couturier et al. 2012 mobulid review.pdf

Journal of Fish Biology (2012) 80, 1075–1119

doi:10.1111/j.1095-8649.2012.03264.x, available online at wileyonlinelibrary.com

Biology, ecology and conservation of the Mobulidae

L. I. E. Couturier*†‡, A. D. Marshall§, F. R. A. Jaine†‖,T. Kashiwagi*¶, S. J. Pierce§, K. A. Townsend**, S. J. Weeks‖,

M. B. Bennett* and A. J. Richardson†‡‡††

*School of Biomedical Sciences, The University of Queensland, St Lucia, Qld 4072,Australia, †Climate Adaptation Flagship, CSIRO Marine and Atmospheric Research,

Ecosciences Precinct, GPO Box 2583, Brisbane, Qld 4102, Australia, §Marine MegafaunaAssociation and ECOCEAN USA, Tofo Beach, Inhambane, Mozambique, ‖Biophysical

Oceanography Group, School of Geography, Planning and Environmental Management, TheUniversity of Queensland, St Lucia, Qld 4072, Australia, ¶Molecular Fisheries Laboratory,

Department of Employment, Economic Development and Innovation, P. O. Box 6097,Brisbane, Qld 4072, Australia, ∗∗School of Biological Sciences, The University of

Queensland, St Lucia, Qld 4072, Australia, ††Centre for Applications in Natural ResourceMathematics, School of Mathematics and Physics, The University of Queensland, St Lucia,Qld 4072, Australia and ‡‡Environmental Decisions Group, School of Biological Sciences,

The University of Queensland, Brisbane, Qld 4072, Australia

The Mobulidae are zooplanktivorous elasmobranchs comprising two recognized species of mantarays (Manta spp.) and nine recognized species of devil rays (Mobula spp.). They are found cir-cumglobally in tropical, subtropical and temperate coastal waters. Although mobulids have beenrecorded for over 400 years, critical knowledge gaps still compromise the ability to assess thestatus of these species. On the basis of a review of 263 publications, a comparative synthesisof the biology and ecology of mobulids was conducted to examine their evolution, taxonomy,distribution, population trends, movements and aggregation, reproduction, growth and longevity,feeding, natural mortality and direct and indirect anthropogenic threats. There has been a markedincrease in the number of published studies on mobulids since c. 1990, particularly for the genusManta, although the genus Mobula remains poorly understood. Mobulid species have many com-mon biological characteristics although their ecologies appear to be species-specific, and some-times region-specific. Movement studies suggest that mobulids are highly mobile and have thepotential to rapidly travel large distances. Fishing pressure is the major threat to many mobu-lid populations, with current levels of exploitation in target fisheries unlikely to be sustainable.Advances in the fields of population genetics, acoustic and satellite tracking, and stable-isotopeand fatty-acid analyses will provide new insights into the biology and ecology of these species.Future research should focus on the uncertain taxonomy of mobulid species, the degree of over-lap between their large-scale movement and human activities such as fisheries and pollution, andthe need for management of inter-jurisdictional fisheries in developing nations to ensure theirlong-term sustainability. Closer collaboration among researchers worldwide is necessary to ensurestandardized sampling and modelling methodologies to underpin global population estimates andstatus. © 2012 The Authors

Journal of Fish Biology © 2012 The Fisheries Society of the British Isles

Key words: by-catch; distribution; elasmobranch; fisheries; Manta; Mobula.

‡Author to whom correspondence should be addressed. Tel.: +61733467975; email: [email protected]

1075© 2012 The AuthorsJournal of Fish Biology © 2012 The Fisheries Society of the British Isles

1076 L . I . E . C O U T U R I E R E T A L .

INTRODUCTION

The Mobulidae, which comprises manta rays (Manta spp.) and devil rays (Mobulaspp.), is a diverse family of planktivorous elasmobranchs occurring worldwide intropical, subtropical and temperate waters. Represented by two genera and 11 recog-nized species, the family contains the largest rays, reaching over 7 m disc width (WD)(Compagno & Last, 1999; Marshall et al., 2009). Although early historical accountspainted devil and manta rays as ‘diabolical creatures’ and ‘ferocious brutes’, accus-ing them of stealing boats and deliberately killing divers (Gill, 1908; Saenz-Arroyoet al., 2006), it is now known that these rays are harmless to humans and feed mainlyon zooplankton. The genus Manta was recently re-described and has at least two dis-tinct species, the reef manta Manta alfredi (Krefft 1868) and the giant manta Mantabirostris (Walbaum 1792), and a third putative species, Manta sp. cf birostris (Mar-shall et al., 2009). Manta individuals can reach a maximum size ranging between 5and 7 m WD. The genus Mobula comprises nine recognized species that attain a WDfrom 1 to 5 m: the pygmy devil ray Mobula eregoodootenkee (Bleeker 1959), theAtlantic devil ray Mobula hypostoma (Bancroft 1831), the spinetail devil ray Mob-ula japanica (Muller & Henle 1841), the shortfin devil ray Mobula kuhlii (Muller& Henle 1841), the giant devil ray Mobula mobular (Bonnaterre 1788), Munk’sdevil ray Mobula munkiana Notarbartolo-di-Sciara 1987, the lesser Guinean devilray Mobula rochebrunei (Vaillant 1979), the Chilean devil ray Mobula tarapacana(Philippi 1893) and the bentfin devil ray Mobula thurstoni (Lloyd 1908).

Although the existence of mobulids has been documented since at least the 17thcentury (Willughby & Ray, 1686), there is surprisingly little information available ontheir biology and ecology. Such baseline knowledge is now urgently needed as dra-matic increases in fishing pressure on most species threatens the stability of manyregional sub-populations (Alava et al., 2002; Dewar, 2002; White et al., 2006a).Fisheries for mobulids are considered to be unsustainable because of large, directedcatches coupled with the low fecundity and conservative life history of this group.Mobulids are landed in large numbers in targeted and by-catch fisheries across theglobe, and the demand for mobulid products in the Asian market is rising (Lack& Sant, 2008, 2009; FAO, 2009). Assessment of the current conservation status ofmobulids is hampered by the paucity of information, resulting in a data-deficient sta-tus for three Mobula species (M. hypostoma, M. kuhlii and M. tarapacana) on IUCNRed List for Threatened Species (Clark et al., 2006a; Bizzarro et al., 2007, 2008).Of those that have been assessed on the IUCN Red List, four species of Mobula arelisted as near threatened (M. eregoodootenkee, M. japanica, M. munkiana and M.thurstoni ), one as vulnerable (M. rochebrunei ) and one as endangered (M. mobular)(Pierce & Bennett, 2003; Bizzarro et al., 2006; Clark et al., 2006b; Notarbartolo-di-Sciara et al., 2006; White et al., 2006b; Valenti & Kyne, 2007). Both Manta specieswere reassessed in 2011 and are listed as globally vulnerable on the IUCN Red List(Marshall et al., 2011a, b).

The Mobulidae are highly mobile epipelagic rays that are challenging to observeand investigate in their extensive oceanic environment. Predicting the occurrence ofthese pelagic species is often difficult with field observations spatially (e.g. one studyarea) and temporally (e.g. by dive time and during the day and night) restricted. Thereare relatively few locations around the world where mobulids are easily observed andapproached [e.g. Gulf of California (Notarbartolo-di-Sciara, 1988); Hawaii (Deakos

© 2012 The AuthorsJournal of Fish Biology © 2012 The Fisheries Society of the British Isles, Journal of Fish Biology 2012, 80, 1075–1119

B I O L O G Y A N D E C O L O G Y O F T H E M O B U L I DA E 1077

et al., 2011); Mozambique: (Marshall et al., 2011c); Maldives: Kitchen-Wheeler,2010); eastern Australia (Couturier et al., 2011)]. Even at well-established aggre-gation sites, obtaining detailed biological information without restraining or killingindividuals can be difficult because of the large size and fast swimming abilities ofmost mobulid species.

Species identification of mobulids has proven problematic because of the closeexternal resemblance of many species that has led to taxonomic ambiguities (e.g.M. munkiana and M. thurstoni, M. mobular and M. japanica, M. alfredi andM. birostris). The taxonomy of the family Mobulidae has a complicated historywith several generic and species synonyms described in the literature. Historically,species descriptions have been limited, often based on a single museum specimenor no type specimen at all. Misidentification of Mobula spp. is common even in thecurrent literature, where synonyms referring to several recognized species are used[e.g. Mohanraj et al. (2009), Raje & Zacharia (2009) and Borrell et al. (2011) referto M. diabolus, which is a synonym of M. mobular and M. kuhlii ]. The recent revi-sion of the genus Manta (Marshall et al., 2009) is particularly important for futurestudies and highlights the fact that past literature needs to be re-evaluated to clarifywhich species is referred to.

Since c. 1990, there has been a marked increase in the number of published studieson the Mobulidae [Fig. 1(a)]. Most of these studies consist of notes on taxonomy,reports on the occurrence of species and reports on by-catch and fisheries [Fig. 1(b)].There are relatively few studies on the life history and ecology of the 11 species,leaving serious gaps in the knowledge needed to understand the important aspectsof biology relevant to population dynamics and conservation requirements. This isparticularly true for the genus Mobula, with only 28 published studies primarilyfocused on at least one of these species since 1980 [Fig. 1(c)]. In contrast, Mantaspp. have received increasing scientific attention over the past decade [Fig. 1(c)].Manta rays are iconic species, and there is growing interest in their ecotourismpotential, especially in coastal communities and developing countries where theycan generate significant economic benefits (Anderson et al., 2010). The predictableaggregative behaviour of these species has enabled the development of this industry,as Manta spp. can be relatively easy to find and readily approached by scuba diversor by snorkellers at the surface. Partially as a result of the growing popular interest inthese species, field research has increased in the last decade at established aggregationsites (e.g. off Australia, Hawaii, Japan, Maldives and Mozambique).

One important feature of Manta spp. is that all individuals bear a unique skinpigmentation pattern on their ventral surface that can be used to differentiate individ-uals using photographic identification (photo-ID) (Kitchen-Wheeler, 2010; Marshallet al., 2011c). Photo-ID has already provided high-quality information on Mantaspp. ecology, population structure and behaviour at several locations around theworld (Kashiwagi et al., 2010, 2011; Marshall & Bennett, 2010a, b; Couturier et al.,2011; Deakos et al., 2011; Marshall et al., 2011c). Although there is an increase inpublished studies on the genus Manta, knowledge of their biology and ecology stillremains limited, with critical biological data missing on longevity, age at maturity,growth rates, diet, short and long-term movement patterns.

The current fragmented information on the biology, ecology and threats to theMobulidae in the published literature compromises an integrated understanding ofthis group and limits assessment of their global status and local and regional



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2

4

6

8

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12

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18 (a)

(b)

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Num

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ies

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20

30

40

50

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1984

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1990

1992

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1996

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2000

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2004

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1

2

3

4

5

6

7

8

9

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1696

1706

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1726

1736

1746

1756

1766

1776

1786

1796

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1826

1836

1846

1856

1866

1876

1886

1896

1906

1916

1926

1936

1946

1956

1966

1976

1986

1996

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Fig. 1. Figure legend on next page.



management needs. This review collates the available information on the Mobul-idae, both published and in the grey literature. It provides a comprehensive synthesisof all basic biological data known, the threats facing these animals and the newapproaches available to help fill some of these knowledge gaps. It is hoped that thiscomparative synthesis will stimulate future research into these understudied speciesand, in so doing, contribute in the longer term to an improved prognosis for theirconservation.

