
s that th
e reefs have
relatively high
cover and a large p
ercentage of m
ature corals,
coral recruitm
ent is gen
erally low. Th
is m
ay be a matter of
concern
for the lon
g-term overall resilien
ce of these reefs, an
d
shou
ld be p
rime target of m
anagem
ent in
tervention
. Th
e low
levels of recruitm
ent m
ay be du
e to a nu
mber of factors,
inclu
din
g: (1) few sou
rces of coral larvae: most reefs in
this
region h
ave a narrow
and sh
allow reef p
rofi le, limitin
g the
total area of reef comm
un
ity and p
otential sou
rce colonies for
reprod
uction
; (2) low reten
tion of d
ispersin
g larvae: strong
curren
ts that rip
north
ward
aroun
d th
e tip of M
adagascar m
ay deliver d
ispersin
g larvae to the E
ast African
main
land in
stead
Chapter 6
64Rapid Assessm
ent Program
Figure 6.7. Coral size distribution by number of colonies counted for each of the m
ost abundant 9 genera (by number of colonies).
Figure 6.8. Coral size distribution by colony area surveyed for most abundant 9 genera (by area).
Coral reef health and status
65 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Figure 6.9. The extent of coral bleaching by location, expressed as the percentage of colonies affected per 100 m2. Error bars are SE of each condition class
(Normal, Pale, etc.).
Figure 6.10. The extent of coral bleaching by location, expressed as colony area per 100 m2 affected. Error bars are SE of each condition class (Norm
al, Pale, etc.).
Chapter 6
66Rapid Assessm
ent Program
Figure 6.11. The extent of coral bleaching by size class, expressed as number of colonies per 100 m
2 affected. Error bars are SE of each condition class (Norm
al, Pale, etc.).
Figure 6.12. The extent of coral bleaching by genus across the 23 focal genera sampled in our m
ethodology, expressed as number of colonies per 100 m
2 affected. Error bars are SE of each condition class (Norm
al, Pale, etc.), numbers above the bars indicate the num
ber of transects in which the genus was found.
Coral reef health and status
67 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
of retainin
g them
locally; and (3) little good
settlemen
t sub-
strate: most reefs sh
owed
a relatively low abu
ndan
ce of bare, h
ard su
bstrate free of sedim
ent, tu
rf, soft coral or spon
ge.W
hile th
e fi rst two of th
ese factors are not likely to ch
ange
du
e to conservation
actions, th
ese reefs can b
e man
aged to
prom
ote coral recruitm
ent by en
surin
g good settlem
ent su
b-
strate. Th
is can b
e prom
oted b
y a) watersh
ed m
anagem
ent,
to redu
ce sedim
entation
and
smoth
ering of su
rfaces other-
wise su
itable for coral settlem
ent, an
d b
) limitin
g fi shin
g p
ressure, to p
romote h
erbivory an
d red
uction
of comp
eti-tion
betw
een tu
rf/macroalgae an
d recru
iting corals. O
ther
comp
onen
ts of the R
AP
also foun
d sed
imen
t infl u
ence an
d
fi shin
g to be overw
helm
ing an
throp
ogenic factors aff ectin
g th
e region’s reefs, reinforcin
g the n
eed to m
anage b
oth fac-
tors to sustain
the reefs.
In term
s of practical m
anagem
ent action
s therefore, w
e recom
men
d th
e followin
g:1.
limitin
g fi shin
g pressu
re in p
riority sites to main
tain a
strong com
mu
nity of grazin
g fi sh;
2.
man
aging u
pstream
watersh
eds to m
inim
ize sedim
ent
delivery to th
e reefs;3.
mon
itoring fi sh
ing, w
atershed
use an
d coral reef h
ealth
to evaluate th
e eff ectiveness of th
ese man
agemen
t action
s; and
4.
selecting ad
dition
al man
agemen
t actions sp
ecifi c to in
divid
ual sites an
d/or location
s, such
as a.
partn
ership
with
the algae-farm
ing com
pan
y at N
osy An
kao, b.
establish
ing com
mu
nity-b
ased m
anagem
ent areas
in A
mb
odivah
ibe, L
oky Bay an
d A
nd
ravina, an
d
c. estab
lishin
g solid an
d liq
uid
waste an
d
mu
nicip
ally-man
aged p
rotected areas in
V
ohem
ar.
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68Rapid Assessm
ent Program
Appendix 1
Coral species recorded by site, during surveys in northeast M
adagascar
Coral species recorded by site, during surveys in northeast Madagascar. Site num
bers correspond to site codes in Table 1 on page 13, and the map on page
25. Sites followed by an ‘a’ or ‘b’ indicate more than one full survey.
Number
Family/Genus
SpeciesSite code
Acroporidae
1A
croporaabrotan
oides6a,1
1,1
3,1
4,1
5,1
6,1
8,2
1,2
4
2A
croporaacu
leus
6b
3A
croporaappressa
6b,9
,24
4A
croporaau
stera1,2
b,6b,7
,8,1
0,1
1,2
4
5A
croporabran
chi6a,9
,10,1
1,1
3,1
4,1
5,1
6,1
7,1
8,2
2,2
3,2
4
6A
croporacerealis
20,2
4
7A
croporaclathrata
5,6
b,8,1
0,1
1,1
3,1
4,1
5,1
8
8A
croporacopiosa
1,2
a,2b,6
a,11,2
0
9A
croporacytherea
2a,1
4,1
6,1
7,2
4
10
Acropora
digitifera2a,1
1,2
0
11
Acropora
divaricata1,2
b,4,5
,6a,6
b,8,1
0,1
1,1
3,1
4,1
5,2
0,2
1,2
3,2
4
12
Acropora
fl orida2b,6
a,6b,8
,9,1
0,1
1,1
3,1
5,1
6,2
4
13
Acropora
gemm
ifera1,2
b,6a,6
b,11,2
0,2
3
14
Acropora
grandis
2a,1
1,1
3,1
8,2
3
15
Acropora
granulosa
1,2
a,2b,4
,5,6
a,6b,7
,8,1
0,1
1,1
6,1
7,1
8,2
1
16
Acropora
hum
ilis1,4
,6b,9
,11,1
4,2
0,2
2,2
3
17
Acropora
hyacinthu
s2b,5
,6a,6
b,7,8
,9,1
0,1
1,1
3,1
6,1
7
18
Acropora
interm
edia5,6
a,11,1
6,1
8,2
3
19
Acropora
latistella1,2
a,2b,4
,5,6
a,6b,7
,8,9
,10,1
1,1
6,1
7,1
8,2
4
20
Acropora
loripes2a
21
Acropora
lutken
i2a,5
,6a,6
b,9,1
6,2
3
22
Acropora
microclados
5,8
23
Acropora
microphthalm
a6b
24
Acropora
muricata
6a,6
b,8,1
1,2
0
25
Acropora
nan
a11,2
4
26
Acropora
nasu
ta11
27
Acropora
retusa
6a,1
1,2
0,2
2,2
3,2
4
table contin
ued on
next page
Coral species recorded by site, during surveys in northeast Madagascar
69 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
28
Acropora
robusta
11,1
3,1
6,1
8
29
Acropora
rosaria2a,2
b,6b
30
Acropora
samoen
sis1,2
a,2b,4
,5,6
a,6b,8
,9,1
1,2
1,2
3,2
4
31
Acropora
secale2a,2
b,4,6
b,7,8
,20,2
4
32
Acropora
squarrosa
5,7
,9,2
1
33
Acropora
subu
lata1,2
b,5,6
b,21
34
Acropora
tenuis
1,2
b,4,5
,6a,6
b,8,1
0,1
1,1
4,1
6,2
4
35
Acropora
valida2a,5
,8,1
1,1
4,1
7,1
8,2
0,2
2
36
Acropora
verweyi
2a,2
1
37
Acropora
zp2a,5
,6a,6
b,8,9
,10,1
1,1
3,1
7,1
8,2
1
38
Acropora
zp4,5
,6a,6
b,8,9
,10,1
1,1
6
39
Acropora
zp1,2
a,14,1
5
40
Acropora
zp-bu
lbous ac
1,2
b,4,2
0,2
2
41
Acropora
zp-hu
milis 2
6a,6
b,7
42
Acropora
zp1
16,1
8
43
Acropora
zp1
-Mayotte
22,2
3,2
4
44
Acropora
zpA
-long rc
2a
45
An
acroporapillai
16
46
Astreopora
expansa
8,1
3,1
4,1
5,1
6,1
8,2
1,2
2,2
3
47
Astreopora
listeri2b,8
,9,1
0
48
Astreopora
myriophthalm
a1,2
b,4,5
,6a,6
b,8,9
,10,1
4,1
5,2
1,2
2,2
3,2
4
49
Astreopora
ocellata8,9
,10
50
Astreopora
randalli
2a,2
b,5,7
51
Astreopora
suggesta
2a,1
7
52
Isoporapalifera
2b,4
,5,6
a,6b,1
0,1
1,1
3,1
5,1
6,2
0,2
1,2
3,2
4
53
Mon
tiporaaequ
itubercu
lata1,6
a,7,1
3,1
8,2
0,2
3,2
4
54
Mon
tiporacalcarea
5,1
7,2
1
55
Mon
tiporacryptu
s6a
56
Mon
tiporadan
ae20
57
Mon
tiporaeffl orescen
s1,6
b,23,2
4
58
Mon
tiporaeff u
sa5,6
b,21
59
Mon
tiporafl ow
eri1,2
b,8,1
7
60
Mon
tiporafoveolata
1,4
,5,8
,10,1
1,1
3,1
4,2
1,2
2
61
Mon
tiporagrisea
6b,1
6,1
8,2
1,2
3
62
Mon
tiporain
formis
1,2
a,2b,4
,6a,6
b,7,1
3,2
1,2
3
63
Mon
tiporam
onasteriata
2b,4
,5,6
a,6b,7
,8,9
,10,1
1,1
3,1
5,1
6,1
8,2
1
64
Mon
tiporan
odosa6b,2
1
65
Mon
tiporaspon
godes1
66
Mon
tiporastilosa
2a,6
b
67
Mon
tiporatu
berculosa
1,2
b,4,5
,6a,8
,9,1
0,1
4,1
7,2
1,2
3
68
Mon
tiporatu
rgescens
1,2
b,14
69
Mon
tiporaun
data2a,2
b,4,5
,6a,6
b,7,8
,9,1
0,1
1,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
2,2
3,2
4
Appendix 1
70Rapid Assessm
ent Program
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
70
Mon
tiporaven
osa5
71
Mon
tiporaverru
cosa2a,2
b,5,6
b,9,1
0,1
1,1
5,1
6,1
7,2
2,2
4
72
Mon
tiporazp
2b,1
7,2
1
Agariciidae
73
Gardin
eroserisplan
ulata
2b,6
b,7,1
7,2
3
74
Leptoseris
foliosa6a,6
b,7,1
0,1
3,1
4,1
5,1
6
75
Leptoseris
glabra2a,4
,7,1
6,1
7,2
4
76
Leptoseris
hawaiien
sis8,1
0,1
4,1
5,1
7
77
Leptoseris
incru
stans
1,2
a,6a,6
b,15,1
6,1
7,1
8
78
Leptoseris
mycetoseroides
1,2
b,4,6
a,6b,8
,9,1
0,1
1,1
6,1
7
79
Leptoseris
scabra1,2
a
80
Leptoseris
solida2a
81
Leptoseris
yabei2a
82
Pachyseris
speciosa1,2
a,2b,4
,6a,6
b,7,8
,9,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
3,2
4
83
Pavon
acactu
s1,1
1,1
4
84
Pavon
aclavu
s1,2
a,6b,7
,20,2
1,2
4
85
Pavon
adecu
ssata11,1
4,2
1
86
Pavon
adu
erdeni
6a,1
6,2
1,2
3,2
4
87
Pavon
aexplan
ulata
2b,5
,6a,6
b,7,9
,10,1
1,1
3,1
4,1
5,1
6,1
7,1
8
88
Pavon
afron
difera11,1
4
89
Pavon
am
aldivensis
