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.

REFERENCES

Allen

, G. R

. 2005. R

eef fi shes of N

orthw

estern M

adagascar.

In: M

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

Bu

lletin of

Biological A

ssessmen

t 31. C

onservation

Intern

ational,

Wash

ington

DC

, USA

. Pp. 3

9-4

8.

AN

GA

P, 2001. P

lan d

e Gestion

du

Réseau

Nation

al des

Aires P

rotégées de M

adagascar. P

arcs Nation

aux M

ada-

gascar, An

tanan

arivo. Mad

agascar.C

oles, S. and

B.E

Brow

n. 2

003. C

oral bleach

ing –

capacity

for acclimatization

and

adap

tation. A

dvan

ces in M

arine

Biology 4

6: 1

83–224.

Con

servation In

ternation

al. 2009. A

ssessing th

e imp

acts of clim

ate chan

ge on M

adagascar’s b

iodiversity an

d

livelihood

s. A w

orkshop

report. M

EE

FT

, CI, W

WF.

An

tanan

arivo. Mad

agascarM

aharavo, J., T

. A. O

liver and

A. R

abearisoa. In

press. A

R

apid

Marin

e Biod

iversity Assessm

ent of N

ortheast

Mad

agascar. Con

servation In

ternation

al. An

tanan

arivo. M

adagascar.

Cooke, A

., O. R

atomah

enin

a, E. R

anaivoson

and

H. R

aza-fi n

drain

ibe. 2

000. C

hap

ter 60. M

adagascar. In

: C. R

. C

. Shep

pard

(ed.). Seas at th

e Millen

niu

m: an

environ

-m

ental evalu

ation. V

olum

e 2. R

egional ch

apters: th

e In

dian

Ocean

to the P

acifi c. Pergam

on, A

msterd

am. P

p. 113-1

31.

Main

a, J. and

D.O

. Obu

ra. 2008. C

limate ch

ange: Sp

atial d

ata for Coastal an

d M

arine E

cosystem V

uln

erability

Assessm

ents in

Mad

agascar. WW

F/C

I. An

tanan

arivo. M

adagascar.

Main

a, J., V. V

enu

s, T.R

. McC

lanah

an, an

d M

. Atew

eber-

han

. 2008. M

odelin

g ecological suscep

tibility of coral

reefs to environ

men

tal stress usin

g remote sen

sing, G

IS an

d in

situ ob

servations: a case stu

dy in

the W

estern

Ind

ian. O

cean.E

col. Mod

. 212: 1

80–199.

McC

lanah

an, T

. R., M

. Atew

eberh

an, C

. Mu

han

do, J.

Main

a, and

S.M. M

oham

med

. 2007. C

limate ch

ange

and

spatio-tem

poral variation

in seaw

ater temp

erature

eff ects on coral b

leachin

g and

mortality. E

col. Mon

ogr. 77: 5

03–525.

McK

enn

a, S., G.R

. Allen

and

H. R

and

rianasolo (ed

s.). 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 B

iological Assessm

ent 3

1. C

onservation

Intern

ational,

Wash

ington

DC

, USA

.O

bu

ra, D.O

. 2009. C

oral Reef R

esilience A

ssessmen

t of the

Nosy H

ara Marin

e Protected

Area, N

orthw

est Mad

agas-car. W

WF

Mad

agascar.O

bu

ra, D.O

. 2005. R

esilience an

d clim

ate chan

ge: les-son

s from coral reefs an

d b

leachin

g in th

e Western

In

dian

Ocean

. Estu

arine C

oastal and

Shelf Scien

ce 63:3

53–372.

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.

REFERENCES

Allen

, G. R

. 2002. D

escription

s of two n

ew sp

ecies of dam

-selfi sh

es (Pom

acentrid

ae: Pom

acentru

s) from M

adagas-

car. Journ

al of Ichth

yology and

Aqu

atic Biology 6

(2):

45-5

2.

Allen

, G. R

. 2005. R

eef fi shes of N

orthw

estern M

adagascar.

In: M

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

Bu

lletin of

Biological A

ssessmen

t 31. C

onservation

Intern

ational,

Wash

ington

DC

, USA

. Pp. 3

9-4

8.