BIOLOGY AND ECOLOGY

E VO L U T I O N

Fossil, morphological and molecular data show that the Mobulidae are one of themost derived groups of elasmobranchs (Compagno, 1977; Maisey, 1984; Nishida,1990; Cappetta et al., 1993; Lovejoy, 1996; McEachran et al., 1996; Shirai, 1996;Dunn et al., 2003; de Carvalho et al., 2004; Gonzalez-Isais & Domínguez, 2004;Maisey et al., 2004; McEachran & Aschliman, 2004; Musick & Eliis, 2005; Nayloret al., 2005; Cicimurri & Knight, 2009; Claeson et al., 2010). No mobulid-likespecies has been found in the fossil record prior to the Cretaceous–Tertiary bound-ary. Although extinct species of Mobula have a fossil record extending back to theLate Oligocene (23·6–25·7 million years before present, b.p.) (Cicimurri & Knight,2009), the earliest record of Manta is from the late early-Pliocene (c. 4·8 Mb.p.)(Snyder et al., 1983; Bourdon, 1999; Purdy et al., 2001; Cappetta & Stringer, 2002),although there have been some species that were erroneously referred to as Mantapreviously (Cappetta, 1970, 1987; Case, 1980; Bourdon, 1999). Further accountson extinct genera can be found in the works of Herman (1979), Bourdon (1999),Cappetta & Stringer (2002) and Cicimurri & Knight (2009).

TA X O N O M I C H I S T O RY

The most recent, detailed taxonomic description of the recognized Mobula spp.can be found in the study of Notarbartolo-di-Sciara (1987b). While the genus Mobulacurrently comprises nine recognized species (Table I), at least 29 different specieshave been proposed previously (Notarbartolo-di-Sciara, 1987b; Pierce & Bennett,2003; Froese & Pauly, 2010; Polack, 2011). The first species to be identified,M. (Raia) mobular, was collected in the late 1700s from the Mediterranean Sea(Bonnaterre, 1788). Six species were described in the 1800s with type localitiesfrom around the globe: Jamaica, Japan, India, Malaysia, western Africa and Chile,and two in the 1900s from India and Mexico. Type material is limited, especially forthe older species, and there are no holotypes to accompany the original descriptionsof M. eregoodootenkee, M. kuhlii, M. mobular, M. tarapacana and M. thurstoni as

Fig. 1. Literature survey on mobulids: (a) all peer-reviewed literature, published technical fishery reports andhistorical literature (pre-1950s, n = 263), (b) by research theme (ordered by frequency) and (c) by recentpublications from the primary literature subdivided by genus Manta ( ) and Mobula ( ) (containingeither Manta or Mobula in the title, n = 96). Literature was sought using ISI resource and GoogleScholar using ‘Mobulid*’, ‘Mobula’, ‘Manta’, ‘Mobulidae’ and ‘devil ray’ as key words, and citationsin key papers (Notarbartolo-di-Sciara, 1987a, b; Marshall et al., 2009).



Tab

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Bio

logi

cal

and

ecol

ogic

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form

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hed

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Com

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(yea

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ed)

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reat

ened

(Pie

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&B

enne

tt,20

03)

Dat

ade

ficie

nt(B

izza

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etal

.,20

08)

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rth

reat

ened

(Whi

teet

al.,

2006

b)

Hab

itat

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gic

coas

tal

(Pie

rce

&B

enne

tt,20

03)

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gic

coas

tal

and

ocea

nic

(Biz

zarr

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al.,

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geat

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urity

(yea

rs)

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now

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idth

(WD

,m

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ity(m

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ale)

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di-S

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)

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(Big

elow

&Sc

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der,

1953

)M

:20

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(Not

arba

rtol

o-di

-Sci

ara,

1987

b)

Num

ber

ofpu

ps(W

Dat

birt

h,m

m)

1(>

241)

(Not

arba

rtol

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1987

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1(>

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(Big

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)(W

hite

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.,20

06a

)

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ter)

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-Sci

ara,

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2010

)


B I O L O G Y A N D E C O L O G Y O F T H E M O B U L I DA E 1081T

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al.,

2008

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.,20

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2007

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ther

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Tab

leI.

Con

tinue

d

Mob

ula

kuhl

iiM

obul

am

obul

ar

Num

ber

ofpu

ps(W

Dat

birt

h,m

m)

1(3

10)

(Com

pagn

o&

Las

t,19

99)

1(1

660)

(Not

arba

rtol

o-di

-Sci

ara

&Se

rena

,19

88)

Die

tU

nkno

wn

Eup

haus

iid(M

egan

ycht

ipha

nes

norv

egic

a),

smal

lfis

hes

(Cel

ona,

2004

;N

otar

bart

olo-

di-S

ciar

aet

al.,

2006

)M

axim

umm

ovem

ent

reco

rded

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now

n33

7km

in12

0da

ys(C

anes

eet

al.,

2011

)K

now

nag

greg

atio

nsU

nkno

wn

Stra

itof

Mes

sina

(Sic

ily):

Lat

esp

ring

toau

tum

n(C

elon

a,20

04;

Can

ese

etal

.,20

11);

sout

hern

Med

iterr

anea

nco

ast

inw

inte

ran

dno

rthe

rnM

edite

rran

ean

coas

tin

spri

ng(H

emid

aet

al.,

2002

,B

raid

ii&

Cap

ape,

2001

)Po

pula

tion

size

Unc

omm

on(C

ompa

gno

&L

ast,

1999

)U

nkno

wn

but

inlo

wde

nsity

(Not

arba

rtol

o-di

-Sci

ara

etal

.,20

06)

Popu

latio

ntr

end

Dec

reas

ing

(Biz

zarr

oet

al.,

2007

)D

ecre

asin

g(N

otar

bart

olo-

di-S

ciar

aet

al.,

2006

)Fi

sher

ies

Eas

tern

Indo

nesi

a(W

hite

&D

harm

adi,

2007

);M

ozam

biqu

e(A

.M

arsh

all,

unpu

bl.

obs.

)N

orth

Afr

ica

(Hem

ida

etal

.,20

02)

By-

catc

hSo

uth

Afr

ica:

shar

kba

ther

nets

(You

ng,

2001

);In

done

sia:

drif

tgi

llnet

s(W

hite

etal

.,20

06a

)

Eur

opea

ntu

napu

rse

sein

efis

hery

(Am

ande

etal

.,20

10),

east

ern

trop

ical

Atla

ntic

Oce

an:

purs

e-se

ine

fishe

ry(M

enar

det

al.,

2000

),Sa

rdin

ia:

tuna

trap

s(S

tora

iet

al.,

2011

).H

igh

mor

talit

yfr

ompe

lagi

cdr

iftn

et(i

llega

l),

long

lines

,pu

rse

sein

esan

dtr

awls

(Bau

chot

,19

87;

Cav

anag

h&

Gib

son,

2007

)C

onse

rvat

ion

actio

nsU

nkno

wn

Prot

ecte

dun

der

U.N

.(B

arce

lona

)C

onve

ntio

nfo

rth

ePr

otec

tion

Of

The

Med

iterr

anea

nSe

aA

gain

stPo

llutio

n19

76A

nnex

eII

Mob

ula

mun

kian

aM

obul

aro

cheb

rune

i

Com

mon

nam

eM

unk’

sde

vil

ray

Les

ser

Gui

nean

devi

lra

yIU

CN

Stat

us(y

ear

asse

ssed

)N

ear

thre

aten

ed(B

izza

rro

etal

.,20

06)

Vul

nera

ble

(Val

enti

&K

yne,

2007

)H

abita

tSh

allo

wco

asta

lw

ater

s(B

izza

rro

etal

.,20

06)

Surf

ace

and

clos

eto

the

botto

m(M

cEac

hran

&S

eret

,19

90)



able

I.C

ontin

ued

Mob

ula

mun

kian

aM

obul

aro

cheb

rune

i

Ges

tatio

nU

nkno

wn

Unk

now

nL

onge

vity

(yea

rs)

Unk

now

nU

nkno

wn

Age

atm

atur

ity(y

ears

)U

nkno

wn

Unk

now

nM

axim

umW

D(m

m)

1100

(Not

arba

rtol

o-di

-Sci

ara,

1987

b)

1330

(Not

arba

rtol

o-di

-Sci

ara,

1987

b)

WD

atm

atur

ity(m

m)

(M,

mal

e;F,

fem

ale)

M:

820

(Not

arba

rtol

o-di

-Sci

ara,

1987

b),

F:91

0(V

illav

icen

cio-

Gar

ayza

r,19

91)

Unk

now

n

Num

ber

ofpu

ps(W

Dat

birt

h,m

m)

1(3

50–

360)

(Not

arba

rtol

o-di

-Sci

ara,

1987

b;

Vill

avic

enci

o-G

aray

zar,

1991

)1

(>35

0)(N

otar

bart

olo-

di-S

ciar

a,19

87b

)

Die

tM

ysid

ium

spp.

(Not

arba

rtol

o-di

-Sci

ara,

1988

)Sm

all

fish

and

plan

kton

(Mai

gret

&Ly

,19

86)

Max

imum

mov

emen

tre

cord

edU

nkno

wn

Unk

now

nK

now

nag

greg

atio

nsG

ulf

ofC

alif

orni

a:m

ainl

yw

inte

r(N

otar

bart

olo-

di-S

ciar

a,19

88)

Sene

gal

coas

t(C

aden

at,

1958

)

Popu

latio

nsi

zeU

nkno

wn

Unk

now

nPo

pula

tion

tren

dU

nkno

wn

Unk

now

nFi

sher

ies

Gul

fof

Cal

ifor

nia

(Not

arba

rtol

o-di

-Sci

ara,

1987

a,b

;19

88;

Biz

zarr

oet

al.,

2009

),E

cuad

or(A

.M

arsh

all,

unpu

bl.

obs.

)

Wes

tern

Afr

ica

(Val

enti

&K

yne,

2007

)

By-

catc

hU

nkno

wn

Gill

net

(Val

enti

&K

yne,

2007

)C

onse

rvat

ion

actio

nsFi

sher

yba

nin

Mex

ico

and

Ecu

ador

Unk

now

n

Mob

ula

tara

paca

naM

obul

ath

urst

oni

Com

mon

nam

eC

hile

ande

vil

ray

Ben

tfin

devi

lra

yIU

CN

Stat

us(y

ear

asse

ssed

)D

ata

defic

ient

(Cla

rket

al.,

2006

a)

Nea

rth

reat

ened

(Cla

rket

al.,

2006

b)

Hab

itat

Oce

anic

and

occa

sion

ally

inco

asta

lw

ater

s(C

lark

etal

.,20

06a

)Sh

allo

wne

ritic

wat

ers

(Cla

rket

al.,

2006

b)

Ges

tatio

nU

nkno

wn

Unk

now

nL

onge

vity

(yea

rs)

Unk

now

nU

nkno

wn



Tab

leI.

Con

tinue

d

Mob

ula

tara

paca

naM

obul

ath

urst

oni

Age

atm

atur

ity(y

ears

)U

nkno

wn

Unk

now

nM

axim

umW

D(m

m)

3700

(Com

pagn

o&

Las

t,19

99)

1800

(Not

arba

rtol

o-di

-Sci

ara,

1987

b)

WD

atm

atur

ity(m

m)

(M,

mal

e;F,

fem

ale)

M:

2340

–25

22,

F:un

know

n(W

hite

etal

.,20

06a

)M

:>

1500

,F:

1538

(Not

arba

rtol

o-di

-Sci

ara,

1987

b;

Whi

teet

al.,

2006

a)

Num

ber

ofpu

ps(W

Dat

birt

h,m

m)

1(>

1052

)(N

otar

bart

olo-

di-S

ciar

a,19

87b

)1

(650

–85

0)(N

otar

bart

olo-

di-S

ciar

a,19

87b

)D

iet

Var

ious

plan

kton

icor

gani

sms,

smal

lfis

h(N

otar

bart

olo-

di-S

ciar

a,19

88)

Shif

tin

diet

acco

rdin

gto

seas

on:

Eup

haus

iida

esp

p.(N

.sim

plex

)in

sum

mer

,M

ysid

ium

spp.