6b,7
,10,1
3,1
4,1
7,2
1,2
4
90
Pavon
avarian
s1,2
b,4,6
a,6b,8
,9,1
8,2
1,2
3
91
Pavon
aven
osa1,2
b,4,6
a,6b,8
,9,1
0,1
1,2
0,2
1,2
3,2
4
92
Pavon
azp
21
Astrocoeniidae
93
Stylocoeniella
armata
1,2
b,4,6
b,10,2
1
94
Stylocoeniella
guen
theri2a,4
,6a
Dendrophylliidae
95
Heteropsam
mia
cochlea18
96
Tubastrea
micran
tha21
97
Tubastrea
zpp
2b,6
a,6b,7
,9,1
0,1
8,2
1
98
Turbin
ariairregu
laris4,5
,8,1
0,1
4,1
8,2
4
99
Turbin
ariam
esenterin
a4,5
,8,9
,13,1
4,1
5,1
6,1
7,1
8,2
1,2
2,2
4
10
0T
urbin
ariapeltata
5,8
,9,1
0,1
7,1
8
10
1T
urbin
ariaren
iformis
8,1
5,1
7,1
8,2
2
10
2T
urbin
ariastellu
lata5,6
a,9,1
0,1
4,1
5,1
7
10
3T
urbin
ariazp
8,9
,10,2
2
Euphyllidae
104
Physogyra
lichtenstein
i1,2
a,5,6
a,6b,7
,9,1
0,1
7,2
0,2
1,2
3,2
4
10
5P
lerogyrasin
uosa
1,2
a,2b,4
,6a,6
b,7,9
,10,2
3
Faviidae
106
Barabattoia
amicoru
m2a,6
b,17
Coral species recorded by site, during surveys in northeast Madagascar
71 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
10
7C
aulastrea
conn
ata8,1
6,1
7
10
8C
yphastreachalcidicu
m2b,5
,13,1
5,2
1,2
3
10
9C
yphastream
icrophthalma
2b,4
,5,8
,9,1
0,1
3,1
4,1
5,1
8,2
1
11
0C
yphastreaserailia
13,2
1,2
2
11
1C
yphastreazp
A5
11
2D
iploastreaheliopora
2b,4
,6a,6
b,8
11
3E
chinopora
cf-small
14,1
5
11
4E
chinopora
forskaliana
6b,1
6
11
5E
chinopora
gemm
acea1,2
a,2b,4
,5,6
a,6b,7
,8,1
0,1
1,1
3,1
4,1
6,2
1
11
6E
chinopora
hirsutissim
a1,2
b,4,5
,6a,6
b,7,8
,9,1
0,1
1,1
3,1
4,1
5,1
6,1
8,2
0,2
1,2
2,2
3,2
4
11
7E
chinopora
robusta
4,5
,6b,7
,10,1
3,1
7
11
8Favia
danae
1,2
a,5,9
,10
11
9Favia
favus
2a,2
b,5,6
a,6b,7
,8,9
,10,1
1,1
4,1
5,1
6,1
7,2
2
12
0Favia
helianthoides
1,2
a,5,6
a,11,1
3,1
4,1
5,1
6,1
8,2
2,2
4
12
1Favia
lizardensis
1,2
b,4,6
a,6b,8
,9,1
0,1
3,1
4,1
6,1
8,2
0,2
1,2
2,2
3
12
2Favia
maritim
a6a,6
b,7,8
,10,1
3,1
4,1
6,1
7,1
8,2
2
12
3Favia
matthai
4,6
a,6b,7
,8,1
4,2
1
12
4Favia
maxim
a21
12
5Favia
pallida2b,5
,6b,7
,8,1
0,1
1,1
4,2
4
12
6Favia
rotum
ana
16,1
7,1
8
12
7Favia
rotun
data1,2
b
12
8Favia
speciosa2b,4
,6b,8
,10,1
3,1
4,1
6,2
0,2
1,2
2,2
3,2
4
12
9Favia
stelligera2a,2
b,4,6
b,13,2
1,2
3,2
4
13
0Favia
trun
catus
1,2
b,4,5
,6a,6
b,22
13
1Favia
veroni
9,1
0,1
3,1
5,1
7,2
2
13
2Favites
abdita4,5
,6a,1
1,1
3,1
4,1
5,1
8,2
3
13
3Favites
acuticolis
14,1
8,2
1
13
4Favites
chinen
sis4,1
6
13
5Favites
complan
ata1,2
a,2b,6
b,16,2
1,2
4
13
6Favites
fl exuosa
1,2
a,2b,4
,6a,6
b,9,1
0,1
6,2
1,2
2,2
3
13
7Favites
halicora6b,2
1,2
4
13
8Favites
parafl exuosa
2b,1
0,2
1,2
4
13
9Favites
pentagon
a1,5
,6a,8
,9,1
0,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
2,2
3,2
4
14
0Favites
russelli
6b,7
,8,9
,13,1
5,1
6,1
7,1
8,2
1,2
2,2
3,2
4
14
1Favites
vasta2a,4
,5,7
,9,2
2,2
3
14
2G
oniastrea
aspera21,2
3
14
3G
oniastrea
australen
sis2a,8
,24
14
4G
oniastrea
deformis
5,9
,15,1
7
14
5G
oniastrea
edwardsi
2a,2
b,4,5
14
6G
oniastrea
min
uta
5,9
,10
14
7G
oniastrea
palauen
sis9,2
1,2
2
14
8G
oniastrea
pectinata
1,2
a,2b,4
,5,6
a,6b,7
,9,1
0,1
1,2
0,2
1,2
3,2
4
Appendix 1
72Rapid Assessm
ent Program
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
14
9G
oniastrea
peresi1,2
b,4,5
,7,8
,9,1
0,1
1,1
5,1
7,2
1,2
2,2
3,2
4
15
0G
oniastrea
retiformis
5,6
b,21,2
4
15
1L
eptastreaaequ
alis2a,2
b,4,5
,6b,8
,9,1
0,1
3,1
4,1
8,2
1,2
2,2
3,2
4
15
2L
eptastreapru
inosa
4,6
a,9,1
0,1
3,1
4,1
5,1
6
15
3L
eptastreapu
rpurea
2a,6
a,6b,2
1
15
4L
eptastreatran
sversa2a,1
6,2
1
15
5L
eptoriaphrygia
2a,2
b,6a,9
,11,1
3,1
4,1
5,1
6,1
8,2
4
15
6M
ontastrea
ann
uligera
6b
15
7M
ontastrea
curta
2a,6
a,16,2
2,2
3,2
4
15
8M
ontastrea
magn
istellata2a,6
b,17
15
9M
ontastrea
serageldini
1,2
a,2b,5
16
0O
ulophyllia
benettae
15,1
7,2
2
16
1O
ulophyllia
crispa1,2
a,2b,4
,5,6
a,6b,8
,9,1
0,1
1,1
3,1
4,1
6,1
7,2
0,2
2
16
2P
latygyraacu
ta9,1
5
16
3P
latygyracarn
osus
14
16
4P
latygyracon
torta13,1
4,1
6
16
5P
latygyracrosslan
di1,2
b,5,6
a,6b,9
,10,1
6
16
6P
latygyradaedalea
1,2
a,2b,4
,5,6
b,7,8
,9,1
0,1
1,1
5,1
6,1
7,2
3
16
7P
latygyralam
ellina
2a,2
b,6a,6
b,23
16
8P
latygyrapin
i1,2
b,5,6
b,8,9
,10,1
6,1
7,1
8,2
0,2
2,2
4
16
9P
latygyraryu
kyuen
sis2a,2
b,6a,9
,14,1
5,2
2,2
4
17
0P
latygyrasin
ensis
1,2
a,2b,4
,5,6
a,6b,1
0,1
1,1
3,1
4,1
6
17
1P
latygyraverw
eyi2a,4
,5,6
a,6b,7
,8,1
3,1
5,1
6
17
2P
lesiastreadevan
tieri4,6
a,6b,8
,9,1
0,1
1,1
4,2
1,2
2
17
3P
lesiastreaversipora
4,5
,6b,9
,13,1
4,1
6,1
7,1
8,2
2,2
4
Fungiidae
174
Cycloseris
costulata
2a,7
,8,1
0
17
5C
ycloserispatelliform
is2a,5
17
6F
un
giacon
cina
1,2
a,2b,6
a,6b,1
1,2
4
17
7F
un
giacoron
a6b
17
8F
un
giadan
ai1,2
a,4,6
a,6b,7
,11
17
9F
un
giafu
ngites
1,2
b,4,6
a,6b,7
,8,1
0,1
1,1
3,1
4,1
6,2
0,2
1,2
4
18
0F
un
giagran
ulosa
1,2
a,2b,6
b,7,8
,9,1
0,1
1,1
3,2
2,2
4
18
1F
un
giapau
moten
sis2a,4
,8,9
,10
18
2F
un
giarepan
da1,4
,6a,6
b,7,9
,11,1
3,1
4,2
0,2
4
18
3F
un
giascabra
1,2
4
18
4F
un
giascru
posa20
18
5F
un
giascu
taria5,9
18
6F
un
giaseychellen
sis1,2
b
18
7H
erpolithalim
ax1,2
b,6a,7
,11
18
8San
dalolitharobu
sta2a
Coral species recorded by site, during surveys in northeast Madagascar
73 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
M
eandrinidae
189
Gyrosm
iliain
terrupta
2a,5
,6b,7
,8,9
,10,1
6,2
2
M
erulinidae
190
Hydn
ophoraexesa
1,2
a,5,6
a,6b,7
,8,1
0,1
1,1
4,1
5,1
6,1
7,2
0,2
1,2
2,2
3,2
4
19
1H
ydnophora
pilosa8,1
0,1
3,1
5,1
6,1
7
19
2M
erulin
aam
pliata1,2
b,4,6
a,6b,8
,9,1
1,2
0,2
1,2
3,2
4
M
ussidae
193
Acan
thastreabrevis
24
19
4A
canthastrea
echinata
1,2
a,2b,5
,6a,6
b,8,9
,10,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
3,2
4
19
5A
canthastrea
faviaformis
1
19
6A
canthastrea
hemprichii
2b,4
,5,6
a,6b,1
0,1
3,1
8
19
7A
canthastrea
ishigakiensis
2a,6
b
19
8A
canthastrea
regularis
14
19
9A
canthastrea
rotun
dofl ora15,1
6,1
8
20
0A
canthastrea
subechin
ata2a,9
,10,1
1
20
1A
canthastrea
zp1
15
20
2B
lastomussa
merletti
2a,2
b,6a,6
b,10,1
4,1
5,1
6,1
7,1
8
20
3C
ynarin
alachrym
alis6a,7
20
4L
obophylliacorym
bosa2b,5
,7,2
4
20
5L
obophylliafl abelliform
is2a,1
3
20
6L
obophylliahataii
6a,6
b,8,1
0,1
7
20
7L
obophylliahem
prichii1,2
a,2b,4
,6a,6
b,7,9
,10,1
1,1
4,1
6,1
7,2
0,2
1,2
2,2
4
20
8L
obophylliapachysepta
5,1
7
20
9L
obophylliarobu
sta2a,6
b,7,1
6,1
7,2
1,2
3
21
0M
icromussa
amaku
sensis
12
21
1Scolym
iavitien
sis2a
21
2G
alaxeaastreata
1,2
a,2b,4
,6a,6
b,7,1
1,2
0,2
1,2
3
21
3G
alaxeafasicu
laris1,2
a,2b,4
,5,6
a,6b,7
,8,9
,10,1
1,1
3,1
4,1
5,1
6,2
0,2
1,2
2,2
3,2
4
Pectiniidae
214
Echin
ophylliaaspera
6a,6
b,8,1
0,1
1,1
3,1
4,1
6,1
8,2
1,2
4
21
5E
chinophyllia
echinata
2a,2
b,5,6
a,6b,8
,9,1
0,1
3,1
5,1
6,1
7,1
8,2
3,2
4
21
6E
chinophyllia
orpheensis
14
21
7M
ycedium
elephantotu
s1,2
a,2b,6
a,7,8
,9,1
0,1
3,1
4,1
5,2
4
21
8M
ycedium
man
caoi2a,2
b,7,2
0,2
2,2
4
21
9M
ycedium
um
bra2a
22
0O
xyporaglabra
2a,6
b,21,2
4
22
1O
xyporalacera
1,2
b,4,6
a,6b,7
,16,1
7,2
3
22
2O
xyporazf-rou
gh4
22
3P
ectinia
africanus
2a,6
b,7,8
,9,1
0,1
3,1
7
Pocilloporidae
224
Pocillopora
damicorn
is1,2
a,2b,4
,6b,8
,9,1
2,1
4,1
8,2
0,2
1,2
2,2
3
22
5P
ocilloporadan
ai20
Appendix 1
74Rapid Assessm
ent Program
continued
table contin
ued on
next page
Number
Family/Genus
SpeciesSite code
22
6P
ocilloporaeydou
xi1,2
a,2b,4
,5,6
a,6b,8
,9,1
0,1
1,1
2,1
3,1
4,1
5,1
6,1
8,2
0,2
1,2
3,2
4
22
7P
ocilloporain
diania
2a,2
b,6b,2
0,2
1
22
8P
ocilloporam
eandrin
a20
22
9P
ocilloporaverru
cosa1,2
a,2b,4
,5,6
a,6b,8
,10,1
1,1
4,1
5,1
8,2
0,2
1,2
3
23
0Seriatopora
caliendru
m1,2
a,2b,4
,6b,1
1,1
6,2
0,2
1
23
1Seriatopora
dentritica
7,1
6
23
2Seriatopora
hystrix1,2
a,2b,4
,5,6
a,6b,8
,11,1
3,1
5,1
6,2
0,2
1,2
4
23
3Seriatopora
stellata2b,7
,20,2
3,2
4
23
4Stylophora
danai
6a,1
5,1
6,1
8
23
5Stylophora
madagascaren
sis2b,1
5,1
6,1
8,2
1,2
3
23
6Stylophora
pistillata4,5
,8,1
0,1
1,1
3,1
4,1
6,1
7,1
8,2
2
23
7Stylophora
subseriata
6a,1
3,1
8
23
8Stylophora
zp13,1
5
Poritidae
239
Alveopora
daedelea2a,2
b,4,5
,6a,6
b,8,1
0,1
1,1
3,1
5,1
7
24
0A
lveoporaspon
giosa2a
24
1A
lveoporatizardi
2a,1
0,1
3,1
6
24
2G
oniopora
albiconus
9,1
8
24
3G
oniopora
colum
na
1,5
,6a,9
,10,2
4
24
4G
oniopora
djiboutien
sis2a,5
,6a,8
,9,1
0,1
7,2
4
24
5G
oniopora
lobata2a,4
,5,6
a,6b,1
1,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
2,2
3,2
4
24
6G
oniopora
planulata
16
24
7G
oniopora
somalien
sis4,6
a,7,8
,10,1
4,1
5,1
6,1
7,2
1,2
2,2
4
24
8G
oniopora
stokesi1,2
b,4,5
,7,8
,9,1
3,1
8,2
0,2
1,2
2,2
4
24
9G
oniopora
stutchbu
ryi2a,4
,6b,7
,9,1
0,1
3,1
5,2
2
25
0G
oniopora
zp.