Allen

, G.R

. and

T.B

. Wern

er. 2002. C

oral reef fi sh assess-

men

t in th

e coral triangle of sou

theastern

Asia. E

nv.

Biol. F

ish. 6

5: 2

09-2

14.

Allen

, G.R

., N. C

ross, C. A

llen, an

d M

. Gom

on. 2

006.

LA

BR

IDA

E: W

rasses. Zoological C

atalogue of A

ustra-

lia. Volu

me 3

5. F

ishes.

Bru

ce R.W

. and

J.E. R

and

all. 1984. Scarid

ae. FA

O Sp

ecies id

entifi cation

sheets for fi sh

ery pu

rposes.

Esch

meyer, W

. N. 2

010. C

atalog of Fish

es 2010. O

n lin

e au

thoritative referen

ce for taxonom

ic fi sh n

ames. C

ali-forn

ia Acad

emy of Scien

ces. Web

site: http

://research.

calacadem

y.org/ichth

yology/catalogC

hoat, J.H

. and

J.E. R

and

all. 1983. A

revision of th

e par-

rotfi shes (fam

ily Scaridae) of th

e Great B

arrier Reef of

Au

stralia with

a descrip

tion of a n

ew sp

ecies. Record

s of th

e Au

stralian M

useu

m 3

8: 1

75-2

39.

Brock, R

.E. 1

982. A

critiqu

e of the visu

al censu

s meth

od for

assessing coral reef fi sh

pop

ulation

s. Bu

ll. Mar. Sci. 3

2:

269–276.

Bellw

ood, D

.R., T

.P. Hu

ghes, C

. Folke, an

d M

. Nyström

. 2004. C

onfron

ting th

e coral reef crisis. Natu

re 429:

827- 8

33.

Table 2.6. Total herbivore abundance (all trophic groups combined). Sites

recomm

ended for focussed protection through no-take zones are bolded.

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

39 A Rapid M

arine Biodiversity Assessment of the Coral Reefs of N

ortheast Madagascar

Ch

aban

et, P. and

P. Du

rville. 2005. R

eef fi sh in

vetory of Ju

an D

e Nova’s N

atural P

ark (Western

Ind

ian O

cean).

Western

Ind

ian O

cean J. M

ar. Sci. 4(2

):145-1

62.

David

son, J., N

. Hill, L

. Mu

aves, S. Mu

caves, I. Silva, A.

Gu

issamu

lo and

A. Sh

aw. 2

006. V

amizi Islan

d M

ozam-

biq

ue: M

arine E

cological Assessm

ent O

ctober 2

006.

ZSL

/Malu

ane R

eport.

En

glish, S., C

. Wilkin

son an

d V

. Baker (ed

s). 1994. Su

rvey m

anu

al for tropical m

arine resou

rces. ASE

AN

-Au

stralia Scien

ce Project, A

ustralian

Institu

te of Marin

e Science,

Tow

nsville.

Fricke, R

., T. M

uloch

au, P. D

urville, P. C

hab

anet, E

. Tessier

and

Y. L

etourn

eur. 2

009. A

nn

otated ch

ecklist of the fi sh

sp

ecies (Pisces) of L

a Réu

nion

, inclu

din

g a Red

List of

threaten

ed an

d d

eclinin

g species. Stu

ttgarter Beiträge

zur N

aturku

nd

e A, N

eue Serie. v. 2

: 1-1

68.

Grah

am N

.A.J., S.K

. Wilson

, S. Jenn

ings, N

.V.C

. Polu

nin

, J. R

obin

son, J.P. B

ijoux an

d T

.M. D

aw. 2

007. L

ag eff ects in

the im

pacts of m

ass coral bleach

ing on

Coral

Reef F

ish, F

isheries, an

d E

cosystems. C

onservation

Bio-

logy 21 (5

): 1291–1300.

Grah

am, N

.A.J., T

.R. M

cClan

ahan

, M.A

. MacN

eil, S.K.

Wilson

, N.V

.C. P

olun

in an

d al. 2

008. C

limate W

ar-m

ing, M

arine P

rotected A

reas and

the O

cean-Scale

Integrity of C

oral Reef E

cosystems. P

LoS O

NE

3(8

): e3

039.