(juv

enile

)in

win

ter,

othe

rpl

ankt

onic

orga

nism

s(N

otar

bart

olo-

di-

Scia

ra,

1988

;Sa

mps

onet

al.,

2010

)M

axim

umm

ovem

ent

reco

rded

Unk

now

nU

nkno

wn

Kno

wn

aggr

egat

ions

Gul

fof

Cal

ifor

nia:

sum

mer

and

autu

mn

(Not

arba

rtol

o-di

-Sci

ara,

1988

)G

ulf

ofC

alif

orni

a:pr

esen

tal

lye

arar

ound

(Not

arba

rtol

o-di

-Sci

ara,

1988

)Po

pula

tion

size

Unk

now

nU

nkno

wn

Popu

latio

ntr

end

Unk

now

nU

nkno

wn

Fish

erie

sL

amak

era,

Indo

nesi

a(D

ewar

,20

02),

east

ern

Indo

nesi

a(W

hite

&D

harm

adi,

2007

);G

ulf

ofC

alif

orni

a(N

otar

bart

olo-

di-S

ciar

a,19

87a

,b

);Se

nega

l(M

arin

eM

egaf

auna

Ass

ocia

tion,

pers

.co

mm

.)

Lam

aker

a,In

done

sia

(Dew

ar,

2002

);G

ulf

ofC

alif

orni

a(N

otar

bart

olo-

di-S

ciar

a,19

87a

,b

;Sa

mps

onet

al.,

2010

),ea

ster

nIn

done

sia

(Whi

te&

Dha

rmad

i,20

07);

Indi

a(P

illai

,19

98,

Moh

anra

jet

al.,

2009

)

By-

catc

hE

urop

ean

tuna

purs

e-se

ine

fishe

ry(A

man

deet

al.,

2010

),In

done

sia

(Whi

teet

al.,

2006

a);

Nor

th-w

est

Afr

ica

(Zee

berg

etal

.,20

06)

Indo

nesi

a(W

hite

etal

.,20

06a

),so

uth-

east

ern

Bra

zil

(Cas

aset

al.,

2006

)

Con

serv

atio

nac

tions

Fish

ery

ban

inM

exic

o,E

cuad

orFi

sher

yba

nin

Mex

ico,

Ecu

ador

Man

taal

fred

iM

anta

biro

stri

s

Com

mon

nam

eR

eef

man

taG

iant

man

taIU

CN

Stat

us(y

ear

asse

ssed

)V

ulne

rabl

e(M

arsh

all

etal

.,20

11a

)V

ulne

rabl

e(M

arsh

all

etal

.,20

11b

)



able

I.C

ontin

ued

Man

taal

fred

iM

anta

biro

stri

s

Hab

itat

Insh

ore,

coas

tal

area

s(M

arsh

all

etal

.,20

09)

Off

shor

e,oc

eani

c,pr

oduc

tive

coas

tline

s(M

arsh

all

etal

.,20

09)

Ges

tatio

n12

–13

mon

ths

(Mar

shal

l&

Ben

nett,

2010

b)

Unk

now

nL

onge

vity

(yea

rs)

>31

(Cla

rk,

2010

)>

20(A

.M

arsh

all,

J.H

olm

erg,

J.M

.B

runn

schw

eile

r&

S.J.

Pier

ce,

unpu

bl.

data

)A

geat

mat

urity

(yea

rs)

M:

3–

6,F:

unkn

own

(Cla

rk,

2010

)U

nkno

wn

Max

imum

WD

(mm

)50

00(M

arsh

all

etal

.,20

09)

>70

00to

9000

(Com

pagn

o&

Las

t,19

99;

Mar

shal

let

al.,

2009

)W

Dat

mat

urity

(mm

)(M

,m

ale;

F,fe

mal

e)M

:27

00–

3000

,F:

3700

–39

00(M

arsh

all

etal

.,20

09;

Cla

rk,

2010

;D

eako

s,20

10b

)M

:38

00,

F:41

30(W

hite

etal

.,20

06a

)

Num

ber

ofpu

ps(W

Dat

birt

h,m

m)

1(1

300

–15

00)

(Mar

shal

let

al.,

2009

)1

(Bee

be&

Tee-

Van

,19

41;

Whi

teet

al.,

2006

a)

Die

tPl

ankt

onic

crus

tace

ans

(Whi

tley,

1936

)Sh

rim

p,cr

abs

and

smal

lfis

hes

(Big

elow

&Sc

hroe

der,

1953

)M

axim

umm

ovem

ent

reco

rded

(km

)>

500

(Cou

turi

eret

al.,

2011

)>

1000

(Rub

inet

al.,

2008

;M

arsh

all

etal

.,20

10)

Kno

wn

aggr

egat

ions

Haw

aii;

Moz

ambi

que;

Mal

dive

s;R

yuky

uIs

land

,Ja

pan;

Yap

;In

done

sia;

east

ern

Aus

tral

ia,

Wes

tern

Aus

tral

ia(M

arsh

all

etal

.,20

09;

Kas

hiw

agi

etal

.,20

11*)

Gul

fof

Cal

ifor

nia,

Mex

ico;

Ecu

ador

;M

ozam

biqu

e;Sr

iL

anka

;In

done

sia

(Mar

shal

let

al.,

2009

;K

ashi

wag

iet

al.,

2011

*)

Popu

latio

nsi

zeTo

tal

popu

latio

nin

Moz

ambi

que:

802

indi

vidu

als

(Mar

shal

let

al.,

2011

c);

max

imum

seas

onal

popu

latio

nin

Haw

aii:

230

indi

vidu

als

(Dea

kos

etal

.,20

11),

min

imum

num

bers

atot

her

loca

tions

inK

itche

n-W

heel

er(2

010)

;C

outu

rier

etal

.(2

011)

;K

ashi

wag

iet

al.

(201

1)

Min

imum

num

bers

inK

ashi

wag

iet

al.

(201

1)

Popu

latio

ntr

end

Eas

tern

Indo

nesi

a:de

crea

sing

(Dew

ar,

2002

)U

nkno

wn



Tab

leI.

Con

tinue

d

Man

taal

fred

iM

anta

biro

stri

s

Fish

erie

sN

orth

Sula

wes

i,In

done

sia

(Ano

n,19

97);

east

ern

Indo

nesi

a(D

ewar

,20

02),

Moz

ambi

que

(Mar

shal

let

al.,

2011

a)

Indo

nesi

a(W

hite

etal

.,20

06a

),In

dia

(Pill

ai,

1998

;M

ohan

raj

etal

.,20

09),

Sri

Lan

ka(P

.H

ilton

,pe

rs.

com

m.)

;Ta

nzan

ia(B

ianc

hi,

1985

;Ji

ddaw

i&

Stan

ley,

1999

;A

.M

arsh

all,

unpu

bl.

obs.

);Pe

ru(M

.H

ardi

ng,

pers

.co

mm

.);

Gha

na(E

ssum

ang,

2010

)B

y-ca

tch

Papu

aN

ewG

uine

a(C

.R

ose,

pers

.co

mm

.);

Phili

ppin

es(D

olar

,20

04);

Kw

aZul

u-N

atal

,So

uth

Afr

ica

(You

ng,

2001

),ea

ster

nA

ustr

alia

(Sum

pton

etal

.,20

11)

Atla

ntic

Oce

an(A

man

deet

al.,

2010

);K

waZ

ulu-

Nat

al,

Sout

hA

fric

a(Y

oung

,20

01);

Wes

tT

haila

nd(A

.M

arsh

all,

unpu

bl.

obs.

);E

cuad

oran

dPe

ru(M

.H

ardi

ng,

pers

.co

mm

.);

Sout

hB

razi

l(Z

erbi

ni&

Kot

as,

1998

);G

eorg

iaan

dea

stFl

orid

a(T

rent

etal

.,19

97;

Car

lson

&L

ee,

2000

);ce

ntra

l-w

este

rnPa

cific

(Coa

net

al.,

2000

)C

onse

rvat

ion

actio

nsM

anta

prot

ecte

dzo

nes

inth

eM

aldi

ves,

Yap

and

Haw

aii

Leg

ally

prot

ecte

din

Haw

aii,

Mex

ico,

Ecu

ador

,N

ewZ

eala

nd,

Phili

ppin

es[s

eeFi

g.(4

)]

*Gen

eral

com

men

tson

abun

danc

eat

20lo

calit

ies

can

befo

und

inK

ashi

wag

iet

al.

(201

1)bu

tth

enu

mbe

rof

phot

ogra

phic

reco

rds

prov

ided

inth

eA

ppen

dix

isno

teq

uiva

lent

topo

pula

tion

size

estim

ates

.



some were not collected and some have been lost (Polack, 2011). This lack of origi-nal material and the on-going uncertainties about species’ ranges and identities, suchas possible confusion of M. mobular with M. japanica, can only be resolved throughcontemporary collection of both morphological examinations and genetic samples.

The genus Manta has a long and confused taxonomic history. Over 10 generic and25 specific synonyms have been used to describe manta rays since Walbaum’s (1792)first description (some authors ascribe the original species designation to Donndorff’s(1798) description of Walbaum’s species). Difficulties associated with transportingand preserving large specimens have resulted in a paucity of preserved specimens,and type material is absent for almost all of the species described since 1792. Thegenus currently includes two recognized species, with the possible existence of athird putative species Manta sp. cf. birostris, proposed to be endemic to the westernAtlantic Ocean and Caribbean Sea (Marshall et al., 2009). The failure to differentiatethe two Manta spp. prior to 2009 has resulted in confusion concerning almost allthe biological information available (Homma et al., 1999; Dewar et al., 2008).

D I S T R I B U T I O N

Mobulids occur in tropical and temperate seas between 40◦ N and 40◦ S, althoughthe majority of species have a tropical to subtropical distribution (Fig. 2). Theserays have an apparent preference for water temperatures of 20–26◦ C (Dewar et al.,2008; Clark, 2010; Canese et al., 2011; Marshall et al., 2011c). Mobula spp. areregarded as pelagic or epipelagic species of coastal waters, and may be encountered inboth shallow-inshore environments and deeper offshore waters (Bizzarro et al., 2007;Cortes & Blum, 2008; Scacco et al., 2009; Canese et al., 2011). Manta alfredi, M.birostris, Mobula japanica, M. tarapacana and M. thurstoni have worldwide distribu-tions, with each species reported from the Pacific, Atlantic and Indian Oceans (Clarket al., 2006a, b; White et al., 2006b; Marshall et al., 2009, 2011a, b; Kashiwagiet al., 2011) [Fig. 2(a), (b)]. The only observation of M. tarapacana from the west-ern Atlantic Ocean is based on an aerial survey off Venezuela (Notarbartolo-di-Sciara& Hillyer, 1989), and confirmation of this species occurrence in the Caribbean Seais required.

Mobula eregoodootenkee and M. kuhlii are both restricted to the Indo-West PacificOcean (Pierce & Bennett, 2003; Bizzarro et al., 2007) [Fig. 2(c)]. Mobula ere-goodootenkee has a contiguous distribution from coastal regions of southern Africa tonorthern Australia, Papua New Guinea and the Philippines (Pierce & Bennett, 2003).In contrast, records for M. kuhlii are patchy over a similar range (Bizzarro et al.,2007), possibly as a result of misidentifications or insufficient sampling [Fig. 2(a)].White et al. (2006a) refer to specimens from Cilacap, Java, Indonesia, as M. cf. kuh-lii, suggesting that there is uncertainty about the species’ identity in this region. Therecord from the Philippines is for a single individual from Iloilo City Fish Port (Com-pagno et al., 2005). Further investigation of the distribution of M. kuhlii is needed.