1
25
1P
oritesan
nae
14
25
2P
oritescylin
drica1,2
a,2b,4
,6b,7
,8,9
,11,1
3,1
4,1
5,1
6
25
3P
oriteslatistella
16
25
4P
oriteslichen
9,1
3,1
5,1
6,1
7,1
8
25
5P
oriteslobata
2b,4
,6a,8
,9,1
0,1
1,1
3,1
4,1
6,2
3
25
6P
oriteslu
tea2a,2
b,4,5
,6a,6
b,8,9
,10,1
1,1
3,1
4,1
5,1
6,1
7,1
8,2
0,2
1,2
2,2
4
25
7P
oritesm
onticu
losa6b
25
8P
oritesn
igrescens
6b,2
2,2
3
25
9P
oritesprofu
ndu
s1,2
a,4,5
,6a,6
b,8,1
0,1
1,1
3,1
4,2
0,2
1,2
2,2
3,2
4
26
0P
oritesru
s1,2
a,2b,4
,6a,6
b,13,1
4,1
5,1
6,1
8,2
0,2
3,2
4
26
1P
oritesstephen
soni
6a
26
2P
oriteszp
-fi ne prof ??
5
Siderastreidae
263
An
omastrea
irregularis
17,1
8
26
4C
oscinaraea
colum
na
1,2
b,4,5
,8,1
1,2
2,2
4
26
5C
oscinaraea
crassa1,2
b,4,5
,6a,6
b,8,9
,10,1
3,1
4,1
5,2
0,2
1,2
2,2
3,2
4
Coral species recorded by site, during surveys in northeast Madagascar
75 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
continued Num
berFam
ily/GenusSpecies
Site code
26
6C
oscinaraea
mcn
eilli1,2
a,2b,5
,6b,7
,8,9
,13,1
4,1
5,1
6,1
8,2
1,2
2,2
4
26
7C
oscinaraea
mon
ile1,2
a,2b,4
,6a,6
b,7,8
,9,1
0,1
3,1
4,1
5,1
6,1
7,1
8,2
1,2
2
26
8C
oscinaraea
wellsi
2a,6
b,7,1
4,1
6,1
7
26
9C
raterasteralaevis
16,1
7,2
4
27
0H
orastreain
dica4,5
,8,1
0,1
7,1
8
27
1P
samm
ocoraalbopicta
6a,1
7
27
2P
samm
ocoracon
tigua
12,1
3,1
4,2
0
27
3P
samm
ocoraexplan
ulata
6a,1
7
27
4P
samm
ocoran
iestraazi2b,6
b,7,9
,13,1
8,2
1
27
5P
samm
ocoraprofu
ndacella
5,6
a,6b,8
,9,1
0,1
3,1
4,1
6,1
8,2
0,2
1,2
2,2
4
Non-scleractinia
276
Tubipora
musicu
m1,2
b,11,1
5,2
0,2
2,2
3
27
7M
illeporadichotom
a4,1
8
27
8M
illeporaexesa
2b,5
,8,1
0,1
1,1
2,1
5,2
0,2
1,2
2,2
3,2
4
27
9M
illeporaplatyphylla
2b,4
,5,6
a,6b,9
,10,1
1,1
2,1
5,2
0,2
1,2
2,2
3,2
4
28
0M
illeporaten
ella1,2
b,4,6
b,20,2
1,2
4
28
1M
illeporazp
-dark, coarse2b
76Rapid Assessm
ent Program
Appendix 2
Coral species lists from northeast
Madagascar, M
adagascar and the w
estern Indian Ocean
table contin
ued on
next page
Coral species lists from northeast M
adagascar, Madagascar and the western Indian Ocean. W
IO – western Indian Ocean (from Veron & Turak 2005). RAP1
– combined listed of the fi rst CI m
arine RAP in 2002 (Veron & Turak 2005), indicated by a “1”, and species reported for Madagascar but not recorded in the
surveys “p”. RAP3 – this study. Note – species for which further work is necessary, with respect to ID-identifi cation.
Family
GenusSpecies
AuthorityW
IORAP1
RAP3Note
AcroporidaeA
cropora
abrotanoides
(Lam
arck 1816)
11
1
aculeu
s(D
ana 1
846)
11
1
acum
inata
(Verrill 1
864)
1ID
anthocercis
(Brook 1
893)
11
ID
appressa(E
hren
berg 1
834)
11
1
arabensis
(Hod
gson &
Carp
enter 1
995)
11
ID
aspera(D
ana 1
846)
1ID
austera
(Dan
a 1846)
11
1
bifurcata
(Nem
enzo 1
971)
1
branchi
(Riegl 1
995)
11
1
brueggem
ann
i(B
rook 1893)
1P
cerealis(D
ana 1
846)
11
1
cf(valencien
nesi
(Miln
e Ed
ward
s & H
aime 1
860)
11
ID
clathrata(B
rook 1891)
11
1
copiosa(N
emen
zo 1967)
11
1
cytherea(D
ana 1
846)
11
1
digitifera(D
ana 1
846)
11
1
divaricata(D
ana 1
846)
11
1
donei
(Veron
& W
allace 1984)
1ID
echinata
(Dan
a 1846)
1P
florida(D
ana 1
846)
11
1
muricata/form
osa(D
ana 1
846)
11
1
forskali(E
hren
berg 1
834)
11
ID
gemm
ifera(B
rook 1892)
11
1
glauca
(Brook 1
893)
1P
grandis
(Brook 1
892)
11
1
granulosa
(Miln
e Ed
ward
s & H
aime 1
860)
1P
1
haimei
(Miln
e Ed
ward
s & H
aime 1
860)
1
hemprichii
(Eh
renberg 1
834)
11
ID
horrida(D
ana 1
846)
11
ID
hum
ilis(D
ana 1
846)
11
1
hyacinthu
s1
indon
esia(W
allace 1997)
1ID
inerm
is(B
rook 1891)
1P
insign
is(N
emen
zo 1967)
11
ID
Coral species lists from northeast M
adagascar, Madagascar and the w
estern Indian Ocean
77 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
continued
Family
GenusSpecies
AuthorityW
IORAP1
RAP3Note
irregularis
(Brook 1
892)
11
ID
lamarcki
(Veron
2000)
1P
latistella(B
rook 1891)
11
1
longicyathu
s(M
ilne E
dw
ards &
Haim
e 1860)
11
ID
loripes(B
rook 1892)
11
1
lutken
i1
ID
macrostom
a(B
rook 1891)
11
ID
massaw
ensis
(Maren
zeller 1906)
1ID
microclados
1ID
microphthalm
a(V
errill 1859)
11
1
millepora
(Eh
renberg 1
834)
1
mirabilis
(Qu
elch 1
886)
1P
mon
ticulosa
(Brü
ggeman
n 1
879)
1P
nan
a(Stu
der 1
878)
1P
1
nasu
ta(D
ana 1
846)
11
1
natalen
sis(R
eigl 1995)
11
ID
nobilis
(Dan
a 1846)
11
1
ocellata(K
lun
zinger 1
879)
11
ID
panicu
lata(V
errill 1902)
1ID
papillarae(L
atypov 1
992)
1ID
parilis(Q
uelch
1886)
1ID
pharaonis
(Miln
e Ed
ward
s & H
aime 1
860)
1P
pingu
is(W
ells 1950)
11
ID
plantagin
ea(L
amarck 1
816)
1P
polystoma
(Brook 1
891)
11
ID
prostrata(D
ana 1
846)
1ID
pulchra
(Brook 1
891)
11
ID
rambleri
(Bassett-Sm
ith 1
890)
1
retusa
(Dan
a 1846)
11
1
robusta
(Dan
a 1846)
11
1
rosaria(D
ana 1
846)
1P
1
roseni
(Wallace 1
999)
1P
samoen
sis(B
rook 1891)
11
1
secale(Stu
der 1
878)
11
1
selago(Stu
der 1
878)
1ID
squarrosa
(Eh
renberg 1
834)
11
1
stoddarti(P
illai & Sch
eer 1976)
11
ID
striata(V
errill 1866)
1P
subu
lata(D
ana 1
846)
11
1
tenuis
(Dan
a 1846)
11
1
un
described1
ID
valida(D
ana 1
846)
11
1
variabilis(K
lun
zinger 1
879)
11
ID
variolosa(K
lun
zinger 1
879)
11
ID
vermicu
lata(N
emen
zo 1967)
11
ID
verweyi
(Veron
& W
allace 1984)
11
1
willisae
(Veron
& W
allace 1984)
11
ID
yongei
(Veron
& W
allace 1984)
11
ID
zp1
ID
zp1
ID
zp1
ID
zp-bu
lbous ac
1ID
zp-hu
milis 2
1ID
zp1
1ID
Appendix 2
78Rapid Assessm
ent Program
table contin
ued on
next page
continued Fam
ilyGenus
SpeciesAuthority
WIO
RAP1RAP3
Notezp
1-M
ayotte1
ID
zpA
-long rc
1ID
An
acroporaforbesi
(Rid
ley 1884)
1P
pillai1
ID
Astreopora
expansa
(Brü
ggeman
n 1
877)
1P
1
listeri(B
ernard
1896)
11
1
myriophthalm
a(L
amarck 1
816)
11
1
ocellata(B
ernard
1896)
11
1
randalli
1
suggesta
(Wells 1
954)
1P
1
Isoporacu
neata
(Dan
a 1846)
1P
palifera(L
amarck 1
816)
11
1
Mon
tiporaaequ
itubercu
lata(B
ernard
1897)
11
1
australien
sis(B
ernard
1897)
1P
calcarea(B
ernard
1897)
11
1
caliculata
(Dan
a 1846)
1ID
cryptus
1ID
danae
(Miln
e Ed
ward
s & H
aime 1
851)
11
1
digitata(D
ana 1
846)
11
ID
efflorescens
(Bern
ard 1
897)
11
1
eff usa
(Dan
a 1846)
11
floweri
(Wells 1
954)
1P
1
foliosa(P
allas 1766)
11
ID
foveolata1
friabilis(B
ernard
1897)
11
ID
grisea(B
ernard
1897)
11
1
hispida(D
ana 1
846)
1
inform
is(B
ernard
1897)
11
1
kellyi(V
eron 2
000)
1P
lobulata
(Bern
ard 1
897)
1P
mean
drina
(Eh
renberg 1
834)
1ID
millepora
(Crossl&
1952)
1P
mollis
(Bern
ard 1
897)
11
ID
mon
asteriata(F
orskäl 1775)
11
1
zf.nodosa
(Dan
a 1846)
11
orientalis
(Nem
enzo 1
967)
1P
peltiformis
(Bern
ard 1
897)
11
ID
spongodes
(Bern
ard 1
897)
11
spum
osa(L
amarck 1
816)
1P
stilosa(E
hren
berg 1
834)
11
tubercu
losa(L
amarck 1
816)
11
1
turgescen
s(B
ernard
1897)
11
1
un
data(B
ernard
1897)
11
1
venosa
(Eh
renberg 1
834)
11
1
verrucosa
(Lam
arck 1816)
11
1
zp1
ID
AgariciidaeC
oeloserism
ayeri(V
augh
an 1
918)
1P
Gardin
eroseriszf.plan
ulata
(Dan
a 1846)
11
1
Leptoseris
explanata
(Yab
e & Su
giyama 1
941)
11
foliosa(D
ineson
1980)
11
1
glabra1
ID-taxon
note
hawaiien
sis(V
augh
an 1
907)
11
1
incru
stans
(Qu
elch 1
886)
11
1
mycetoseroides
(Wells 1
954)
11
1
papyracea(D
ana 1
846)
1P
Coral species lists from northeast M
adagascar, Madagascar and the w
estern Indian Ocean
79 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
continued
Family
GenusSpecies
AuthorityW
IORAP1
RAP3Note
scabra(V
augh
an 1
907)
11
1
solida(Q
uelch
1886)
1P
1
yabei(P
illai & Sch
eer 1976)
1P
1
Pachyseris
rugosa
(Lam
arck 1801)
11
synon
ymyzed
speciosa(D
ana 1
846)
11
1
Pavon
abipartita
(Nem
enzo 1
980)
11
ID
cactus
(Forskål 1
775)
1P
1
clavus
(Dan
a 1846)
11
1
decussata
(Dan
a 1846)
11
1
duerden
i(V
augh
an 1
907)
11
1
explanulata
(Lam
arck 1816)
11
1
frondifera
(Lam
arck 1816)
11
1
maldiven
sis(G
ardin
er 1905)
11
1
varians
(Verrill 1
864)
11
1
venosa
(Eh
renberg 1
834)
11
1
zp1
ID
AstrocoeniidaeM
adraciskirbyi
(Veron
& P
ichon
1976)
1
zp.