Green

, A.L

. and

D.R

. Bellw

ood. 2

009. M

onitorin

g fun

c-tion

al group

s of herb

ivorous reef fi sh

es as ind

icators of coral reef resilien

ce – A

practical gu

ide for coral reef

man

agers in th

e Asia P

acifi c region. IU

CN

workin

g grou

p on

Clim

ate Ch

ange an

d C

oral Reefs. IU

CN

, G

land

, Switzerlan

d.

Harm

elin-V

ivien, M

.L., J.G

. Harm

elin, C

. Ch

auvet, C

. D

uval, R

. Galzin

, P. Lejeu

ne, G

. Barn

abe, F. B

lanc, R

. G

hevalier, J. D

uclerc an

d G

. Lasserre. 1

985. E

valuation

visu

elle des p

eup

lemen

ts et pop

ulation

s de p

oissons:

meth

odes et p

roblem

es. Review

of Ecology (T

erre Vie)

40: 4

68–539.

Heem

stra and

J.E. R

and

all. 1993. G

roup

ers of the W

orld

(Family Serran

idae, Su

bfam

ily Ep

inep

helin

ae). An

an

notated

and

illustrated

catalogue of th

e group

er. FA

O

Fish

eries Synop

sis 125. F

AO

, Rom

e. H

ugh

es, T.P., D

. R. B

ellwood

, C. F

olke, R.S. Sten

eck, and

J. W

ilson, 2

005. N

ew p

aradigm

s for sup

portin

g the

resilience of m

arine ecosystem

s. Tren

ds in

Ecology an

d

Evolu

tion 2

0(7

): 380-3

86.

Ku

iter, R. an

d H

. Deb

elius. 2

006. W

orld A

tlas of Marin

e F

ishes. C

onch

Books.

Lieske, E

. and

R. M

yers. 1996. C

oral Reef F

ishes. P

rinceton

U

niversity P

ress. 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 B

iological Assessm

ent 3

1. C

onservation

Intern

ational,

Wash

ington

DC

, USA

.

McC

lanah

an, T

.R., B

. Kau

nd

a-Arara an

d J. O

. Om

ukoto.

2010. C

omp

osition an

d d

iversity of fi sh an

d fi sh

catches

in closu

res and

open

-access fi sheries of K

enya. F

isheries

Man

agemen

t and

Ecology 1

7: 6

3-7

6.

Ru

ss, G.R

. 2002. Yet A

noth

er Review

of Marin

e Reserves

as Reef F

ishery M

anagem

ent T

ools. In: Sale P.F. (ed

). 2002. C

oral Reef F

ishes: D

ynam

ics and

Diversity in

a C

omp

lex Ecosystem

. Acad

emic P

ress. Pp. 4

21-4

43.

Obu

ra, D. 2

004. B

iodiversity Su

rveys of the C

oral Reefs

of Mn

azi Bay - R

uvu

ma E

stuary M

arine P

ark. IUC

N

EA

RO

Pu

blication

. O

bu

ra, D.O

. 2008. Scleractin

ian coral fau

na of th

e Western

In

dian

Ocean

. In: O

bu

ra, D.O

., J. Tam

eland

er, , and

O

. Lin

den

(eds).T

en years after b

leachin

g – facin

g the

conseq

uen

ces of climate ch

ange in

the In

dian

Ocean

. C

OR

DIO

Status R

eport 2

008. C

OR

DIO

(Coastal

Ocean

s Research

and

Develop

men

t in th

e Ind

ian

Ocean

) / Sida-SA

RE

C. M

ombasa. K

enya. P

p. 139-1

47.

Obu

ra, D.O

., M.A

. Samoilys, J. L

utjeh

arms an

d J. H

ermes.

2008. Is th

ere a Western

Ind

ian O

cean “C

oral Tri-

angle”? M

arine Scien

ce for Man

agemen

t/WIO

MSA

P

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

CN

w

orking grou

p on

Clim

ate Ch

ange an

d C

oral Reefs.

IUC

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

Dep

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.

REFERENCES

Coles, S.L

. and

B.E

. Brow

n. 2

003. C

oral bleach

ing - C

apa-

city for acclimatization

and

adap

tation. A

dv. M

ar. Biol.

46:1

83-2

23.