Mobula hypostoma is a widely distributed species endemic to coastal and shelfwaters of the western Atlantic Ocean, from North Carolina, U.S.A., through muchof the Gulf of Mexico, Greater and Lesser Antilles, to northern Argentina in thesouth (McEachran et al., 2002; Bizzarro et al., 2008) [Fig. 2(c)]. Mobula munkiana isendemic to tropical coastal and oceanic waters in the eastern Pacific Ocean, extendingto the waters of Cocos, Malpelo and Galapagos Islands (Bizzarro et al., 2006;



Smithsonian Tropical Research Institute, 2011) [Fig. 2(c)]. Mobula rochebrunei is apelagic species found primarily along the central and south-eastern Atlantic coast ofAfrica (Cadenat, 1958; Maigret & Ly, 1986; McEachran & Seret, 1990), althoughthere are isolated records of the species from coastal waters off Parana State, Brazil(Valenti & Kyne, 2007) [Fig. 2(c)]. Further verification of this species in the westernAtlantic Ocean is required.

Mobula mobular is an offshore, epipelagic species that occasionally frequentsshallow coastal waters. Some authors consider this species to be endemic to theMediterranean Sea (Notarbartolo-di-Sciara & Bianchi, 1998), whereas others suggestthat its range extends to waters of the eastern Atlantic Ocean (Scacco et al., 2009).Reports of this species encompass Ireland in the north, Senegal in the south andremote offshore islands including the Azores, Canary Islands, Cape Verde Islandsand Madeira (Santos et al., 1997; Wirtz et al., 2008) [Fig. 2(c)]. The species haseven been suggested to occur in waters off Cuba, the U.S.A. and India (Froese &Pauly, 2010; Zacharia & Kandan, 2010). Many of these records need to be verifiedas they are unconfirmed and misidentification cannot be excluded because of theclose morphological resemblance with M. japanica (Notarbartolo-di-Sciara, 1987b

Notarbartolo-di-Sciara et al., 2006). Further taxonomic description of both M. mob-ular and M. japanica is likely to aid field workers, particularly as the separationof both species is only based on the observation of one specimen of M. mobular(Panceri & De Sanctis, 1869; Notarbartolo-di-Sciara, 1987b). An author survey ofMobula images taken by recreational divers and accessible on the internet indicatesthat this species occurs at many sites in the Mediterranean Sea (e.g. Crete, Corsicaand Sardinia) for which there are no formal scientific records.

P O P U L AT I O N T R E N D S

The current status of each mobulid species is unclear, as research into popula-tion trends is only in its infancy. Understanding the status of these wide-rangingspecies across their entire distributions remains a challenge. Estimates of the worldglobal catch of mobulids have increased from c. 900 t in 2000 to >3300 t in 2007(FAO, 2009; Lack & Sant, 2009). Despite the increasing catches worldwide, dramaticdeclines in mobulid catches have been documented in some areas (e.g. Philippines:Alava et al., 2002), suggesting serial depletions through over-fishing. There are nosolid quantitative data to determine long-term population trends with the exceptionof M. hypostoma in the Atlantic Ocean region, which is thought to be increasingbased on trawl surveys conducted since 1989 (Bizzarro et al., 2008).

The use of photo-ID methodology has enabled minimum estimates of M. alfredipopulations at several locations around the world (Kashiwagi et al., 2011). Currentlocal population sizes of M. alfredi in Hawaii indicate that up to 230 are resident tothe island of Maui within one sampling period of c. 3 months (Deakos et al., 2011).In Mozambique, the annual population estimates of M. alfredi vary between 149and 454, with a total population estimate of 890 (Marshall et al., 2011c) (Table I).Although all current estimates of population size are for M. alfredi, minimum num-bers of individuals identified at aggregation sites exist for M. birostris (Kashiwagiet al., 2011). Without significant natural markings on which to base photo-ID studies,efforts to quantify numbers of Mobula spp. are effectively limited to fisheries data,aerials surveys and studies that employ conventional tags. Such approaches have yetto produce reliable population estimates for these species.



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M OV E M E N T S A N D AG G R E G AT I O N S

Movement patterns and swimming capacities of most mobulid species are poorlyunderstood. All mobulids are believed to undertake relatively large-scale movements,travelling from one productive area to another, and some species aggregate at specificlocations (Notarbartolo-di-Sciara, 1988; Celona, 2004; Couturier et al., 2011). Whilemembers of each species may be seen alone, or with a few other individuals, mostspecies have been observed travelling and feeding in schools. Schools may contain afew to hundreds of individuals and aggregate seasonally in large numbers at differentlocations throughout their ranges (Table I). Predictable aggregations of mobulidsare mainly associated with local productivity and food availability (Notarbartolo-di-Sciara, 1988; Celona, 2004; Sleeman et al., 2007; Dewar et al., 2008; Marshall,2009; Anderson et al., 2011; Couturier et al., 2011; Marshall et al., 2011c).

The aggregative behaviour of Mobula spp. remains largely understudied, withmost information on potential aggregation drawn from fisheries data in which somespecies are caught more frequently during particular seasons (e.g. Gulf of Califor-nia: Notarbartolo-di-Sciara, 1987a; Mumbai: Raje & Zacharia, 2009). The seasonaloccurrence of M. munkiana in winter in the Gulf of California and M. japanica andM. tarapacana in summer appears to be linked to the seasonal abundance of differenttypes of zooplanktonic prey (Notarbartolo-di-Sciara, 1988). Mobula thurstoni is theonly species known to occur all year around at this location (Notarbartolo-di-Sciara,1988). Smaller individuals are mostly caught during winter suggesting temporalsize segregation of this species in the Gulf of California (Notarbartolo-di-Sciara,1988). Mobula mobular seasonally aggregates between late spring and summer inthe Messina Strait, central Mediterranean Sea (Celona, 2004; Canese et al., 2011).There are also suggestions that the species undertakes seasonal north–south migra-tions within the Mediterranean Sea in relation to water temperature or productivity(Hemida et al., 2002; Celona, 2004; Scacco et al., 2009).

The limited information available on the movement of Mobula spp. includes onepublished study of M. mobular (Canese et al., 2011) and one conference abstract onM. japanica (Freund et al., 2000). Satellite tag data for three M. mobular individualsin the Messina Strait, central Mediterranean Sea, have shown that they spent >80%of their time within the upper 50 m of the water column and dived to 700 m depth(Canese et al., 2011). One tagged individual travelled 278 km in 60 days, while thetwo others travelled 298 and 337 km in 120 days (Canese et al., 2011). Using acous-tic telemetry techniques in the southern Gulf of California, M. japanica has been

Fig. 2. Distributions of mobulid species. (a) Distributions of the two known species of manta rays, Mantaalfredi ( ) and Manta birostris ( ). This figure incorporates data from Kashiwagi et al. (2011) and thenew IUCN Red List assessment for these species (Marshall et al., 2011a, b). In some regions, both speciesoccur close to one another and are colour-coded accordingly ( ) [Kashiwagi et al. (2011) provides furtherdetail about sympatric and allopatric occurrence]. (b) Distribution of wide-ranging species of Mobula:Mobula japanica ( ), Mobula tarapacana ( ) and Mobula thurstoni ( ); adapted and modified fromClark et al. (2006a, b) and White et al. (2006b). (c) Distribution of regionally endemic species ofMobula: Mobula eregoodootenkee ( ), Mobula hypostoma ( ), Mobula kuhlii ( ), Mobula mobular( ), Mobula munkiana ( ) and Mobula rochebrunei ( ); adapted and modified from Pierce & Bennett(2003), Bizzarro et al. (2006, 2007, 2008), Notarbartolo-di-Sciara et al. (2006) and Valenti & Kyne(2007). Question marks indicate some uncertainty about the identity of M. mobular in the North AtlanticOcean, and M. kuhlii off Java, Indonesia.



shown to move relatively fast, travelling up to 50 km in 24 h with swim speeds ofup to 8·3 km h−1 (Freund et al., 2000). Tagged animals spent most of their timeat relatively shallow depths, with occasional deeper excursions to 445 m (Freundet al., 2000).

Manta spp. tend to form predictable aggregations. Aggregations and movements inboth M. alfredi and M. birostris species are associated with seasonal food availability,current circulation patterns, seawater temperature, mating behaviour and cleaning-station visits (Sleeman et al., 2007; Dewar et al., 2008; Luiz et al., 2009; Andersonet al., 2011; Couturier et al., 2011). Over the past few years, movements and swim-ming behaviour of Manta spp. have been documented at several locations (Dewaret al., 2008; Kashiwagi et al., 2008; Couturier et al., 2011; Deakos et al., 2011;A. Marshall, J. Holmerg, J. M. Brunnschweiler & S. J. Pierce, unpubl. data). Mantaspp. are regularly present around shallow-reef cleaning stations and coastal feedinggrounds during daylight hours and potentially move to deeper and offshore watersat night (Dewar et al., 2008; Marshall et al., 2009; Clark, 2010; Anderson et al.,2011).

Long-term sighting records of M. alfredi at established aggregation sites suggestthat this species is more resident than M. birostris, preferring inshore tropical watersand exhibiting relatively small home ranges and movements (Homma et al., 1999;Dewar et al., 2008; Kitchen-Wheeler, 2008; Anderson et al., 2011; Couturier et al.,2011; Deakos et al., 2011; Marshall et al., 2011c). Using acoustic telemetry, vanDuinkerken (2010) found that M. alfredi can travel up to 70 km in a day and photo-ID studies have shown that this species undertakes seasonal migrations of up to500 km between known aggregation sites (Homma et al., 1999; Kashiwagi et al.,2010; Kitchen-Wheeler, 2010; Anderson et al., 2011; Couturier et al., 2011). In theHawaiian Islands, M. alfredi appears not to travel between Maui and Hawai’i Islands,only 49 km apart, despite intensive monitoring between 2005 and 2009 in both places(Clark, 2010; Deakos et al., 2011). Movements of M. alfredi may be restricted hereby bathymetric features or regional circulation patterns (Deakos et al., 2011).

Manta birostris is a larger, more oceanic and probably more migratory speciesthan M. alfredi, with individuals regularly sighted around offshore islands, oceanicseamounts and submarine ridge systems (Yano et al., 1999; Rubin, 2002; Marshallet al., 2009; Kashiwagi et al., 2010, 2011). In addition, rare or seasonal sightingsof M. birostris at locations such as northern New Zealand (Duffy & Abbott, 2003),southern Brazil (Luiz et al., 2009), the Azores and the Similan Islands (A. Mar-shall, pers. obs.), and the eastern coast of the U.S.A. (Bigelow & Schroeder, 1953)suggest that this species undergoes extensive migrations. Preliminary pop-off satel-lite tag studies have recorded broad-scale movements of >1000 km (Rubin et al.,2008; A. Marshall, J. Holmerg, J. M. Brunnschweiler & S. J. Pierce, unpubl. data).Together with international photo-ID projects, these data have so far revealed littleinterchange between regional populations, and whether M. birostris crosses oceanbasins is still unknown.

R E P RO D U C T I O N

The Mobulidae, as all chondrichthyans, employ internal fertilization to reproduce(Wourms, 1977; Carrier et al., 2004; Pratt et al., 2005). Data on the time for claspersto extend past the pelvic fins, together with observation of mating behaviour, suggest



that male M. alfredi mature at 3–6 years in Hawaii (Clark, 2010). Field observationsin Mozambique and Japan indicate a complex mating process for both Manta species,which involves a ritualized sequence of chasing, biting, copulating, post-copulationholding and separation (Yano et al., 1999; Marshall & Bennett, 2010b). Matingbehaviour in the wild has rarely been observed and so far has only been documentedfor M. alfredi and M. birostris (Yano et al., 1999; Marshall & Bennett, 2010b).