()1
Stylocoeniella
armata
(Eh
renberg 1
834)
11
1
guen
theri(B
assett-Smith
1890)
11
1
DendrophylliidaeH
eterocyathus
aequicostatu
s(M
ilne E
dw
ards &
Haim
e 1848)
1
Heteropsam
mia
cochlea(Sp
engler 1
781)
11
1
Tubastrea
micran
tha1
zpp
1
Turbin
ariafron
dens
(Dan
a 1846)
11
ID
irregularis
( Bern
ard)
1
mesen
terina
(Lam
arck 1816)
11
1
peltata(E
sper 1
794)
11
1
reniform
is(B
ernard
1896)
11
1
stellulata
(Lam
arck 1816)
11
1
un
described1
1ID
-taxon n
ote
EuphilliidaeC
atalaphylliajardin
ei(Saville-K
ent 1
893)
11
1
Euphyllia
glabrescens
(Ch
amisso &
Eysen
hard
t 1821)
1P
1
Physogyra
lichtenstein
i(M
ilne E
dw
ards &
Haim
e 1851)
11
1
Plerogyra
sinuosa
(Dan
a 1846)
11
1
FaviidaeB
arabattoiaam
icorum
(Miln
e Ed
ward
s & H
aime 1
850)
11
Cau
lastreacon
nata
(Ortm
ann
1892)
11
1
furcata
(Dan
a 1846)
1P
tum
ida(M
atthai 1
928)
1
Cyphastrea
chalcidium
(Forskål 1
775)
11
1
microphthalm
a(L
amarck 1
816)
11
1
serailea(F
orskål 1775)
11
1
zp.
()1
ID
Diploastrea
heliopora(L
amarck 1
816)
11
1
Echin
oporacf-sm
all1
forskaliana
(Miln
e Ed
ward
s & H
aime 1
850)
11
1
fruticu
losa(E
hren
berg 1
834)
1P
gemm
acea(L
amarck 1
816)
11
1
hirsutissim
a(M
ilne E
dw
ards &
Haim
e 1849)
11
1
lamellosa
(Esp
er 1795)
11
mam
miform
is(N
emen
zo 1959)
1P
robusta
1
Favia
danae
(Verrill 1
872)
11
1
Appendix 2
80Rapid Assessm
ent Program
table contin
ued on
next page
continued
Family
GenusSpecies
AuthorityW
IORAP1
RAP3Note
favus
(Forskål 1
775)
11
1
helianthoides
(Wells 1
954)
11
1
laxa(K
lun
zinger 1
879)
11
ID
lizardensis
(Veron
& P
ichon
1977)
11
1
maritim
a(N
emen
zo 1971)
11
1
matthai
(Vau
ghan
1918)
11
1
maxim
a1
ID
pallida(D
ana 1
846)
11
1
rotum
ana
(Gard
iner 1
899)
11
1
rotun
data1
ID-taxon
note
speciosa(D
ana 1
846)
11
1
stelligera(D
ana 1
846)
11
1
trun
catus
(Veron
2000)
11
1
veroni
(Moll &
Borel-B
est 1984)
11
1
Favites
abdita(E
llis & Sol&
er 1786)
11
1
acuticolis
1ID
chinen
sis(V
errill 1866)
11
1
complan
ata(E
hren
berg 1
834)
11
1
flexuosa
(Dan
a 1846)
11
1
halicora(E
hren
berg 1
834)
11
1
micropen
tagona
(Veron
2000)
1ID
paraflexuosa
(Veron
2000)
1P
1
pentagon
a(E
sper 1
794)
11
1
russelli
(Wells 1
954)
11
1
spinosa
(Klu
nzin
ger 1879)
11
vasta(K
lun
zinger 1
879)
11
1
Gon
iastreaaspera
(Verrill 1
905)
11
1
australen
sis(M
ilne E
dw
ards &
Haim
e 1857)
11
1
colum
ella(C
rossl& 1
948)
1P
deformis
1ID
edwardsi
(Ch
evalier 1971)
11
1
min
uta
(Veron
2000)
1P
1
palauen
sis(Y
abe &
Sugiyam
a 1936)
1P
1
pectinata
(Eh
renberg 1
834)
11
1
peresi(Fau
re & P
ichon
1978)
11
1
retiformis
(Lam
arck 1816)
11
1
thecata(V
eron D
eVan
tier & T
urak 2
000)
1
Leptastrea
aequalis
(Veron
2000)
11
1
bottae(M
ilne E
dw
ards &
Haim
e 1849)
11
pruin
osa(C
rossl& 1
952)
11
purpu
rea(D
ana 1
846)
11
1
transversa
(Klu
nzin
ger 1879)
11
1
Leptoria
phrygia(E
llis & Sol&
er 1786)
11
1
Mon
tastreaan
nuligera
(Miln
e Ed
ward
s & H
aime 1
849)
1P
1
coleman
i(V
eron 2
000)
1P
curta
(Dan
a 1846)
11
1
magn
istellata(C
hevalier 1
971)
11
1
serageldini
(Veron
2000)
11
1
valencien
nesi
(Miln
e Ed
ward
s & H
aime 1
848)
11
ID
Oulophyllia
benn
ettae(V
eron P
ichon
)1
ID
crispa(L
amarck 1
816)
11
1
Platygyra
acuta
(Veron
2000)
11
1
carnosu
s(V
eron 2
000)
1P
1
contorta
1
Coral species lists from northeast M
adagascar, Madagascar and the w
estern Indian Ocean
81 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
continued Fam
ilyGenus
SpeciesAuthority
WIO
RAP1RAP3
Notecrosslan
di(M
atthai 1
928)
11
1
daedalea(E
llis & Sol&
er 1786)
11
1
lamellin
a(E
hren
berg 1
834)
11
1
pini
(Ch
evalier 1975)
11
1
ryukyu
ensis
(Yab
e & Su
giyama 1
936)
11
1
sinen
sis(M
ilne E
dw
ards &
Haim
e 1849)
11
1
verweyi
(Wijsm
an-B
est 1976)
11
Plesiastrea
devantieri
(Veron
2000)
11
versipora(L
amarck 1
816)
11
1
zun
know
nzp
.gen
us/sp
ecies un
describ
ed1
FungiidaeC
antharellu
sdoederlein
i(M
arenzeller 1
907)
1
Cycloseris
costulata
(Ortm
ann
1889)
11
1
curvata
(Hoeksem
a 1989)
11
ID
cyclolites(L
amarck 1
801)
11
ID
erosa(D
öderlein
1901)
11
ID
patelliformis
(Bosch
ma 1
923)
1P
1
sinen
sis(M
ilne E
dw
ards &
Haim
e 1851)
1
somervillei
(Gard
iner 1
909)
11
ID
tenuis
(Dan
a 1846)
11
ID
vaughan
i(B
oschm
a 1923)
11
ID
Diaseris
distorta(M
ichelin
1843)
1P
fragilis(A
lcock 1893)
1P
Fun
giacon
cinn
a(V
errill 1864)
11
1
corona
(Död
erlein 1
901)
11
danai
(Miln
e Ed
ward
s & H
aime 1
851)
11
1
fun
gites(L
inn
eaus 1
758)
11
1
granulosa
(Klu
nzin
ger 1879)
11
1
horrida(D
ana 1
846)
11
ID
klun
zingeri
(Död
erlein 1
901)
11
ID
molu
ccensis
(Horst 1
919)
11
ID
paum
otensis
(Stutch
bu
ry 1833)
11
1
puishan
i(V
eron &
DeV
antier 2
000)
1
repanda
(Dan
a 1846)
11
1
scabra(D
öderlein
1901)
11
1
scruposa
(Klu
nzin
ger 1879)
11
1
scutaria
(Lam
arck 1801)
11
1
seychellensis
(Hoeksem
a 1993)
11
1
Halom
itrapileu
s(L
inn
aeus 1
758)
11
Herpolitha
limax
(Hou
ttuyn
1772)
11
1
weberi
(Horst 1
921)
11
synon
ymyzed
Podabacia
crustacea
(Pallas 1
766)
11
motu
porensis
(Veron
1990)
11
Polyphyllia
talpina
(Lam
arck 1801)
11
Sandalolitha
robusta
1
Meandrinidae
Cten
ellachagiu
s(M
atthai 1
928)
1
Gyrosm
iliain
terrupta
(Eh
renberg 1
834)
11
1
Merulinidae
Hydn
ophoraexesa
(Pallas 1
766)
11
1
microcon
os(L
amarck 1
816)
11
1ID
-taxon n
ote
rigida(D
ana 1
846)
1P
Meru
lina
ampliata
(Ellis &
Sol&er 1
786)
11
1
scabricula
(Dan
a 1846)
11
ID
Mussidae
Acan
thastreabrevis
(Miln
e Ed
ward
s & H
aime 1
849)
11
1
echinata
(Dan
a 1846)
11
1
faviaformis
1
hemprichii
(Eh
renberg 1
834)
11
1
Appendix 2
82Rapid Assessm
ent Program
table contin
ued on
next page
continued Fam
ilyGenus
SpeciesAuthority
WIO
RAP1RAP3
Notehillae
(Wells 1
955)
11
ID
ishigakiensis
(Veron
1990)
11
1
lordhowen
sis(V
eron &
Pich
on 1
982)
11
ID
regularis
1
rotun
dofl ora1
subechin
ata1
zp1
1ID
Blastom
ussa
merleti
(Wells 1
961)
11
un
described1
ID
Cyn
arina
lacrymalis
(Miln
e Ed
ward
s & H
aime 1
848)
11
1
Lobophyllia
corymbosa
(Forskål 1
775)
11
1
fl abelliformis
1ID
hataii(Y
abe &
Sugiyam
a 1936)
11
1
hemprichii
(Eh
renberg 1
834)
11
1
pachysepta1
ID
robusta
(Yab
e & Su
giyama 1
936)
11
1
Microm
ussa
zf.amaku
sensis
(Veron
1990)
11
Scolymia
zf.vitiensis
(Brü
ggeman
n 1
877)
11
1
Symphyllia
agaricia(M
ilne E
dw
ards &
Haim
e 1849)
11
erythraea(K
lun
zinger 1
879)
1
recta(D
ana 1
846)
11
valencien
nesii
(Miln
e Ed
ward
s & H
aime 1
849)
1P
OculinidaeG
alaxeaastreata
(Lam
arck 1816)
11
1
fascicularis
(Lin
naeu
s 1767)
11
1
PectinidaeE
chinophyllia
aspera(E
llis & Sol&
er 1788)
11
1
echinata
(Saville-Ken
t 1871)
11
1
orpheensis
(Veron
& P
ichon
1980)
11
1
Mycediu
melephatotu
s(P
allas 1766)
11
1
man
caoi(N
emen
zo 1979)
11
1
Pectiniidaeum
bra1
PectinidaeO
xyporacrassispin
osa(N
emen
zo 1979)
11
ID-taxon
note
glabra(N
emen
zo 1959)
11
lacera(V
errill 1864)
11
1
Pectin
iaafrican
us
(Veron
2000)
11
1
lactuca
(Pallas 1
766)
1P
PocilloporidaeP
ocilloporaan
keli(Sch
eer & P
illai 1974)
1ID
damicorn
is(L
inn
aeus 1
758)
11
1
danai
1ID
eydouxi
(Miln
e Ed
ward
s & H
aime 1
860)
11
1
fun
giformis
(Veron
2000)
1P
indian
ia(V
eron 2
000)
11
1
mean
drina
1ID
verrucosa
(Ellis &
Sol&er 1
786)
11
1
zf.kelleheri( V
eron 2
000)
1ID
Seriatoporaacu
leata(Q
uelch
1886)
1ID
caliendru
m(E
hren
berg 1
834)
11
1
dentritica
1
guttatu
s(V
eron 2
000)
1
hystrix(D
ana 1
846)
11
1
stellata1
ID
un
described1
ID
Stylophoradan
ai1
madagascaren
sis(V
eron 2
000)
11
1
pistillata(E
sper 1
797)
11
1
subseriata
(Eh
renberg 1
834)
11
1
Coral species lists from northeast M
adagascar, Madagascar and the w
estern Indian Ocean
83 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
continued Fam
ilyGenus
SpeciesAuthority
WIO
RAP1RAP3
Notew
ellsi(Sch
eer 1964)
1P
zp1
ID
PoritidaeA
lveoporaallin
gi(H
off meister 1
925)
11
ID
daedalea(F
orskål 1775)
11
1
excelsa(V
errill 1863)
1ID
fenestrata
(Lam
arck 1816)
11
ID
tizardi(B
assett-Smith
1890)
11
1
un
described1
ID
zf.spongiosa
(Dan
a 1846)
11
1
Gon
ioporaalbicon
us
(Veron
2000)
11
1
burgosi
(Nem
enzo 1
955)
11
ID
colum
na
(Dan
a 1846)
11
1
djiboutien
sis(V
augh
an 1
907)
11
1
lobata(M
ilne E
dw
ards &
Haim
e 1860)
11
1
min
or(C
rossl& 1
952)
11
ID
planulata
(Eh
renberg 1
834)
11
1
savignyi
(Dan
a 1846)
1ID
somalien
sis(V
augh
an 1
907)
11
1
stokesi(M
ilne E
dw
ards &
Haim
e 1851)
11
1
stutchbu
ryi(W
ells 1955)
11
tenuiden
s(Q
uelch
1886)
11
ID
zf.tenella
(Qu
elch 1
886)
1ID
zp.