Hoegh

-Gu

ldberg, O

, P. J. Mu

mby, A

. J. Hooten

, R. S. Ste-

neck, P. G

reenfi eld

, E. G

omez, C

. D. H

arvell, P. F. Sale, A

. J. Ed

ward

s, K. C

aldeira, N

. Kn

owlton

, C. M

. Eakin

, R

. Iglesias-Prieto, N

. Mu

thiga, R

. H. B

radbu

ry, A. D

ubi

and

M. E

. Hatziolos. 2

007. C

oral Reefs U

nd

er Rap

id

Clim

ate Ch

ange an

d O

cean A

cidifi cation

. Science 3

18

(5857): 1

737-1

742

Kn

owlton

, N.K

., R.E

. Brain

ard, R

. Fish

er, M. M

oews, L

. P

laisance an

d M

.J. Caley. 2

010. C

oral Reef B

iodiver-

sity. In: M

cIntyre, A

. (ed.). L

ife in th

e World

’s Ocean

s: D

iversity, Distrib

ution

, and

Abu

nd

ance. B

lackwell

Pu

blish

ing.

Liu

G, A

. E. Stron

g, W. Skirvin

g, and

L. F. A

rzayus, 2

005.

Overview

of NO

AA

coral reef watch

program

’s near-real

time satellite glob

al coral bleach

ing m

onitorin

g activi-ties. P

roc .10th

Int. C

oral Reef Sym

p, Okin

awa, Jap

an,

2004. 1

:1783-1

793.

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.

McC

lanah

an, T

. R., M

. Atew

eberh

an, C

. Ru

iz Sebastia´n

, N

. A. J. G

raham

, S. K. W

ilson, J. H

. Bru

ggeman

n an

d

M. M

. M. G

uillau

me. 2

007. P

redictab

ility of coral bleach

ing from

synop

tic satellite and

in situ

temp

erature

observation

s. Coral R

eefs 26:6

95-7

01

McK

enn

a, S., G.R

. Allen

and

H. R

and

rianasolo (ed

s.). 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 B

iological Assessm

ent 3

1. C

onservation

Intern

ational,

Wash

ington

DC

, USA

.M

oberg, F. 1

999. E

cological goods an

d services of coral reef

ecosystems. E

nviron

men

tal Econ

omics 2

9 (2

): 215-2

33

NO

AA

. 2007. N

ational C

limatic D

ata Cen

ter. Web

site: h

ttp:// w

ww

.ncd

c.noaa.gov/oa/n

cdc.h

tml

Obu

ra, D.O

. and

G. G

rimsd

itch.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

CN

w

orking grou

p on

Clim

ate Ch

ange an

d C

oral Reefs.

IUC

N, G

land

, Switzerlan

d.

Obu

ra, D.O

. 2009. C

oral Reef R

esilience A

ssessmen

t of the

Nosy H

ara Marin

e Protected

Area, N

orthw

est Mad

a-gascar. C

OR

DIO

East A

frica Rep

ort for “Bu

ildin

g R

esilient M

arine P

rotected A

reas in M

adagascar” W

WF

P

roject MG

922”. M

adagascar.

Qu

od, J.P., Y. D

ahalan

i, L. B

igot, J. B. N

icet, S. Ah

a-m

ada 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.

Strong, A

. E., G

. Liu

, J. Meyer, J. C

. Hen

dee, an

d D

. Sasko, 2004. C

oral Reef W

atch 2

002. B

ulletin

of Marin

e Sci-en

ce 75(2

): 259-2

68.

Skirving, W

.J., A.E

. Strong, G

. Liu

, C. L

iu, F. A

rzayus, J.

Sapp

er, and

E. B

ayler. 2006. E

xtreme even

ts and

pertu

r-bation

s of coastal ecosystems: Sea su

rface temp

erature

chan

ge and

coral bleach

ing. In

: Rich

ardson

, L. L

. and

E

.F. LeD

rew (ed

s.). Rem

ote Sensin

g of Aqu

atic Coastal

Ecosystem

Processes, K

luw

er pu

blish

ers. W

ebster, F. J. an

d K

. McM

ahon

. 2002. 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.

Wilkin

son, C

. (ed.). 2

004. Statu

s of coral reefs of the W

orld:

2004. V

olum

e 1. Statu

s of coral reefs of the W

orld.

Au

stralian In

stitute of M

arine Scien

ce. Tow

nsville.

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