Mobulids are aplacental viviparous (Gill, 1908; Wourms, 1977; Dulvy & Reynolds,1997). Embryos develop in the uterus, where they initially feed on the yolk and arethen nourished with protein and lipid-rich histotroph or uterine milk, secreted byuterine villi (Wourms, 1977; Dulvy & Reynolds, 1997; Compagno & Last, 1999).The gestation period of most species is unknown, but the reproductive cycle is likelyto last for c. 1 year, with occasional resting period of ≥2 years between pregnan-cies (Notarbartolo-di-Sciara, 1988; Marshall & Bennett, 2010b). Marshall & Bennett(2010b) reported a gestation period in M. alfredi of c. 1 year in the wild. Similargestation periods were observed under captivity in the Churaumi Aquarium, Oki-nawa, Japan, where a captive M. alfredi gave birth four times, after 366–374 daysof gestation (Okinawa Churaumi Aquarium, 2010). All mobulid species normallygive birth to a single pup, but may produce twins on rare occasions (Gill, 1908;Coles, 1913; Notarbartolo-di-Sciara, 1987b; Marshall & Bennett, 2010b). Data onsize at birth and size at maturity for each species are summarized in Table I.

AG E , G ROW T H A N D L O N G E V I T Y

Mobulids are widely presumed to be long lived and slow growing, in keeping withtheir relatively large sizes and low reproductive rates (White et al., 2006a; Deakoset al., 2011). This hypothesis remains largely untested. As yet, there have been nopublished studies on the ageing and growth of Mobula spp. or Manta spp. despitetheir prevalence in fisheries. This dearth of information may be partially explainedby the apparent difficulty in ageing these species. The conventional ageing techniqueapplicable to most elasmobranch species is to section vertebral centra in the thoracicregion (Cailliet et al., 2006). Generally, growth-band pairs are visible to a greater orlesser degree within centra obtained from the anterior vertebrae of sharks (Goldman,2004) and posterior vertebrae of batoids (White et al., 2001; Pierce & Bennett, 2010).It appears that at least some mobulid species have a highly derived vertebral struc-ture that makes it difficult to use this technique (Cuevas-Zimbron, 2007). Dissectionsof M. alfredi (S. Pierce, unpubl. obs.) and M. birostris (A. Marshall, unpubl. obs.)suggest that Manta species do not have obvious calcified centra. Mobula japanicaspecimens from Mexico have been successfully aged based on band-pair counts fromcaudal vertebrae with the maximum number of 14 band pairs found in a female of2300 mm WD (Cuevas-Zimbron, 2007; Cuevas-Zimbron et al., 2008).

Unlike many elasmobranch species, for which maximum age is modelled fromgrowth curves, long-term diver visits to some areas where Manta spp. are com-monly sighted provide field observations of minimum longevity. Photo-ID surveysof M. alfredi off Japan and Hawaii indicate that this species can live for at least30 years (Ito, 1987; 2000; Kashiwagi et al., 2008; Clark, 2010; Marshall et al.,2011a) while individuals of M. birostris have been re-sighted using photo-ID upto 20 years after their initial identification (Marshall et al., 2011b). These observa-tions place both Manta species amongst the longest-lived batoids (Smith et al., 2007;



Pierce & Bennett, 2010) and, as the age at first identification of these animals wasunknown, actual longevity is likely to be exceptional within this group.

F E E D I N G

Behaviour and strategiesDespite regular observations of feeding activities, the feeding behaviour and strate-

gies of mobulids have only been cursorily described. As in all planktivorous elas-mobranchs, mobulids possess gills modified into complex rigid sieving plates forfiltering plankton (Bigelow & Schroeder, 1953; Cortes et al., 2008). Descriptionsof these branchial sieve plates can be found in Bigelow & Schroeder (1953) for M.birostris and M. hypostoma. When feeding, mobulids are generally observed to swimforward with an open mouth creating a passive water flow through the gill-rakerapparatus, a behaviour referred to as ram filter feeding (Sanderson & Wassersug,1990; 1993; Cortes et al., 2008). Cephalic lobes are used to guide water into themouth, where food particles then get sieved by the gill rakers before the waterexits the oropharyngeal cavity through the gill slits (Coles, 1916; Paig-Tran et al.,2011). Anecdotal reports on feeding behaviour of Manta spp. reveal varying for-aging strategies between locations, most likely related to prey density and foragingefficiency (Coles, 1916; Deakos, 2010a). For example, hundreds of M. alfredi havebeen reported to aggregate in the Maldives where they perform chain feeding, fol-lowing each other in a circling movement from surface to bottom creating a cyclonicmotion (Law, 2010). Observations of feeding behaviour in localized looping motionand bottom skimming were also made at several Manta spp. aggregation sites, high-lighting the adaptive responses of these animals according to prey distribution in thewater column (Deakos, 2010a; Osada, 2010).

DietStomach content analyses indicate that mobulids feed on zooplankton and small

fishes (Whitley, 1936; Bigelow & Schroeder, 1953; Notarbartolo-di-Sciara, 1988;Celona, 2004). Those studies highlight that several mobulid species appear to targethigh abundances of zooplankton found seasonally at certain locations (Notarbartolo-di-Sciara, 1988; Celona, 2004; Sampson et al., 2010). Aggregations of several Mob-ula species in the Gulf of California coincide with the peak abundance of theeuphausiid Nyctiphanes simplex (Hansen 1912), and this prey item dominates thestomach contents of several Mobula spp. caught in that region (Notarbartolo-di-Sciara, 1988). Mobula munkiana is the only species in this location that is presentonly during winter, feeding exclusively on mysids, Mysidium spp., the most abundantplanktonic prey available at this time of the year (Notarbartolo-di-Sciara, 1988). Sim-ilarly, M. mobular feeds in areas where the euphausiid Meganychtiphanes norvegicais in high density, and stomach content analysis confirmed M. norvegica as thedominant prey item (Celona, 2004). Mobula thurstoni stomach contents in the Gulfof California were dominated by either mysid shrimps or euphausiids, although thetwo types of prey were never found together in the same stomach (Notarbartolo-di-Sciara, 1988). Mobula thurstoni mainly feed on N. simplex during summer, when itis most abundant, and on Mysidium spp. during winter. These observations suggestthat M. thurstoni is able to adapt and shift diet, according to the dominant foodavailable in a particular location.



Results from stomach content analyses corroborate stable isotope analyses thatindicate that Mobula species mainly feed on zooplankton (Sampson et al., 2010;Borrell et al., 2011). These two stable isotope studies are to be interpreted with cau-tion as calculations involving the trophic enrichment factor could be inappropriate(Hussey et al., 2010). Isotopic values in tissues of M. japanica and M. thurstoniare relatively stable, indicating a consistent diet across individuals (Sampson et al.,2010). Borrell et al. (2011) found that Mobula species caught in fisheries off Ver-vaval, north-west India, had stable-isotope signatures consistent with those expectedfor zooplankton feeders. It is not possible to determine which Mobula spp. is referredto in that study as the species name, M. diabolus, is a synonym used to designateseveral Mobula species (Notarbartolo-di-Sciara, 1987b). Data on the diet and trophicposition of species can contribute to improve ecological understanding and can helpsupport conservation management plans for areas where the temporal and spatialabundance and distribution of prey are understood.

NAT U R A L M O RTA L I T Y

Natural predation on the Mobulidae appears to be low and opportunistic withsharks probably the most common cause. Non-fatal shark-inflicted injuries are reg-ularly observed on manta rays (Homma et al., 1999; Marshall & Bennett, 2010a;Deakos et al., 2011) and occasionally on mobulids (A. Marshall, unpubl. obs.). Thefrequency of shark attacks on M. alfredi individuals appears to differ among loca-tions, with 76% of the identified population in Mozambique bearing a shark-inflictedinjury (Marshall & Bennett, 2010a), compared with 24% off Maui, Hawaii (Deakoset al., 2011). Attackers of M. alfredi are thought to be primarily larger sharks such asgrey reef sharks Carcharhinus amblyrhynchos (Bleeker 1856), silvertip sharks Car-charhinus albimarginatus (Ruppell 1837), bull sharks Carcharhinus leucas (Muller& Henle 1839), tiger sharks Galeocerdo cuvier (Peron & LeSueur 1822) and greatwhite sharks Carcharodon carcharias (L. 1758) (Marshall & Bennett, 2010a). Gale-ocerdo cuvier and C. amblyrhynchos have been observed attacking and feeding onManta spp. (Ebert, 2003; Marshall & Bennett, 2010a; Deakos et al., 2011). The rateof delayed mortality induced by shark attacks is unknown, but physical injuries andthe redirection of energy into wound healing may affect reproduction in M. alfredi(Marshall & Bennett, 2010a). In addition, remains of unidentified mobulids havebeen found in the stomach contents of C. leucas (Cliff & Dudley, 1991). Killerwhales Orcinus orca are also known to occasionally feed on mobulids (Fertl et al.,1996; Homma et al., 1999; Visser & Bonoccorso, 2003).

THREATS

Mobulids are affected by numerous human activities including directed fishing,incidental capture as by-catch, entanglement in marine debris and boat strikes.Mobulids are easy to target in fisheries because of their large size, aggregativebehaviour, predictable habitat use and lack of human avoidance. Mobulid productsare valuable items in international trade markets. Their gill rakers are particularlysought after and are used in Asian medicinal products (Zhongguo yao yong dongwu zhi xie zuo zu bian zhu, 1983; Shen et al., 2003; Rajapackiam et al., 2007a).



This market has resulted in directed fisheries for nearly all mobulid species. Arti-sanal fisheries also target both genera for food and local products (White et al.,2006a; Essumang, 2009, 2010; Marshall et al., 2011c). Mobulids are also incidentalby-catch in both large-scale fisheries and small netting programmes such as shark-control bather-protection nets (Paterson, 1990; Krogh & Reid, 1996; Young, 2001;Sumpton et al., 2011).

As a result of sustained pressure from directed fishing and by-catch, certainsub-populations have rapidly declined including in India, Indonesia, Mexico, thePhilippines and Mozambique (Anon, 1997; Alava et al., 2002; Mohanraj et al., 2009;Marshall et al., 2011a, b). Particularly threatening to mobulid populations is the ten-dency for fisheries to target individuals in critical habitats or aggregation sites, wherethey can be caught quickly in large numbers. On the basis of seasonal abundance ataggregation sites and the size of fisheries catches, regional populations appear to berelatively small, and localized declines are unlikely to be significantly mitigated byimmigration and reproduction. This situation is exacerbated by the conservative lifehistory of these rays, which constrains their ability to recover from a depleted state.

D I R E C T E D F I S H E R I E S

Fishing methodsMobulids are currently killed or captured by a variety of methods including har-

pooning, netting and trawling. Gillnet fisheries are the most common and efficientmethod, with each haul taking large numbers of individuals (Alava et al., 2002).Target fisheries for M. thurstoni in Kerala State, south-west India, can capture up to65 mobulids per gillnet per haul, extrapolating to thousands of fishes caught annu-ally (K. Bineesh, pers. comm.). Gillnets are widely used in most areas where largenumbers of mobulids are landed, including Indonesia (White et al., 2006a; White &Dharmadi, 2007), the Gulf of California (Bizzarro et al., 2009), India (Rajapackiamet al., 2007a; CMFRI, 2009), western Africa (Essumang, 2010) and eastern Africa(A. Marshall & S. Pierce, unpubl. obs.). In some regions, mobulids are directly fishedin large trap nets set in major migratory channels such as in the Tangkoko NatureReserve in the Manado region of North Sulawesi, Indonesia (Anon, 1997). Someartisanal fisheries still employ more traditional techniques to capture mobulids suchas harpoons, hand spears, gaff hooks and hook and lines (Alava et al., 2002). Har-pooning is commonly used to catch both species of Manta as well as M. kuhlii insouthern Mozambique for local consumption (A. Marshall, unpubl. obs.). In north-east India, seasonal organized harpoon fisheries for Mobula spp. occur in AndhraPradesh State and the Indian Union Territory of Lakshadweep, north of the Maldives(Pillai, 1998). Although the technique does not appear to be widely applied, thecapture of mobulids by trawlers has been reported in both the Kerala region, India(Nair, 2003), and Tanzania (Bianchi, 1985; Jiddawi & Stanley, 1999).