1ID
Porites
ann
ae(C
rossl& 1
952)
11
attenuata
(Nem
enzo 1
955 )
ID
australien
sis(V
augh
an 1
918)
11
ID
colum
naris
(Klu
nzin
ger 1879)
1ID
cum
ulatu
s(N
emen
zo 1955)
1ID
cylindrica
(Dan
a 1846)
11
1
deformis
(Nem
enzo 1
955)
1ID
densa
(Vau
ghan
1918)
1ID
echinulata
(Klu
ngzin
ger 1879)
11
ID
everman
ni
(Vau
ghan
1907)
1ID
flavus
(Veron
2000)
1ID
horizontalata
(Hoff m
eister 1925)
1ID
latistellata(Q
uelch
1886)
11
1
lichen(D
ana 1
846)
11
1
lobata(D
ana 1
846)
11
1
lutea
(Miln
e Ed
ward
s & H
aime 1
851)
11
1
mayeri
(Vau
ghan
1918)
11
ID
mon
ticulosa
(Dan
a 1846)
11
1
murrayen
sis(V
augh
an 1
918)
11
ID
napopora
(Veron
2000)
1ID
negrosen
sis(V
eron 1
990)
1ID
nigrescen
s(D
ana 1
846)
11
1
ornata
(Nem
enzo 1
971)
1ID
profun
dus
(Reh
berg 1
892)
11
1
rugosa
(Fen
ner &
Veron
2000)
1ID
rus
(Forskål 1
775)
11
1
silliman
iana
(Nem
enzo 1
976)
11
ID
solida(F
orskäl 1775)
11
ID
somalien
sis(G
ravier 1911)
11
ID
stephenson
i(C
rossl& 1
952)
11
tubercu
losa(V
eron 2
000)
1ID
vaughan
i(C
rossl& 1
952)
1ID
Appendix 2
84Rapid Assessm
ent Program
continued Fam
ilyGenus
SpeciesAuthority
WIO
RAP1RAP3
Notezp.1
()1
ID
zp.2()
1ID
zp-fi n
e prof??1
ID
Poritipora
paliformis
(Veron
2000)
11
Stylaraeapu
nctata
(Lin
neau
s 1758)
1P
SiderastreidaeA
nom
astraeairregu
laris(M
arenzeller 1
901)
1P
1
Coscin
araeacolu
mn
a(D
ana 1
846)
11
1
crassa(V
eron &
Pich
on 1
980)
1P
1
mon
ile(F
oskål 1775)
11
1
un
described1
ID
wellsi
(Veron
& P
ichon
1980)
11
1
zp.m
cneilli
()1
1ID
-taxon n
ote
Craterastera
laevis1
new
Horastrea
indica
(Pich
on 1
971)
11
1
Psam
mocora
albopicta1
new
contigu
a(E
sper 1
797)
11
1
digitata(M
ilne E
dw
ards &
Haim
e 1851)
1P
synon
ymyzed
explanulata
(Horst 1
922)
11
1
haimean
a(M
ilne E
dw
ards &
Haim
e 1851)
11
synon
ymyzed
nierstraszi
(Horst 1
921)
1P
1
obtusan
gula
(Lam
arck 1816)
1P
synon
ymyzed
profun
dacella(G
ardin
er 1898)
11
1
superfi
cialis(G
ardin
er 1898)
11
synon
ymyzed
zf.vaughan
i(Y
abe &
Sugiyam
a 1936)
1syn
onym
yzed
Pseudosiderastrea
tayami
(Yab
e & Su
giyama 1
935)
1P
Siderastreasavign
yana
(Miln
e Ed
ward
s & H
aime 1
850)
1P
TrachyphyllidaeT
rachyphylliageoff royi
(Au
dou
in 1
826)
11
85A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Appendix 3
List of coral species collected for taxonom
y/genetic analyses
table contin
ued on
next page
GenusSpecies
#
Acropora
cf. arabensis?
1
abrotanoides
1
appressa1
austera
1
branchi
1
cf cerealis/nasu
ta1
cf. arabensis/open
1
cf. fi ne/verw
eyi, roun
ded corallites1
cf. hum
ilis/gemm
ifera1
cf. stag-interm
edia?2
cf.hum
ilis 21
cf.raspy staghorn1
cf.scaly1
copiosa12
cytherea/irregularis (robu
st)1
digitifera1
digitifera/samoen
sis1
divaricata1
fl orida1
gemm
ifera7
grandis
1
hum
ilis1
hum
ilis/retusa
1
hyacinthu
s/subu
lata1
latistella1
loripes/granulosa
1
lutken
i1
mon
ticulosa
1
muricata
13
nan
a1
GenusSpecies
#
nasu
ta1
retusa
1
rosaria1
samoen
sis1
secale1
sp (sam
/dig?)1
sp-blu
e1
sp-M
ayotte1
sp. 1
squarrosa
1
tenuis
1
valida?2
valida/squarrosa?
1
valida/variabilis/appressa1
variabilis??1
An
omastrea
irregularis
6
Blastom
ussa
merletti
6
Coscin
araea
mon
ile15
mon
ile/crassa?1
sp1
sp. (old nam
e mcn
eilli?)4
wellsi
2
Echin
opora
hirsutissim
a2
robusta/forskalian
a (large)2
sp (sm
all)4
Favites
acu
ta2
pentagon
a17
Galaxea
astreata
4
fascicularis
4
Tentative species names and num
ber of fragments collected of each species.
Appendix 3
86Rapid Assessm
ent Program
GenusSpecies
#
Gyrosm
iliain
terrupta
4
Horastrea
indica
5
Isoporapalifera
2
Leptoseris
explanulata
2
foliosa3
incru
stans
2
mycetoseroides
2
Sp (in
cl. Craterastrea laevis)
9
sp (sol)
1
Mycediu
m
cf.um
bra?1
man
caoi1
Pavon
a
(blank)
1
cactus
10
decussata
4
duerden
i1
explanulata
1
frondifera
5
maldiven
sis2
Platygyra
carnosu
s??1
sp (verw
eyi??)1
sp (verw
eyi/ryukyu
sensis??)
2
Plesiastrea
devan
tieri2
versipora1
Pocillopora
damicorn
is1
eydouxi
4
indian
ia1
mean
drina
1
verrucosa
1
verrucosa (B
)1
Psam
mocora
albopicta2
explanulata
1
nierstrazi (fi n
e)3
profun
dacella16
Seriatopora
hystrix1
stellata1
Stylophora
cf.danai
4
madagascaren
sis5
pistillata22
pistillata (mordax)
2
Turbin
aria
mesen
terina
1
peltata1
sp1 (irregu
laris)3
Total sam
ples
28
0
continued
87A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Appendix 4
Reef fi sh species list, northeast M
adagascar
Record column indicates when species have been observed in M
adagascar: 2010 = new record for M
adagascar as seen during RAP 3. 2002 = observed by
Allen in 2002 in NW M
adagascar; * = new record for M
adagascar reported by Allen from NW
Madagascar (M
cKenna and Allen 2005). P = previously recorded
for Madagascar (collated by Allen 2005). Cf. =
identifi cation uncertain, looks similar to the nam
ed Pacifi c species.
SPECIESRECORD
TO
RP
ED
INID
AE
Torpedo fu
scomacu
lata Peters, 1
855
2002
DASYATIDIDAE
Taen
iura lym
ma (F
orsskål, 1775)
2002
Taen
iura sp.