Targeted speciesAll mobulids are sought after and targeted, generally captured using

non-discriminatory methods such as gillnets. Any mobulid species present withina fishing area tend to be exploited. In Indonesia, for example, mobulid landingsincluded all species known to occur in the area (i.e. M. japanica, M. tarapacana,M. kuhlii, M. thurstoni, M. alfredi and M. birostris) (Dewar, 2002; White &



Dharmadi, 2007). Similarly in the Gulf of California, 94% of the number of elasmo-branchs caught consisted of the four Mobula species present in the area, M. thurstoni,M. japanica, M. tarapacana and M. munkiana (Notarbartolo-di-Sciara, 1987b).

Value and useDespite these intense fisheries, mobulid meat is not highly valued for consump-

tion. It is often sold at relatively low prices or used as shark bait. Only the branchialfilaments are highly prized and are sent to Asia (Booda, 1984; Dewar, 2002; Rubin,2002; White et al., 2006a; Rajapackiam et al., 2007a; Mohanraj et al., 2009). Forexample, mobulids in Chennai, south-east India, are sold on the seashore. Gill rak-ers are then removed washed and dried before being sold for comparatively highprices to traders for export to Hong Kong, Singapore, Thailand, Malaysia and China(Rajapackiam et al., 2007a; Mohanraj et al., 2009). In Sri Lanka, the trade for mob-ulid gill rakers has become more lucrative than the shark-fin trade (Hilton, 2008).

Reported catchesAlthough a few studies have focused on quantifying the catch of mobulids in par-

ticular regions (e.g. eastern Indonesia, the Philippines and India), such reports remainscarce. The few studies that have been conducted project considerable numbers, with>1000 rays caught per year in some areas (Alava et al., 2002; Dewar, 2002; Whiteet al., 2006a).

Species of Mobula and Manta are caught in large numbers in Indian waters,raising concerns for the sustainability of the local targeted fishing practices (Fig. 3).The annual catch of elasmobranchs in India is currently estimated at c. 70 000 tyear−1 (Banerjee et al., 2008), representing 10% of the world annual catch (FAO,2009; Lack & Sant, 2009). Surveys revealed that between 2001 and 2006, 5% of theelasmobranch biomass caught along the Chennai coast were Mobula spp. (Mohanrajet al., 2009). Over 340 t of Mobula spp. can be caught at a single landing station inone fishing season, as was recorded at Nagapattinam from January to March 2009(CMFRI, 2009). Substantial landings can also occur in a single day. For example,c. 33 t of rays were landed in one day in July 2009 at the Tuticorin fishery harbour,with Mobula spp. (referred to as M. mobular but probably M. japanica) constituting11% of the catch (Zacharia & Kandan, 2010).

Thousands of mobulids are caught annually in Indonesian waters leading to sev-eral hundred tonnes of mobulid products. Catch of mobulids by targeted fisheriesalong eastern Indonesia was estimated to be 4110 individuals annually, equivalent toa biomass of 544 t and representing 38% of the total batoid fishery biomass (Whiteet al., 2006a; White & Dharmadi, 2007). On a smaller scale, a report of the fisheryaround Alor Island, south-east Indonesia, revealed that the recent demand for mob-ulid products in the Asian market has led to increased fishing effort on the fishesthrough improvement of fishing techniques and the equipment used (Dewar, 2002).Annual catch in this region is estimated to be 1500 individuals (range 1050–2400Manta spp.) (Dewar, 2002). Additionally, a study of the catch composition in NorthSulawesi, Indonesia, between March 1996 and February 1997 reported 1424 Mantaspp. caught within that year. While this practice was temporarily banned, it re-startedillegally in late 1997 and fishing effort has moved to new unmonitored locations(Anon, 1997). Similar landings have been reported from other locations, such as the



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Fig. 3. Distribution map of recent and current fisheries and by-catch locations of mobulids. All directed andtrade-driven fisheries on mobulids: 1 Mexico, La Paz (Booda, 1984; Notarbartolo-di-Sciara, 1987a;Rubin, 2002; Bizzarro et al., 2009; Erisman et al., 2010; Sampson et al., 2010; Cartamil et al., 2011), 2

Ecuador (M. Harding, pers. comm.), 3 Peru (M. Harding, pers. comm.), 4 Mediterranean Sea, coast ofAlgeria (Hemida et al., 2002), 5 Senegal (Cadenat, 1958; Marine Megafauna Association, pers. comm.),6 Ghana (Essumang, 2009; 2010), 7 Mozambique (A. Marshall, unpubl. obs.), 8 Tanzania (Bianchi,1985; Jiddawi & Stanley, 1999), 9 India, Veraval region (Sivaprakasam, 1964; Kunjipalu & Boopen-dranath, 1981; Said Koya et al., 1993; Pillai, 1998; Borrell et al., 2011), 10 India, Mumbai region (Rane,2002; Raje & Zacharia, 2009), 11 India, Karwar (Pillai, 1998), 12 India, Union Territory of Lakshad-weep (Said Koya et al., 1993; Pillai, 1998), 13 India, Kerala region, Vizhinjam (Pillai, 1998; Nair, 2003;Bineesh, pers comm.), 14 India, Gulf of Mannar and Tuticorin (Rajapackiam et al., 1990; Rajapack-iam & Balasubramanian, 1994; Pillai, 1998; Abdussamad et al., 2006; Arumugam & Balasubramanian,2006; Zacharia & Kandan, 2010), 15 India, Nagapattinam (CMFRI, 2009), 16 India, Pondicherry (Pillai,1998), 17 India, Chennai (Said Koya et al., 1993; Pillai, 1998; Rajapackiam et al., 2007a, b; CMFRI,2009; Mohanraj et al., 2009; Kizhakudan et al., 2010), 18 Sri Lanka, Marissa (P. Hilton, pers. comm.),19 Thailand, north-west coast (Compagno & Last, 1999), 20 Indonesia, Java (Barnes, 2005; White et al.,2006a; White & Dharmadi, 2007), 21 Indonesia, Bali (White et al., 2006a; White & Dharmadi, 2007),22 Indonesia, East Lombok (White et al., 2006a; White & Dharmadi, 2007), 23 Indonesia, Alor Island(Dewar, 2002), 24 Indonesia, North Sulawesi (Anon, 1997), 25 Philippines, Bohol Sea (Dolar, 1994; Alavaet al., 2002). All locations where mobulids have been reported in by-catch fisheries: 1 Mexico, Gulf ofCalifornia (Sampson et al., 2010), 2 Mexico, Gulf of Tehuantepec (Sarmiento-Nafate et al., 2007), 3

east Pacific Ocean, Isla de la Plata, Ecuador (M. Harding, pers. comm.), 4 south-eastern Brazil (Zerbini& Kotas, 1998; Amorim et al., 2002; Casas et al., 2006), 5 western Atlantic Ocean, south-eastern U.S.(Trent et al., 1997; Carlson & Lee, 2000; Beerkircher et al., 2002, 2009; Baremore et al., 2007; Schwartz,2011), 6 Mediterranean Sea (Bradaii & Capape, 2001; Celona, 2004; Akyol et al., 2005; Scacco et al.,2009; Akyol & Ceyhan, 2011; Storai et al., 2011), 7 European fisheries off Mauritania (Zeeberg et al.,2006), 8 European purse-seine tuna fishery (Rey & Munozchapuli, 1992; Menard et al., 2000; Amandeet al., 2010), 9 South African coast, Natal (Dudley & Cliff, 1993; Dudley & Gribble, 1999; Young,2001), 10 Mozambique (Olsen et al., 2009; A. Marshall & S. Pierce, unpubl. obs.), 11 west Indian Ocean(Romanov, 2002), 12 Gulf of Aden (Bonfil & Abdallah, 2004), 13 The Gulf (Henderson & Reeve, 2011;Moore, 2011), 14 west coast of Thailand and Myanmar (A. Marshall, unpubl. obs.), 15 eastern Indonesia(Dharmadi et al., 2008; Blaber et al., 2009), 16 Philippines (Dolar, 1994; P. Hilton, pers. comm.), 17 PapuaNew Guinea (C. Rose, pers. comm.), 18 eastern Australia (Paterson, 1990; Krogh & Reid, 1996; Harryet al., 2011; Sumpton et al., 2011), 19 northern New Zealand (Paulin et al., 1982), 20 central-westernPacific tropical purse-seine fishery (Coan et al., 2000), 21 Chinese Pacific longline fishery (Dai & Zhu,2008). , The by-catch area for the corresponding fishery number.



Pamilacan Island in the Philippines where up to 1000 mobulids, particularly Mantaspp. and a few species of the genus Mobula, are harvested each year by directedfisheries (Alava et al., 2002).

Status of current fisheriesAlthough targeted mobulid capture has been banned in a few areas, many direct

fisheries remain (Fig. 3). Targeted fisheries are prevalent at many locations aroundIndia, particularly along the coast of Chennai, Tuticorin, Mumbai and Veraval, withinthe Union Territory of Lakshadweep and in the Andhra Pradesh and Kerala regions(Sivaprakasam, 1964; Said Koya et al., 1993; Rajapackiam & Balasubramanian,1994; Pillai, 1998; Nair, 2003; Rajapackiam et al., 2007a, b; CMFRI, 2009; Mohanrajet al., 2009; Zacharia & Kandan, 2010). Some of these fisheries are highly orga-nized and efficient, such as a new fishery formed along the Chennai coast that usesmechanized gillnets, following the high demand for mobulid products (Pillai, 1998;Rajapackiam et al., 2007a).

After years of exploitation, mobulid catches have begun to decline in mostfished regions suggesting serial depletions. Although fishing effort has been increas-ing, annual landing of rays (including Mobula spp.) has been declining in severalareas of India, including in the Kerala region (Nair, 2003) and along the Chennai(Mohanraj et al., 2009) and Tuticorin coasts (Zacharia & Kandan, 2010). Fisheriessurveys in Mumbai waters revealed that fishes were landed at a rate of 0·65 kg h−1

in 1990, but only 0·24 kg h−1 in 2004 (Raje & Zacharia, 2009). A dramatic reduc-tion in mobulid capture was also reported in eastern Indonesia (Dewar, 2002; Whiteet al., 2006a). Barnes (2005) reported that manta rays were historically fished byindigenous villagers in this region, with up to 360 animals caught in one year, butthe fishes became gradually scarcer until the catch rate reached zero.

Management and regulationsIn response to the decline in mobulid catches, some regions have banned directed

fisheries, established marine protected areas and enacted protection legislation forsome species (Fig. 4). The enforcement of those bans remains uncertain in manyregions and illegal capture of mobulids still occurs and remains unreported (Anon,1997, Alava et al., 2002, Bizzarro et al., 2009, Smith et al., 2009; Erisman et al.,2010). Moreover, many of the current management and interjurisdictional regulationsfor the Manta genus need to be reassessed, as most were put into place prior tothe re-description of the two species. Consequently, M. birostris is the only Mantaspecies mentioned and legally protected by some of this legislation (Fig. 4) and M.alfredi remains unprotected although this species more commonly occurs in tropicaland subtropical coastal areas and thus is widely exposed to fisheries. In regionswhere bans have successfully been enforced, such as in the Maldives, fisheries havebeen replaced with equally or more lucrative tourism industries (Anderson et al.,2010).

B Y- C AT C H

Mobulids are caught as by-catch in purse-seine and trawl fisheries as well as net-ting programmes throughout their distributions (Fig. 3). The most threatened mobulid



1 2

3

4

5

6

7 8

9

Fig. 4. Distribution map of current fishing bans, management regulations and specific species protection formobulids: 1 Hawaii, June 2009: introduction of criminal penalties and administrative fines for knowinglykilling or capturing Manta spp. within state waters, House Bill 366 signed into law as Act 092; 2 Mexico,May 2007: the Mexican official standard rules that regulate the shark and ray fisheries in Mexicanwaters, Pesca responsable de tiburones y rayas. Especificaciones para su Aprovechamiento, 2005. NormaOficial Mexicana Nom029-Pesc-2006, NOM 029 provides specific protection for Manta birostris, Mobulajapanica, Mobula thurstoni, Mobula munkiana, Mobula hypostoma and Mobula tarapacana in Mexicanwaters; 3 Ecuador, August 2010: ban on fishing, taking, keeping incidental catch for M. birostris,M. japanica, M. thurstoni, M. munkiana and M. tarapacana, La subsecretaría de recursos pesquerosconsiderando. Acuerdo 093; 4 Mediterranean, 1995: Mobula mobular protected under U.N. (Barcelona)Convention for the Protection Of The Mediterranean Sea Against Pollution Annexe II; 5 Republic ofMaldives, June 1995: export ban on ray products (preventing commercial fisheries). Regulation NoA-23/95; January 1996: export of ray skins prohibited Regulation No A-26/95 (of 15.7.95); June 2009:marine park areas created at two sites recognized as critical habitat for Manta alfredi ; 6 Western Australia,Manta spp. protected from any fishing (Fisheries Act) and disturbance and harassment (Environment andConservation Act) within marine parks only; 7 Philippines, March 1998: ban on the taking or catching,selling, purchasing and possessing, transporting and exporting of whale sharks (Rhincodon typus) andM. birostris. Fisheries Administrative Order 193 (implanted in 1998 then lifted in 1999 before being re-established in 2002. Mobula spp. are not protected under this ban); 8 Yap, Micronesia, 2008: protectedarea for Manta spp. and their habitat [includes 16 main islands and 145 islets representing 21 349 km2

(8243 square miles)]. Manta Ray Sanctuary and Protection Act of 2008. Bill No 7-69,D1. Yap State LawNo. 7-36 (note: not yet added to the Yap State Code, new Chapter 12 of title 18 of the Yap State Codeto be created see law 7-36 in Seventh State Legislature); 9 New Zealand, 2010: M. birostris and M.japanica fully protected within New Zealand waters. Wildlife Act 1953, Schedule 7A.

ray, M. mobular, is subject to a high mortality rate as a result of incidental capture(Notarbartolo-di-Sciara et al., 2006). This ray is frequently caught as a by-catch in theMediterranean Sea and eastern Atlantic Ocean (although this may be M. japanica)and is likely to be at unsustainable levels (Cadenat, 1958; Menard et al., 2000;Bradaii & Capape, 2001; Celona, 2004; Akyol et al., 2005; Notarbartolo-di-Sciaraet al., 2006; Scacco et al., 2009; Amande et al., 2010; Storai et al., 2011).

Tuna purse-seine fisheries are one of the main contributors to mobulid by-catchwith several species regularly caught in relatively large numbers. In the western



Indian Ocean, between 53 and 112 t of mobulids are caught each year in thesefisheries (Romanov, 2002). It appears that the largest species of mobulids (Mantaspp., M. mobular, M. tarapacana and M. japanica) are being caught by these off-shore fisheries. Globally distributed species such as Manta spp. and M. japanicaare regularly caught in most oceans, including the eastern Atlantic Ocean (Amandeet al., 2010), central-western Pacific Ocean (Coan et al., 2000), western Indian Ocean(Romanov, 2002), off Papua New Guinea (C. Rose, pers. comm.) and northern NewZealand (Paulin et al., 1982).

Gillnet fisheries also contribute to regular non-target landing of mobulid speciesat many locations. Reports of mobulid by-catch in these net fisheries have beenidentified from the Philippines (Dolar, 1994), Indonesia (White et al., 2006a) and offthe south-eastern U.S.A. (Trent et al., 1997; Carlson & Lee, 2000; Baremore et al.,2007) (Fig. 3). Industrial trawlers are also likely to have a considerable effect onmobulid species with up to 620 ‘manta rays’ caught per year by trawlers operatingoff the north-west African coast (Zeeberg et al., 2006). This report indicates thecatch to be M. birostris, but a photograph purporting to be a Manta sp. is actuallya Mobula, most likely M. tarapacana, and further consideration of the raw databy the authors suggests that all catches were Mobula specimens. Mobulid speciesare also occasionally caught in longline fisheries in the Atlantic Ocean (Rey &Munozchapuli, 1992; Trent et al., 1997; Carlson & Lee, 2000; Beerkircher et al.,2002, 2009; Baremore et al., 2007; Schwartz, 2011).

Mobulids are regularly recorded as incidental catches in shark-control nets offboth Australian and South African coasts (Paterson, 1990; Dudley & Cliff, 1993;Krogh & Reid, 1996; Dudley & Gribble, 1999; Young, 2001; Sumpton et al., 2011).Young (2001) reported that the KwaZulu-Natal shark-control nets caught 1191 manta(Manta spp.) and 440 devil rays (Mobula spp.) between 1981 and 2000, with upto three individuals caught per day. Mobulids constituted 12% of the total catchby number from these nets between 1981 and 1990, with a mean annual catch of66 individuals and an average mortality rate of 33%. Of the 440 Mobula caught,19 were identified as M. kuhlii, four as M. japanica and one as M. eregoodooten-kee, leaving over 94% of the catch unidentified to species (Young, 2001). Similarlyin Queensland, Australia, Sumpton et al. (2011) found that 93 mobulids from bothgenera were caught between 1992 and 2008 in shark-control nets, with a mortal-ity rate of 41% for Manta spp. and 89% for Mobula spp. Although incidentalcatch and by-catch might not represent a major threat to mobulid populations, itis an additional stressor on top of the targeted fisheries and other anthropogenicactivities.

OT H E R T H R E AT S

Cryptic threats such as mooring-line entanglement and boat strikes can also woundmobulids, decrease fitness and contribute to mortality (Marshall & Bennett, 2010a;Deakos et al., 2011). Many other threats have the potential to harm mobulids ata global scale such as habitat degradation, climate change, pollution, ingestionof microplastics and irresponsible tourism practices. Unmanaged, rapidly growingtourism (including in-water interactions and recreational boating traffic) is likely toaffect Manta spp. use of and visit rates to critical cleaning and feeding habitats(Anderson et al., 2010; Osada, 2010; Deakos et al., 2011).



Marine debris harms and kills a large number of marine animals through entan-glement and ingestion (Laist, 1987; Hiruki et al., 1993; Cliff et al., 2002; Derraik,2002; Page et al., 2004; Chaloupka et al., 2008; Boerger et al., 2010). Mobulids canbecome entangled, especially Manta spp., which are regularly observed hooked bylost fishing lines (Marshall & Bennett, 2010a; Deakos et al., 2011). In Maui, Hawaii,10% of the M. alfredi population has amputated or non-functioning cephalic fins,most likely caused from entanglement in monofilament fishing line (Deakos et al.,2011). Such debris may also greatly affect the fitness and survival of marine species,especially since many are already affected by other threats such as fishing (Cliffet al., 2002; Derraik, 2002).

Ingestion of plastic debris by sea turtles, seabirds, sharks and fishes has been welldocumented (Laist, 1987; Derraik, 2002), and can cause gastrointestinal blockages,ulcerations, internal perforation, malnutrition and death (Fry et al., 1987; Baird &Hooker, 2000; Mascarenhas et al., 2004; Boerger et al., 2010). Filter feeders, such asmobulids, may be repeatedly exposed to ingestion of microplastic debris mixed withintheir natural food source. For example, microplastic debris sampled in the NorthPacific Ocean central gyre has a similar particle size distribution as zooplankton andthe mass of debris can outweigh by about six times that of plankton (Moore et al.,2001). Planktivorous animals such as basking sharks Cetorhinus maximus (Gunnerus1765) and baleen whales have been observed feeding in areas where marine debriswas reported, and are at high risk of plastic ingestion (Choy & Adams, 1995; Gregory,2009). Additionally, traces of heavy metals such as platinum, mercury and arsenic,known to be highly detrimental to human health, were found in high concentrationsin tissues of mobulids off the Ghanaian coast (Essumang, 2009; 2010). Effects ofthese metals on mobulids are unknown.

It has been suggested that mobulids have low vulnerability to climate change com-pared with other sharks and rays, as they do not appear to have high habitat specificityor rigidity to adaptation (Chin et al., 2010). As zooplanktivores, however, they arelikely to be influenced by the change in the abundance, phenology and distributionof their planktonic food as the global oceans warm (Hays et al., 2005). Further,the local and regional oceanographic conditions that cause seasonal zooplanktonhotspots targeted by mobulids could alter under climate change (Richardson, 2008)and potentially influence migration pathways and timing.

USING ADVANCED TECHNOLOGY TO STIMULATE MOBULIDRESEARCH

The mobility, vast pelagic habit and taxonomic uncertainties surrounding mobulidsare major impediments to the gathering of information on the biology and ecologyof this group. The collection of biological, behavioural, ecological and physiologicalinformation on these species has traditionally been limited to studies of individualsin inshore and surface waters, during the day, and to dead specimens for anatomicalwork. With the recent demand for more information on these species and rapidadvances in technology, innovative techniques now allow researchers to examinenew aspects of mobulid habitat use, seasonality, abundance and migrations, with amore comprehensive insight into their world.



P O P U L AT I O N DY NA M I C S , H A B I TAT U S E A N D M OV E M E N T S

Photographic identificationPhoto-ID is a cost-effective and non-invasive approach in wide use for the long-

term study of animal populations and individual behaviours. This technique is effec-tive for individual identification of species that bear markings or scarring patternsthat remain consistent through time. With the advent of inexpensive digital camerasequipped with underwater housings, this technology has become standard gear forrecreational divers. With thousands of divers in the oceans at any one time, it ispossible to gather information on vulnerable species such as mobulids quickly andinexpensively. Photographic research is often supported by eco-tourism operatorsand the dive community and thus increases environmental education and awareness(Couturier et al., 2011). Photo-ID has been used to examine various aspects of thebiology and ecology of elasmobranchs (Castro & Rosa, 2005; Domeier & Nasby-Lycas, 2007; Bansemer & Bennett, 2008; Dudgeon et al., 2008) and recently, variousaspects of M. alfredi and M. birostris. For manta rays, long-term photo-ID surveyswith appropriate sampling design can allow for the estimation of local populationsizes (Deakos et al., 2011; Marshall et al., 2011c), the examination of visitation pat-terns to specific sites (Luiz et al., 2009; Kitchen-Wheeler, 2010; O’Shea et al., 2010),the detection of movements between aggregation sites (Homma et al., 1999; Kashi-wagi et al., 2010; Couturier et al., 2011) and population reproductive variables suchas gestation period, sex ratio and male maturity (Clark, 2010; Marshall & Bennett,2010b).