2010
MYLIOBATIDAE
Rhin
optera javanica M
üller &
Hen
le, 1841
2010
MURAENIDAE
Gym
nothorax javan
icus (B
leeker, 1865)*
2002
HOLOCENTRIDAE
Myripristis adu
sta Bleeker, 1
853
2002
M. bern
dti Jordan
& E
verman
n, 1
902
2002
M. violacea B
leeker, 1851
2002
N. sam
mara (F
orsskål, 1775)
2002
Sargocentron
caudim
aculatu
m (R
üp
pell, 1
835)
2002
S. spiniferu
m (F
orsskål, 1775)
2002
AULOSTOMIDAE
Aulostom
us chin
ensis (L
inn
aeus, 1
766)
2002
SCORPAENIDAE
Pterois volitan
s (Lin
naeu
s, 1758)
2010
PLATYCEPHALIDAE
Pap
illoculicep
s longicep
s (Eh
renberg, 1
829)
2002
SERRANIDAE
Aethaloperca rogaa (F
orsskål, 1775)
2002
An
yperodon leu
cogramm
icus (V
alencien
nes, 1
828)
2002
Cephalopholis argu
s Bloch
& Sch
neid
er, 1801
2002
SPECIESRECORD
C. m
iniata (F
orsskål, 1775)
2002
C. u
rodeta subsp. n
igripinn
is (Forster 1
801)
2002
C. sexm
aculata R
üp
pell, 1
828
2002
Epin
ephelus caeru
leopun
ctatus (B
loch, 1
790)
2002
E. fasciatu
s (Forsskål, 1
775)
2002
E. fu
scoguttatu
s (Forsskål, 1
775)
2002
E. m
ultin
otatus (P
eters, 1876)
P
Gram
mistes sexlin
eatus (Th
ü
nberg, 1
792)
2002
Plectropom
us laevis (L
acepèd
e, 1802)
2002
P. pun
ctatus Q
uoy &
Gaim
ard, 1
824
2002
Variola lou
ti (Forsskål, 1
775)
2002
CIRRHITIDAE
Paracirrhites forsteri (Sch
neid
er, 1801)
2002
PRIACANTHIDAE
Priacan
thus ham
rur (F
orsskål, 1775)
2002
CARANGIDAE
Caran
goides ferdau (F
orsskål, 1775)
2002
C. fu
lvoguttatu
s (Forsskål, 1
775)
P
Caran
x melam
pygus C
uvier, 1
833
2002
C. papu
ensis A
lleyne &
Macleay, 1
877
2002
Elegatis bipin
nulata (Q
uoy &
Gaim
ard, 1
824)
P
Scomberoides lysan
(Forsskål, 1
775)
P
Selar crum
enophthalm
us (B
loch, 1
793)
2002
LUTJANIDAE
Aprion
virescens V
alencien
nes, 1
830
2002
Lutjan
us argen
timacu
latus (F
orsskål, 1775)
2002
table contin
ued on
next page
Appendix 4
88Rapid Assessm
ent Program
table contin
ued on
next page
Fish, continued
SPECIESRECORD
L. bohar (F
orsskål, 1775)
2002
L. ehren
burgi (P
eters, 1869)*
2002
L. fu
lvifl amm
a (Forsskål, 1
775)
2002
L. fu
lvus (Sch
neid
er, 1801)
2002
L. gibbu
s (Forsskål, 1
775)
2002
L. kasm
ira (Forsskål, 1
775)
2002
L. m
onostigm
a (Cu
vier, 1828)
2002
L. rivu
latus (C
uvier, 1
828)
2002
Macolor n
iger (Forsskål, 1
775)
2002
CAESIONIDAE
Caesio caeru
laurea L
acepèd
e, 1802
2002
C. lu
naris C
uvier, 1
830
2002
C. xan
thonota B
leeker, 1853
2002
Pterocaesio chrysozon
a (Cu
vier, 1830)
2010
P. marri Sch
ultz, 1
953
2010
P. pisang (B
leeker, 1853)*
2002
P. tile (Cu
vier, 1830)
2002
HAEMULIDAE
Diagram
ma pictu
m (Th
ü
nberg, 1
792)
2002
Plectorhin
chus albovittatu
s (Rü
pp
ell, 1838)
2010
P. gaterinus (F
orsskål, 1775)
2002
P. gibbosus (L
acepèd
e, 1802)
P
P. plagiodesmus (F
owler, 1
935)
P
P. schotaf (Forsskål, 1
775)
P
LETHRINIDAE
Gn
athodentex au
rolineatu
s Lacep
ède, 1
802
2002
Gym
nocran
ius griseu
s (Tem
min
ck & Sch
legel, 1843)
2010
L. harak (F
orsskål, 1775)
2002
L. len
tjan (L
acepèd
e, 1802)
2002
L. m
ahsena (F
orsskål, 1775)
2002
L. m
icrodon V
alencien
nes, 1
830
2002
L. n
ebulosu
s (Forsskål, 1
775)
2002
L. obsoletu
s (Forsskål, 1
775)
2002
L. olivaceou
s Valen
cienn
es, 1830
2002
Mon
otaxis grandocu
lis (Forsskål, 1
775)
2002
NEMIPTERIDAE
Scolopsis bimacu
latus R
üp
pell, 1
828
2002
S. gahnam
(Forsskål, 1
775)
2002
MULLIDAE
Mulloidichthys fl avolin
eatus (L
acepèd
e, 1802)
2002
M. van
icolensis (V
alencien
nes, 1
831)
2010
Paru
peneu
s barberinus (L
acepèd
e, 1801)
2002
P. bifasciatus (L
acepèd
e, 1801)
2002
P. cyclostomus (L
acepèd
e, 1802)
2002
SPECIESRECORD
P. heptacanthu
s (Lacep
ède, 1
801)
2002
P. pleurostigm
a (Ben
nett, 1
830)
2002
P. rubescen
s (Lacep
ède, 1
801)
2002
PEMPHERIDAE
Pem
pheris vanicolen
sis Cu
vier, 1831
2002
KYPHOSIDAE
Kyphosu
s cinerascen
s (Forsskål, 1
775)
2002
K. vaigien
sis (Qu
oy & G
aimard
, 1825)
P
CHAETODONTIDAE
Chaetodon
auriga F
orsskål, 1775
2002
C. blackbu
rni (D
esjardin
s 1836)
2002
C. falcu
la Bloch
, 1793
2002
C. gu
ttatissimus B
enn
ett, 1823
2002
C. in
terruptu
s (Ah
l 1923)
2002
C. klein
ii Bloch
, 1790
2002
C. leu
copleura P
layfair, 1867
2010
C. lin
eolatus C
uvier, 1
831
2002
C. lu
nula L
acepèd
e, 1803
2002
C. m
adagaskariensis A
hl, 1
923
2002
C. m
elann
otus Sch
neid
er, 1801
2002
C. m
eyeri Schn
eider, 1
801
2002
C. trifascialis Q
uoy &
Gaim
ard, 1
824
2002
C. trifasciatu
s Park, 1
797
2002
C. vagabu
ndu
s Lin
naeu
s, 1758
2002
C. xan
thocephalus B
enn
ett, 1832
2002
C. zan
zibarensis P
layfair, 1867
2002
Forcipiger fl avissim
us Jord
an &
McG
regor, 1898
2002
F. longirostris (B
rousson
et, 1782)
2010
Hem
itaurichthys zoster (B
enn
ett, 1831)
2002
Hen
iochus acu
min
atus (L
inn
aeus, 1
758)
2002
H. m
onoceros C
uvier, 1
831
2002
POMACANTHIDAE
Apolem
ichthys trimacu
latus (L
acepèd
e, 1831)
2002
Cen
tropyge bispinosa (G
ün
ther, 1
860)
2002
C. m
ultispin
is (Playfair, 1
867)
2002
Pom
acanthu
s chrysuru
s Cu
vier, 1831
2002
P. imperator (B
loch, 1
787)
2002
P. semicircu
latus C
uvier, 1
831
2002
POMACENTRIDAE
Abu
defduf sexfasciatu
s Lacep
ède, 1
802
2002
A. sordidu
s (Forsskål, 1
775)
2002
A. sparoides (C
uvier, 1
830)
2002
A. vaigien
sis (Qu
oy & G
aimard
, 1825)
2002
Reef fish species list, northeast Madagascar
89 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
Fish, continued
SPECIESRECORD
Am
blyglyphidodon leu
cogaster (Bleeker, 1
847)
Am
phiprion akallopisos (B
leeker, 1853)
2002
A. latifasciatu
s Allen
, 1972
2002
Chrom
is atripectoralis Welan
der &
Schu
ltz, 1951*
2002
C. cf. leu
cura G
ilbert, 1
905
2010
C. dim
idiata (Klu
nzin
ger, 1871)
2002
C. lepidolepis B
leeker, 1877
2002
C. n
igroanalis R
and
all, 1989*
2002
C. n
igrura Sm
ith, 1
960
2002
C. opercu
laris (Gü
nth
er, 1867)
2002
C. tern
atensis (B
leeker, 1856)
2002
C. viridis (C
uvier, 1
830)
2002
C. w
eberi Fow
ler & B
ean, 1
928
2002
Chrysiptera biocellata (Q
uoy &
Gaim
ard, 1
824)
2002
Dascyllu
s aruan
us (L
inn
aeus, 1
758)
2002
D. carn
eus F
ischer, 1
885
2002
D. trim
aculatu
s (Rü
pp
ell, 1928)
2002
Neoglyphidodon
melas (C
uvier, 1
830)
2002
Neopom
acentru
s azysron (B
leeker, 1877)
2002
Plectroglyphidodon
dickii (Lién
ard, 1
839)
2002
P. johnston
ianus F
owler &
Ball, 1
924*
2002
Pom
acentru
s baenschi A
llen, 1
991*
2002
P. caeruleu
s Qu
oy & G
aimard
, 1825
2002
P. caeru
leopun
ctatus (A
llen in
press)
2002
P. cf. javanicu
s Allen
, 1991
2010
P. cf. similis A
llen, 1
991
2010
P. pavo (Bloch
, 1787)
2002
P. sulfu
reus K
lun
zinger, 1
871
2002
P. trilineatu
s Cu
vier, 1830
2002
P. lacrymatu
s (Qu
oy & G
aimard
, 1824)
2002
Pristotis obtu
sirostris (Gu
nter 1
862)
2010
Stegastes fasciolatus (O
gilby, 1
889)
2002
S. nigrican
s (Lacep
ède, 1
802)
2002
Stegastes sp.12010
LABRIDAE
An
ampses caeru
leopun
ctatus R
üp
pell, 1
828
2002
A. geographicu
s Valen
cienn
es, 1840
A. lin
eatus R
and
all, 1972*
2002
A. m
elanuru
s Bleeker, 1
857
2010
A. m
eleagrides Valen
cienn
es, 1840
2002
Bodian
us an
thioides (Ben
nett, 1
831)
2002
B. axillaris (B
enn
ett, 1831)
2002
B. bilu
nulatu
s Lacep
ède, 1
801)
2002
SPECIESRECORD
B. dian
a (Lacep
ède, 1
802)
2002
B. m
acrouru
s (Lacep
ède, 1
801)
P
Cheilin
us chloru
rus (B
loch, 1
791)
2002
C. fasciatu
s (Bloch
, 1791)
2002
C. oxycephalu
s (Bleeker, 1
853)
2002
C. trilobatu
s Lacep
ède, 1
802
2002
C. u
ndu
latus R
üp
pell, 1
835
2002
Coris cau
dimacu
la (Qu
oy & G
aimard
, 1834)
2002
C. cu
vieri (Ben
nett, 1
831)
2002
C. freiri (P
layfair & G
ün
ther, 1
867)
2010
Epibu
lus in
sidiator (Pallas, 1
770)
2002
Gom
phosus caeru
leus L
acepèd
e, 1801
2002
Halichoeres cosm
etus R
and
all & Sm
ith, 1
982*
2002
H. doliatu
s (Lacep
ède, 1
801)*
2002
H. du
ssum
ieri (Valen
cienn
es,1839)
2002
H. hortu
lanus (L
acepèd
e, 1802)
2002
H. m
arginatu
s (Rü
pp
ell, 1835)
2002
Hem
igymn
us m
elapterus (B
loch, 1
791)
2002
H. n
ebulosu
s (Valen
cienn
es, 1839)
2002
H. scapu
laris (Ben
nett, 1
832)
2002
H. zeylon
icus (B
enn
ett, 1833)*
2002
Hem
igymn
us fasciatu
s (Bloch
, 1792)
2002
Hologym
nosu
s ann
ulatu
s (Lacep
ède, 1
801)
2002
Labrichthys u
nilin
eatus (G
uich
enot, 1
847)
2002
Labrichthys sp.1
2010
Labroides bicolor F
owler &
Bean
, 1928
2002
L. dim
idiatus (V
alencien
nes, 1
839)
2002
Macropharyn
godon vivien
ae Ran
dall 1
978*
2002
Macropharyn
godon cf. m
elas2010
Notolabru
s cf. parilus (R
ichard
son, 1
850)
2010
Novacu
lichthys taenioru
s (Lacep
ède, 1
802)
2002
Oxycheilin
us bim
aculatu
s (Valen
cienn
es, 1840)
2002
O. digram
mus (L
acepèd
e, 1801)
2002
O. m
entalis (R
üp
pell, 1
828)
2002
Pseu
docheilinus evan
idus Jord
an &
Everm
ann
, 1902
2002
P. hexataenia (B
leeker, 1857)
2002
Pseu
docheilinus sp.
2010
Pseu
dojuloides cf. atavai R
and
all & R
and
all, 1981
2010
P.cerasinus (Sn
yder, 1
904)
2002
Pteragogu
s fl agellifera (Valen
cienn
es, 1839)
2002
Pteragogu
s sp.2010
Stethojulis albovittata (B
onn
aterre, 1785)
2002
S. bandan
ensis (B
leeker, 1851)
2010
Appendix 4
90Rapid Assessm
ent Program
Fish, continued
SPECIESRECORD
S. strigiventer (B
enn
ett, 1832)
2002
Th alassom
a amblycephalu
m (B
leeker, 1856)
2002
T. hardw
icke (Ben
nett, 1
828)
2002
T. hebraicu
m (L
acepèd
e, 1801)
2002
T. lu
nare (L
inn
aeus, 1
758)
2002
Th alassom
a sp.12010
SCARIDAE
Calotom
us carolin
us (V
alencien
nes, 1
839)
2002
C. spin
idens (Q
uoy &
Gaim
ard, 1
824)
P
Cetoscaru
s bicolor (Rü
pp
ell, 1828)
2002
Chloru
rus atrilu
nula R
and
all & B
ruce, 1
983*
2002
C. cyan
escens (V
alencien
nes, 1
840)
2002
C. sordidu
s (Forsskål, 1
775)
2002
C. stron
gylocephalus (B
leeker, 1862)
2002
Hipposcaru
s harid (Forsskål, 1
775)
2002
Scarus cau
dofasciatus (G
ün
ther, 1
862)
2002
S. falcipinn
is (Playfair, 1
867)
2002
S. frenatu
s Lacep
ède, 1
802
2002
S. ghobban F
orsskål, 1775
2002
S. niger F
orsskål, 1775
2002
S. prasiognathu
s Valen
cienn
es, 1840
2010
S. psittacus F
orsskål, 1775
2002
S. rubroviolaceu
s Bleeker, 1
849
2002
S. russelli V
alencien
nes, 1
840*
2002
S. scaber Valen
cienn
es, 1840
2002
S. tricolor Bleeker, 1
849
2002
S. virdifurcatu
s (Smith
, 1956)*
2002
MICRODESM
IDAE
Nem
ateleotris magn
ifi ca Fow
ler, 1938
2010
Ptereleotris evides (Jord
an &
Hu
bbs, 1
925)
2010
EPHIPPIDAE
Platax teira (F
orsskål, 1775)
2002
SIG
AN
IDA
E
Siganus corallin
us (V
alencien
nes, 1
835)
2010
S. luridu
s (Rü
pp
ell, 1829)
2002
S. cf. spinus (L
inn
aeus, 1
758)
2010
S. stellatus (F
orsskål, 1775)
2002
S. sutor (V
alencien
nes, 1
835)
2002
Siganus sp.
2010
ACANTHURIDAE
A. blochii V
alencien
nes, 1
835
2002
A. du
ssum
ieri Valen
cienn
es, 1835
2002
SPECIESRECORD
A. gu
ttatus F
orster, 1801
2010
A. leu
costernon
Ben
nett, 1
832
2002
A. lin
eatus (L
inn
aeus, 1
758)
2002
A. m
ata (Cu
vier, 1829)
2002
A. n
igricaudu
s Du
ncker &
Moh
r, 1929
2002
A. n
igrofuscu
s (Forsskål, 1
775)
2002
A. ten
nen
ti Gü
nth
er, 1861
2002
A. thom
psoni (F
owler, 1
923)*
2002
A. triostegu
s (Lin
naeu
s, 1758)
2002
A. xan
thopterus V
alencien
nes, 1
835
2002
Cten
ochaetus bin
otatus R
and
all, 1955*
2002
C striatu
s (Qu
oy & G
aimard
, 1824)
2010
C. tru
ncatu
s Ran
dall &
Clem
ents 2
001
2002
Naso an
nulatu
s (Qu
oy & G
aimard
, 1825)
2002
N. brachycen
tron (V
alencien
nes, 1
835)
2002
N. brevirostris (V
alencien
nes, 1
835)
2002
N. fagen
i Morrow
, 1954
2010
N. hexacan
thus (B
leeker, 1855)*
2002
N. litu
ratus F
orster 1801
2002
N. tu
berosus L
acepèd
e, 1802
P
N. u
nicorn
is (Forsskål, 1
775)
2002
N. vlam
ingii (V
alencien
nes, 1
835)
2002
Zebrasom
a desjardinii (B
enn
ett, 1836)
2002
Z. gem
matu
m (V
alencien
nes, 1
835)
P
Z. scopas (C
uvier, 1
829)
2002
Z. veliferu
m (B
loch, 1
795)
2010
SPHYRAENIDAE
Sphyraena qen
ie Klu
nzin
ger, 1870
2002
SCOMBRIDAE
Rastrelliger kan
agurta (C
uvier, 1
816)
P
Scomberom
orus com
merson
(Lacep
ède, 1
800)
2002
BALISTIDAE
Abalistes stellatu
s (Lacep
ède, 1
798)
P
Balistapu
s un
dulatu
s (Park, 1
797)
2002
Balistoides con
spicillum
(Bloch
& Sch
neid
er, 1801)
2002
B. viridescen
s (Bloch
& Sch
neid
er, 1801)
2002
Melichthys in
dicus R
and
all & K
lausew
itz, 1973
2002
Melichthys n
iger (Bloch
, 1786)
2010
Odon
us n
iger (Rü
pp
ell, 1836)
P
Pseu
dobalistes fuscu
s (Bloch
& Sch
neid
er, 1801)
2002
Rhin
ecanthu
s aculeatu
s (Lin
naeu
s, 1758)
2002
Suffl am
en bu
rsa (Bloch
& Sch
neid
er, 1801)
2002
Reef fish species list, northeast Madagascar
91 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Fish, continued
SPECIESRECORD
S. chrysoptera (Bloch
& Sch
neid
er, 1801)
2002
S. fraenatu
s (Latreille, 1
804)
2002
MONACANTHIDAE
Alu
terus scriptu
s (Osb
eck, 1765)
P
Am
anses scopas (C
uvier, 1
829)
2002
Can
therines du
merilii (H
ollard, 1
854)*
2002
C. pardalis (R
üp
pell, 1
837)
P
Oxym
onacan
thus longirostris (B
loch &
Schn
eider, 1
801)
2002
Param
onacan
thus sp.
2010
Pervagor jan
thinosom
a (Bleeker, 1
854)
2002
OSTRACIIDAE
Ostracion
cubicu
s Lin
naeu
s, 1758
2002
O. m
eleagris Shaw
, 1796
2002
TETRAODONTIDAE
Arothron
hispidus (L
inn
aeus, 1
758)
2002
A. n
igropun
ctatus (B
loch &
Schn
eider, 1
801)
2002
Can
thigaster amboin
ensis (B
leeker, 1864)
2010
C. ben
netti (B
leeker, 1854)
2002
C. petersii (A
llen in
press)
2010
C. sm
ithae Allen
& R
and
all, 1977
2010
C. solan
dri (Rich
ardson
, 1844)
2002
C. valen
tini (B
leeker, 1853)
2002
92Rapid Assessm
ent Program
Appendix 5
Echinoderm species list, northeast
Madagascar
table contin
ued on
next page
Species list – Echinoderms
Asteroidea (sea stars)
Aqu
ilonastra bu
rtonii [b
ut see O
’Lou
ghlin
& R
owe 2
006]
Asteropsis carin
ifera
Culcita schm
ideliana
Echin
aster cf. calosus
From
ia indica (P
errier, 1869)
From
ia milleporella (L
amarck, 1
816)
From
ia sp.
Lin
ckia laevigata
Mon
achaster sanderi (M
eissner, 1
892)
Neopherdin
a cf. cum
ingi (G
ray, 1940)
Crinoidea (crinoids)
Com
aster cf. gracilis
Dorom
etra mau
ritiana
Lam
prometre palm
ata
Echinoidea (sea urchins)
Astropyga radiata
Diadem
a savignyi
Diadem
a setosum
Echin
ometra m
athaei
Echin
ostrephus m
olaris
Echin
otrix calamaris
Echin
otrix diadema
Eucidaris m
etularia
Toxopn
eustes pileolu
s
Tripn
eustes gratilla
Euryalidae (basket star)
Euryale aspera (L
amarck 1
816)
Species list – Echinoderms
Holothuroidea (sea cucumbers)
Actin
opiga mau
ritiana
Actin
opyga cf. miliaris
Bohadschia m
armorata (Jaeger, 1
833)
Bohadschia sim
ilis
Bohadschia vitien
sis
Euapta godeff royi
Holothu
ria atra
Holothu
ria cynarescen
s (Bran
dt, 1
835)
Holothu
ria edulis
Holothu
ria hilla
Holothu
ria leucospilota (B
rand
t, 1835)
Holothu
ria pervicax (Selenka, 1
867)
Holothu
ria scabra versicolor
Holothu
ria verrucosa (Selen
ka, 1867)
Microthele fu
scogilva (Ch
erbon
nier, 1
980)
Microthele fu
scopun
ctata (Jaeger, 1833)
Synapta m
aculata (C
ham
isso & E
ysenh
ardt, 1
821)
Th elen
ota anax (C
lark, 1921)
Ophiuroidea (brittle stars)
Gorgon
ocephale sp.
Macrophiotrix belli (D
ördelin
, 1896)
Macrophiotrix propin
qua (L
yman
,1861)
Macrophiotrix ru
gosa (Clarck, 1
938)
Ophiarachn
a incrasata
Ophiarachn
ella gorgonia
Ophiarachn
ella macracan
tha (H.L
.Clarck,1
909)
Ophiarthru
m elegan
s (Peters, 1
851)
Echinoderm species list, northeast M
adagascar
93 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
continued
Species list – Echinoderms
Ophiocen
trus dilatatu
s
Ophiocom
a dentata (M
uller &
Troch
el)
Ophiocom
a erinaceu
s
Ophiocom
a scolopendrin
a (Lm
arck
Ophiom
astix (zèbre)
Ophiom
astix caryophyllata
Ophiom
astix koehleri
Ophiom
astix variabilis
Ophiom
astix venosa
Ophiom
ixa australis
Ophion
ereis dubia (M
uler &
Troch
el)
Ophion
ereis porrecta (Lym
an, 1
860)
Ophioplocu
s imbricatu
s
Ophiothela tigris
Ophiotrix cf faveolata
Ophiotrix n
ereidina (L
amarck, 1
816)
Ophiotrix proteu
s
Ophiotrix pu
rpurea (M
artens, 1
867)
Ophiotrix trilin
eata
94Rapid Assessm
ent Program
Tab
le begin
s on n
ext page.
Appendix 6
Echinoderm distribution by site, northeast
Madagascar
Echinoderm distribution by site, northeast M
adagascar
95 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table continued on next page
Echinoderms.
Family Species
Ambodivahibe Loky Vohemar Andravina Loky Ankao
Ambo
diva
hibe
(int
érie
ur-o
uest
) : p
long
ée
Ambo
diva
hibe
(mili
eu-E
st) :
plo
ngée
Ambo
diva
hibe
(mili
eu- E
st) :
apn
ée
Ambo
diva
hibe
(Ext
. Est
) : p
long
ée
Ambo
diva
hibe
(Ext
.Oue
st) :
plo
ngée
Baie
de
Loky
(réc
if fr
ange
ant E
st :
pent
e ex
t)
Baie
de
Loky
(réc
if fr
ange
ant E
st :
Plat
ier)
Baie
de
Loky
(réc
if fr
ange
ant :
Pen
te in
t, îlo
t)
Baie
de
Loky
(Prê
s ilô
t pen
te in
t).
Baie
de
Loky
(ext
érie
ur o
uest
)
Loky
Pla
tier a
vec
bonn
e co
uver
ture
cor
allie
nne)
Loky
Pla
tier 3
mèt
res
de p
rofo
ndeu
r (su
r le
reto
ur)
Pent
e ex
tern
e, s
ortie
pas
se V
ohém
ar
Plat
ier c
ompa
ct E
st (a
pnée
)
Plat
ier r
écifa
l prè
s du
Pha
re v
erte
(apn
ée)
Pent
e ex
tern
e, s
ortie
de
la p
asse
prè
s de
la d
euxi
ème
épav
e
Pent
e ex
tern
e A
mba
vani
hara
gna
Baie
d’A
ndra
vina
(côt
é Es
t, pr
ès d
e la
fala
ise,
P.M
)
Baie
d’A
ndra
vina
(prè
s de
l’îlo
t, A.
M))
Reto
ur à
Bai
e de
Lok
y (r
écif
fran
gean
t Est
)
Baie
de
Loky
(lag
on p
eu p
rofo
nd, a
rriè
re ré
cif,
plon
gée
de n
uit)
Nosy
Ank
ao (L
agon
, dép
ress
ion
d’ar
rière
réci
f)
Nosy
Ank
ao (L
agon
sud
-est
)
Nosy
Ank
ao (P
ente
ext
erne
Est
)
Nosy
Ank
ao (p
ente
peu
pro
fond
e de
vant
vill
age
Ibis
)
Nosy
Ank
ao (P
latie
r réc
ifal –
oues
t)
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Total
Asteroidea Asterina burtoni 1 1
Asteroidea Asteropsis carinifera 1 1
Asteroidea Culcita schmideliana 1 1 2
Asteroidea Echinaster cf. calosus 1 1
Asteroidea Fromia indica (Perrier, 1869) 1 1
Asteroidea Fromia milleporella (Lamarck, 1816) 1 1 2
Asteroidea Fromia sp. 1 1
Asteroidea Linckia laevigata 1 1
Asteroidea Monachaster sanderi (Meissner, 1892) 1 1 2
Asteroidea Neopherdina cf.cumingi (Gray, 1940) 1 1
Crinoidea Comaster cf. gracilis 1 1
Crinoidea Dorometra mauritiana 1 1
Crinoidea Lamprometre palmata 1 1
Echinoidea Astropyga radiata 1 1
Echinoidea Diadema savignyi a
Echinoidea Diadema setosum 1 1
Echinoidea Echinometra mathaei 1 1 1 1 1 5
Echinoidea Echinostrephus molaris 1 1 1 1 1 1 1 1 1 1 1 1 12
Echinoidea Echinotrix calamaris 1 1 1 3
Echinoidea Echinotrix diadema 1 1 1 1 1 5
Echinoidea Eucidaris metularia 1 1 1 1 4
Echinoidea Toxopneustes pileolus 1 1 1 3
Echinoidea Tripneustes gratilla 1 1
Euryalidae Euryale aspera (Lamarck 1816) 1 1
Holothuroidea Actinopiga mauritiana 1 1 2
Holothuroidea Actinopyga cf. miliaris 1 1
Holothuroidea Bohadschia marmorata (Jaeger, 1833) 1 1 1 1 4
Appendix 6
96Rapid Assessm
ent Program
Echinoderms, continued
table continued on next page
Family Species
Ambodivahibe Loky Vohemar Andravina Loky Ankao
Ambo
diva
hibe
(int
érie
ur-o
uest
) : p
long
ée
Ambo
diva
hibe
(mili
eu-E
st) :
plo
ngée
Ambo
diva
hibe
(mili
eu- E
st) :
apn
ée
Ambo
diva
hibe
(Ext
. Est
) : p
long
ée
Ambo
diva
hibe
(Ext
.Oue
st) :
plo
ngée
Baie
de
Loky
(réc
if fr
ange
ant E
st :
pent
e ex
t)
Baie
de
Loky
(réc
if fr
ange
ant E
st :
Plat
ier)
Baie
de
Loky
(réc
if fr
ange
ant :
Pen
te in
t, îlo
t)
Baie
de
Loky
(Prê
s ilô
t pen
te in
t).