Paired-laser photogrammetry is a technique that can be combined with photo-IDto provide data on the structure of a population. This technique uses photographsto indirectly make measurements of an object or, in this case, animals’ morphomet-rics. Two parallel laser pointers mounted to a camera, project two beams of lightonto a targeted animal, creating a scale bar that can be used to extrapolate an esti-mated size of the individual (Deakos, 2010b; Rohner et al., 2011). When paired-laserphotogrammetry and photo-ID are used together, they provide valuable data on thesize structure of a population and could in the future enable the estimation of sizeat maturity (e.g. size of elasmobranch v. pregnancy or claspers development) andgrowth rate of a species (over multiple year surveys) (Deakos, 2010b; Marshall &Bennett, 2010b; Marshall et al., 2011c).

Most photo-ID databases are limited to a particular aggregation area, and thuscannot be used to answer questions associated with the large-scale movements ofManta spp. To assess movements across ocean basins, regional or global databasesare needed. A good example is the ECOCEAN global whale shark Rhincodon typusSmith 1828 database (www.whaleshark.org; Arzoumanian et al., 2005; Holmberget al., 2009), which has >35 000 photos of 3226 individuals. A similar ECOCEANglobal database for Manta spp. (www.mantamatcher.org) has just been launched.Increasing use of such databases in the future will facilitate studies of regionalpopulation size and exchange among Manta spp. aggregation sites, as has beeninvestigated in R. typus (Brooks et al., 2010).

Acoustic telemetryAcoustic telemetry involves the use of acoustic receivers to record the presence

of an animal equipped with an acoustic transmitter within the detection range of the



receiver. It has become an increasingly popular tool to monitor the occurrence pat-terns, home ranges, fine-scale movements and habitat use of various aquatic species,including elasmobranchs (Klimley et al., 1988; Heupel et al., 2004; Huveneers et al.,2006; Vaudo & Lowe, 2006). There are two main types of acoustic telemetry: (1) thepassive monitoring of the occurrence of a tagged animal within an area via an auto-mated array of acoustic receivers (Heupel et al., 2006) and (2) the active or manualtracking of movements of a tagged animal using a directional hydrophone and acous-tic receiver (Cartamil et al., 2003). To date, these techniques have only been used ina few studies investigating mobulids. Passive acoustic telemetry has a great potentialfor investigating site fidelity patterns, seasonal occurrence and habitat use at par-ticular monitored locations of mobulids (Dewar et al., 2008; Marshall, 2009; vanDuinkerken, 2010). In addition, seasonal movements among sites by tagged indi-viduals can be identified when several locations are being monitored simultaneously(Dewar et al., 2008). Active acoustic telemetry can provide information over a shortperiod of time on the fine-scale movements of animals, including horizontal distancetravelled, vertical movement in the water column and swimming speed of the trackedindividual (Freund et al., 2000).

Satellite telemetryRecent advances in the fields of wildlife tracking, remote sensing, geographic

information systems and ecological modelling now allow researchers to monitorremotely the position and environmental conditions of free-ranging animals as theymove undisturbed. In recent years, satellite-tracking technology has permitted obser-vations of some previously undescribed aspects of the lives of large marine ver-tebrates, including predatory fishes, marine birds, marine mammals and sea turtles(Weimerskirch et al., 2002; Block et al., 2011). To date, only one published study hasinvestigated movements and behaviour of a Mobula species using satellite telemetry(Canese et al., 2011), and some preliminary results have been presented at confer-ences for Mobula spp. (Freund et al., 2000) and Manta spp. (Rubin et al., 2008;Jaine et al., 2011; F. R. A. Jaine, L. I. E. Couturier, S. J. Weeks, A. J. Richardson,K. A. Townsend & M. B. Bennett, unpubl. data; A. Marshall, J. Holmerg, J. M.Brunnschweiler & S. J. Pierce, unpubl. data). These studies have highlighted thepotential for satellite telemetry to provide data on the migratory routes and distancestravelled by individuals in a definite period of time. Satellite tags can also recordinformation on the swimming behaviour and dive profile of tagged rays (Freundet al., 2000; et al., 2010; Canese et al., 2011; Jaine et al., 2011; F. R. A. Jaine, L.I. E. Couturier, S. J. Weeks, A. J. Richardson, K. A. Townsend & M. B. Bennett,unpubl. data).

P O P U L AT I O N S T RU C T U R E

GeneticsMolecular analysis has become a standard tool to investigate populations of elas-

mobranchs (Heist, 2004a; Portnoy, 2010). An advantage of genetic approaches isthat only a small tissue sample collected from living or deceased animals con-tains genomic information that can be investigated for genetic relationships at theindividual, population, species and higher taxonomic levels (Heist, 2004a). Geneticmethods will be used increasingly in the future to analyse population structure and



connectivity, aiding fisheries management and conservation decisions (Heist, 2004b;Ovenden et al., 2009, 2010). Although there are several population genetics stud-ies on the Mobulidae commenced or underway (Poortvliet et al., 2011), there arecurrently no published studies.

Age and growthAge and growth data are fundamental to future analyses of the sustainability of

mobulid fisheries and parameterization of demographic models. If the unique verte-bral structure of some species precludes obtaining age and growth data from fishedspecimens, as preliminary observations suggest, long-term photographic sight–resightstudies should focus on this topic. Coupling the deployment of portable in-water mea-surement systems, such as camera-mounted parallel lasers (Deakos, 2010b), withstudies of population ecology (Marshall et al., 2011c) to obtain growth rate andsurvivorship data are effective, if currently labour-intensive, means of estimatingthese parameters from field observations. The lack of field studies on Mobula spp.and the apparent lack of individually identifiable features in these species meanthat continuing efforts should be made to obtain vertebral structure data from fishedspecimens.

F E E D I N G E C O L O G Y

Most published data on the diet of mobulids are based on stomach-content analysisand have a number of limitations. It is often difficult or unethical to obtain stomachsfrom animals threatened or protected in many countries. This means that often onlya few stomachs are available for analysis, often from individuals washed up deadon the shore. Even in countries where they are the target of directed fisheries, itcan be difficult to obtain samples because of distrust in researchers. Once stomachshave been collected, their contents will only represent recent feeding, and individualprey items are often difficult to identify because of digestion, with hard body partsoften all that remain. Ideally, stomach contents are needed from different periods,locations, size classes and both sexes to describe adequately the diet of a species.

Fatty-acid and stable-isotope signatures are two indirect methods used to investi-gate the diet preference and trophic position of marine predators and have consider-able potential in elucidating more light on the diet of mobulids, if correctly applied.Both methods have the advantage of only requiring a relatively small amount oftissue collected in the field without killing the fishes. Importantly, because thesetechniques allow the collection of samples from more individuals, they can detectpotential shifts in the diet of a species and can contribute to understanding the dynam-ics behind animal movements (Kelly, 1999; MacNeil et al., 2005; Budge et al., 2006;Iverson, 2009).

Stable isotopesRatios of stable isotopes of nitrogen (15N:14N or δ15N) and carbon (13C:12C or

δ13C) in a consumer’s tissues are related to its assimilated food and are an indexof its trophic position in the ecosystem. δ15N and, to a lesser extent, δ13C, show apredictable stepwise enrichment with each increasing trophic level (Hobson, 1999;Kelly, 1999; Dahl et al., 2003; Herman et al., 2005; Perga et al., 2006). This tech-nique indicated that the trophic level of three mobulid species was between 3·4 and



3·6 (second level consumer) (Sampson et al., 2010; Borrell et al., 2011). Values ofδ13C provide information on the origin of the carbon entering the food web, whetherit is from marine, freshwater or terrestrial origin (Fry & Sherr, 1984) or garneredfrom benthic or pelagic feeding (Dahl et al., 2003). For example, based on theirknown diet, the origin of δ13C values of Mobula spp. was extrapolated to be fromthe pelagic and epipelagic environment (Borrell et al., 2011).

Fatty acidsDiets of a number of predators have been examined using fatty-acid analysis,

including species of marine mammal (Herman et al., 2005), seabird (Dahl et al.,2003) and elasmobranch (Semeniuk et al., 2007). Fatty acids are the principal con-stituents of most lipid molecules and degrade little when ingested. In the higherpredator, fatty acids are generally assimilated intact and are mostly either useddirectly as an energy supply or are stored in adipose tissues (Iverson et al., 2004;Iverson, 2009). Hence, fatty-acid composition from prey will most likely directlyinfluence the fatty-acid composition of the stored fat of its consumer (Iverson et al.,2004; Budge et al., 2006; Iverson, 2009). Comparison of fatty-acid signatures of arange of potential prey species with its likely predators may provide dietary infor-mation beyond that revealed by gut-content analysis alone, such as shifts in diet andprey specialization. This method has not yet been applied to mobulid ray species,but could reveal new insights into possible prey specialization.

CONCLUSIONS

Although the diverse Mobulidae family appears to be a successful group in theirbroad distribution and relative abundance, rising human pressures are a major threatto most of these species. Collectively, catch reports suggest that a large number ofmobulids are being removed from the marine environment. Of particular concern isthat most catch by directed fisheries in many parts of the world remains unreportedand unregulated. Reproductive adaptations of mobulids place these species amongstthe least fecund elasmobranchs. This, combined with slow growth and an apparentlow natural predation, suggests that mobulids are not likely to be able to cope withhigh fishing mortality.

Movement and aggregation studies suggest that mobulids are highly mobile andhave the potential to travel large distances in relatively short periods of time. Fur-ther studies are needed to develop a more complete picture of the movements ofmobulid species to evaluate the exposure to and overlap with human activities suchas fisheries and pollution. Knowing the location of migration routes and aggregationsites can provide useful information for their protection through the establishmentof marine-protected areas (Game et al., 2009, 2010; Hobday et al., 2011). Althoughprotected areas have typically been fixed in time and space and prevented mostfisheries exploitation, migratory species that use particular aggregation sites couldbenefit from protection that would restrict harmful fishing practices at the time ofyear that mobulids visit. More work is needed to understand the reasons behindmobulid aggregations to underpin effective conservation and management models.

Fundamental questions still exist regarding the taxonomy and basic species-specificbiological variables such as reproduction and habitat requirements. Mobulids have



many common biological characteristics, although many aspects of their ecologyappear to be region and species-specific. Standardized techniques such as acousticand satellite telemetry and stable-isotope and fatty-acid analyses can be applied toall species and will provide new insight into the differences in the biology of thesespecies.

Aggregations of mobulids provide opportunities for researchers to access manyindividuals for tagging programmes, genetic studies, intraspecific behavioural inter-actions and non-invasive dietary analyses. Scientific collaboration is crucial in orderto generate estimates of global population status, and to standardize methodologiesrequired for data input in ecosystem models. Implementation of fisheries manage-ment is critical along with the concomitant need for management strategies capableof sustaining the Mobulidae into the future. To ensure their long-term sustainability,these slow-growing species demand greater scientific involvement in fisheries andconservation management.

The authors thank P. Bassett, K. Hogg, H. Mitchell, K. Stephens and C. Bustamante fortheir precious help with the creation of maps and fisheries information. The authors are alsograteful to P. Kyne, C. Rohner, M. Mauffrey and A. Gutteridge for their useful comments andadvice, to P. Nichols for his guidance on fatty acids work and to K. K. Binesh, P. McCann,C. Rose, M. Harding, G. Kodja, S. Yaha, I. Semesi, N. Michivo, C. Anderson, P. Hiltonand N. Iddawi for advice and information supplied. A.D.M. and S.J.P.’s work on this reviewwas supported by Save Our Seas Foundation, one anonymous donor, Ocean Revolution andCasa Barry Lodge, Mozambique. Support for this work which includes Project Manta, aresearch programme based at the University of Queensland, was provided by ARC LinkageGrant (LP110100712), ARC Future Fellowship (FT099172), Sea World Research and RescueFoundation Inc., EarthWatch Australia, Brother Pty Ltd, Lady Elliot Island Eco-Resort, MantaLodge and Scuba Centre, Project AWARE.

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couturier et al. 2012 mobulid review.pdf

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