Baie
de
Loky
(ext
érie
ur o
uest
)
Loky
Pla
tier a
vec
bonn
e co
uver
ture
cor
allie
nne)
Loky
Pla
tier 3
mèt
res
de p
rofo
ndeu
r (su
r le
reto
ur)
Pent
e ex
tern
e, s
ortie
pas
se V
ohém
ar
Plat
ier c
ompa
ct E
st (a
pnée
)
Plat
ier r
écifa
l prè
s du
Pha
re v
erte
(apn
ée)
Pent
e ex
tern
e, s
ortie
de
la p
asse
prè
s de
la d
euxi
ème
épav
e
Pent
e ex
tern
e A
mba
vani
hara
gna
Baie
d’A
ndra
vina
(côt
é Es
t, pr
ès d
e la
fala
ise,
P.M
)
Baie
d’A
ndra
vina
(prè
s de
l’îlo
t, A.
M))
Reto
ur à
Bai
e de
Lok
y (r
écif
fran
gean
t Est
)
Baie
de
Loky
(lag
on p
eu p
rofo
nd, a
rriè
re ré
cif,
plon
gée
de n
uit)
Nosy
Ank
ao (L
agon
, dép
ress
ion
d’ar
rière
réci
f)
Nosy
Ank
ao (L
agon
sud
-est
)
Nosy
Ank
ao (P
ente
ext
erne
Est
)
Nosy
Ank
ao (p
ente
peu
pro
fond
e de
vant
vill
age
Ibis
)
Nosy
Ank
ao (P
latie
r réc
ifal –
oues
t)
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Total
Holothuroidea Bohadschia similis 1 1
Holothuroidea Bohadschia vitiensis 1 1 2
Holothuroidea Euapta godeff royi a
Holothuroidea Holothuria atra 1 1 1 1 1 5
Holothuroidea Holothuria cynarascens (Brandt, 1835) 1 1
Holothuroidea Holothuria edulis 1 1 2
Holothuroidea Holothuria hila 1 1
Holothuroidea Holothuria leucospilota (Brandt, 1835) 1 1
Holothuroidea Holothuria pervicax (Selenka, 1867) 1 1
Holothuroidea Holothuria scabra versicolor 1 1
Holothuroidea Holothuria verrucosa (Selenka, 1867) 1 1 2
Holothuroidea Microthele fuscogilva (Cherbonnier, 1980) 1 1
Holothuroidea Microthele fuscopunctata (Jaeger, 1833) 1 1
Holothuroidea Synapta maculata (Chamisso&Eysenhardt, 1821) 1 1 2
Holothuroidea Th elenota anax (Clark, 1921) 1 1
Ophiuroidea Gorgonocephale sp. 1 1
Ophiuroidea Macrophiotrix belli (Dördelin, 1896) 1 1
Ophiuroidea Macrophiotrix propinqua (Lyman,1861) 1 1 2
Ophiuroidea Macrophiotrix rugosa (Clarck, 1938) 1 1 2
Ophiuroidea Ophiarachna incrasata 1 1 1 3
Ophiuroidea Ophiarachnella gorgonia 1 1 1 1 1 1 6
Ophiuroidea Ophiarachnella macracantha (H.L.Clarck,1909) 1 1 2
Ophiuroidea Ophiarthrum elegans (Peters, 1851) 1 1 1 1 1 5
Ophiuroidea Ophiocentrus dilatatus 1 1
Ophiuroidea Ophiocoma dentata (Muller & Trochel) 1 1
Ophiuroidea Ophiocoma erinaceus 1 1 1 1 1 1 1 7
Ophiuroidea Ophiocoma scolopendrina (Lmarck) 1 1
Ophiuroidea Ophiomastix (zèbre) 1 1 1 1 1 5
Echinoderm distribution by site, northeast M
adagascar
97 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
Echinoderms, continued
Family Species
Ambodivahibe Loky Vohemar Andravina Loky Ankao
Ambo
diva
hibe
(int
érie
ur-o
uest
) : p
long
ée
Ambo
diva
hibe
(mili
eu-E
st) :
plo
ngée
Ambo
diva
hibe
(mili
eu- E
st) :
apn
ée
Ambo
diva
hibe
(Ext
. Est
) : p
long
ée
Ambo
diva
hibe
(Ext
.Oue
st) :
plo
ngée
Baie
de
Loky
(réc
if fr
ange
ant E
st :
pent
e ex
t)
Baie
de
Loky
(réc
if fr
ange
ant E
st :
Plat
ier)
Baie
de
Loky
(réc
if fr
ange
ant :
Pen
te in
t, îlo
t)
Baie
de
Loky
(Prê
s ilô
t pen
te in
t).
Baie
de
Loky
(ext
érie
ur o
uest
)
Loky
Pla
tier a
vec
bonn
e co
uver
ture
cor
allie
nne)
Loky
Pla
tier 3
mèt
res
de p
rofo
ndeu
r (su
r le
reto
ur)
Pent
e ex
tern
e, s
ortie
pas
se V
ohém
ar
Plat
ier c
ompa
ct E
st (a
pnée
)
Plat
ier r
écifa
l prè
s du
Pha
re v
erte
(apn
ée)
Pent
e ex
tern
e, s
ortie
de
la p
asse
prè
s de
la d
euxi
ème
épav
e
Pent
e ex
tern
e A
mba
vani
hara
gna
Baie
d’A
ndra
vina
(côt
é Es
t, pr
ès d
e la
fala
ise,
P.M
)
Baie
d’A
ndra
vina
(prè
s de
l’îlo
t, A.
M))
Reto
ur à
Bai
e de
Lok
y (r
écif
fran
gean
t Est
)
Baie
de
Loky
(lag
on p
eu p
rofo
nd, a
rriè
re ré
cif,
plon
gée
de n
uit)
Nosy
Ank
ao (L
agon
, dép
ress
ion
d’ar
rière
réci
f)
Nosy
Ank
ao (L
agon
sud
-est
)
Nosy
Ank
ao (P
ente
ext
erne
Est
)
Nosy
Ank
ao (p
ente
peu
pro
fond
e de
vant
vill
age
Ibis
)
Nosy
Ank
ao (P
latie
r réc
ifal –
oues
t)
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Total
Ophiuroidea Ophiomastix caryophyllata* 1 1 1 1 1 1 6
Ophiuroidea Ophiomastix koehleri 1 1 1 3
Ophiuroidea Ophiomastix variabilis 1 1
Ophiuroidea Ophiomastix venosa 1 1
Ophiuroidea Ophiomixa australis 1 1 1 1 1 5
Ophiuroidea Ophionereis dubia (Muler &Trochel) 1 1
Ophiuroidea Ophionereis porrecta (Lyman, 1860) 1 1 1 1 4
Ophiuroidea Ophioplocus imbricatus 1 1
Ophiuroidea Ophiothela tigris 1 1 2
Ophiuroidea Ophiotrix cf faveolata 1 1
Ophiuroidea Ophiotrix nereidina (Lamarck, 1816) 1 1
Ophiuroidea Ophiotrix proteus 1 1
Ophiuroidea Ophiotrix purpurea (Martens, 1867) 1 1 2
Ophiuroidea Ophiotrix trilineata 1 1
6 13 3 12 1 2 6 10 6 18 2 3 2 15 4 9 2 5 2 7 6 3 3 1 2 4 146
98Rapid Assessm
ent Program
Appendix 7
Algae and seagrass species distribution
by location, northeast Madagsacar
NOTES
AMBV
LOKY BayVohem
arAndravina
Nosy Ankao
CHLOROPHYCEAE
1U
lva lactuca
chan
nel
++
2A
nadyom
ene stellata
+
3C
haetomorpha in
dica+++
4C
ladophora sp.+
5B
oergensen
ia forbesii+++
+
6B
oodlea composita
+
7C
aulerpa racem
osa+
++++
8C
aulerpa len
tillifera
9C
aulerpa sertu
laroides++
+++
10
Cau
lerpa taxifolia++
+++
+++
+++
11
Cau
lerpa serrulata
++
+++
12
Cau
lerpa num
mularia
++
13
Cau
lerpa brachypus
cultu
red+
+++
14
Udotea in
dica+++
+++
++
15
Udotea sp.
+++
16
Halim
eda macroloba
+++
+++
17
Halim
eda discoides+
+++
18
Halim
eda opun
tia++
19
Halim
eda cun
eata+++
20
Dictyosphaeria cavern
osa++
21
Dictyosphaeria verslu
ysii++
+
22
En
theromorpha fl exu
osa+++
23
Valon
ia sp.+
+
24
Valon
ia sp1.
25
Ven
tricaria (Valon
ia) ventricosa
++
++
26
Bryopsis sp.
++
27
Codiu
m sp.
+
28
Avrain
villea erecta +++
++
29
Neom
eris vanbosseae
+++
++
++
30
Acetabu
laria pavula
+
table contin
ued on
next page
Algae species list. Key: + few, +
+ com
mon, +
++
abundant
Algae and seagrass species distribution by location, northeast Madagsacar
99 A Rapid M
arine Biodiversity Assessment of the Coral Reefs of N
ortheast Madagascar
table contin
ued on
next page
Algae, continued
NOTES
AMBV
LOKY BayVohem
arAndravina
Nosy Ankao
31
Tydem
ania expedition
is
32
Microdyction
sp.++
RHODOPHYCEAE
1N
eurym
enia fraxin
ifolia+++
2A
man
sia rhodantha
++
3H
ypnea pan
nosa
++
++
4H
ypnea corn
uata
cultu
red+++
+++
+++
++++
5H
ypnea n
idifi ca
6C
arpopeltis mallardii
+++
7G
alaxaura obtu
sata+++
8G
alaxaura fascicu
lata+++
9G
alaxaura m
arginata
+++
10
Phen
ophyllum
sp.++
11
Lithophyllu
m sp.
++
++
12
Neogon
iolithon brassica
+++
13
Lau
rencia sp
1.
+
14
Lau
rencia sp
2.
++
++
15
Actin
otrichia fragilis++
+
16
Am
phiroa anceps
+++
+
17
Am
phiroa fragilissima
+
18
Am
phiroa sp.++
19
Jania adhaeren
s+++
20
Carpopeltis m
allardiid
eep+
21
Crypton
emia u
ndu
lata+++
22
Peysson
nelia capen
sis+++
23
Peysson
nelia sim
ulan
s
24
Galaxau
ra elongata
26
Galaxau
ra sp.
27
Ptilophora sp.
28
Marten
sia elegans
++
29
Acan
thophora spicifera
30
Chon
drophycus papillosu
s
31
Tolypiocladia sp.
32
Gracilaria salicorn
ia+
33
Gracilaria corticata
34
Gracilaria acru
ata
35
Phacelocarpu
s tristichus
++
36
Cham
pia indica
+++
37
Cham
pia sp.+++
38
En
dosiphonia horrida
+
39
Gelidiella acerosa
++
40
Sarconem
a sp.
41
Digen
ia simplex
Appendix 7
100Rapid Assessm
ent Program
Algae, continued
NOTES
AMBV
LOKY BayVohem
arAndravina
Nosy Ankao
42
Fau
chea peltata+
43
Sarconem
a fi liforme
44
Osm
un
daria ???
PHEOPHYCEAE
1D
ictyota hum
ifusa
+++
++
2D
ictyota sp1.
++
3D
ictyota sp2.
4D
ictyota sp3.
5D
ictyota sp4.
6Padin
a boryana
+
7Padin
a gymn
ospora
8L
obophora sp.++
9T
urbin
aria decurren
s+++
10
Turbin
aria conides
++
11
Sargassum
sp.++
Seagrass
NOTESAM
BVLOKY Bay
Vohemar
AndravinaNosy Ankao
SEAGRASSES
1C
ymodocea rotu
ndata
yy
y
2C
ymodocea serru
latay
yy
y
3H
alodule u
nin
ervisy
yy
4H
alodule w
rightiiy
yy
5H
alophila ovalisy
yy
y
6H
alophila stipulacea
yy
yy
7Syrin
godium
isoetifolium
yy
y
8Th alassia hem
prichiiy
yy
9Th alassoden
dron ciliatu
my
yy
10
Zostera capen
sisy
yy