The conserva tion and su s t a i n a bleuse of marine re sou rces is a high l i gh ted goal on a growing nu m ber of n a tional and intern a tional policy agen d a s .Un fo rtunately, ef fo rts to assess pro gress, as well as to stra tegi c a lly plan and pri o ri ti ze new marine conserva tion measu res, have be en hampered by thelack of a detailed, co m preh en s ive bi o ge o graphic sys tem to classify the oceans. Here we repo rt on a new global sys tem for coastal and shel f a reas: theMa rine Eco regions of the Wo rl d , or MEOW, a nested system of 12 realms, 62 provinces, and 232 eco regions. This sys tem provides considera bly bet ters pa tial resol u tion than earl i er global systems, yet it pre serves many common el em ents and can be cro s s - referen ced to many regional bi o ge o graphicclassifications. The design a tion of terre s trial eco regions has revol u ti o n i zed pri o ri ty set ting and planning for terre s trial co n serva tion; we anti ci pa tesimilar ben efits from the use of a coh erent and cred i ble marine sys tem .
Mapped classifications of patterns in biodivers i t y
h ave long been an important tool in fields from evo luti on a ry studies to con s erva ti on planning (Forbes 1856,Wa ll ace 1876, S pell erberg and Saw yer 1999, Lo u rie and Vi n cent 2004). The use of su ch sys tems (notably, the wi delyc i ted sys tem devel oped by Ol s on et al. [2001]) in broadscale con s erva ti on , h owever, has largely been re s tri cted to terre s-trial studies (Ch a pe et al. 2 0 0 3 , Ha zen and An t h a m a t ten2 0 0 4 , Hoe k s tra et al. 2 0 0 5 , Bu r gess et al. 2 0 0 6 , L a m oreux eta l . 2 0 0 6 ) . In the marine envi ron m en t , ex i s ting gl obal cl a s s i-f i c a ti on sys tems remain limited in their spatial re s o luti on .Some are incon s i s tent in their spatial covera ge or met h od-o l ogical approach . The few publ i c a ti ons that have attem ptedto use bi ogeogra phic regi on a l i z a ti on in gl obal marine con s erva ti on planning (e.g. , Kell eh er et al. 1 9 9 5 , Ol s on andDi n ers tein 2002) have been qu a l i t a tive , and have ex pre s s edcon cern abo ut the lack of an adequ a te gl obal cl a s s i f i c a ti on .
In the absen ce of com pelling gl obal covera ge , nu m erous regi onal cl a s s i f i c a ti ons have been cre a ted to meet regi on a lplanning need s . Th i s , of co u rs e , does not satisfy the need fora gl obal sys tem that is con s i s tent ac ross the many mari n erealms and coastal zon e s .
Bi ogeogra phic cl a s s i f i c a ti ons are essen tial for devel op i n geco l ogi c a lly repre s en t a tive sys tems of pro tected are a s , as re-qu i red by intern a ti onal agreem ents su ch as the Conven ti onon Bi o l ogical Divers i ty ’s Programme of Work on Pro tectedAreas and the Ramsar Conven ti on on Wet l a n d s . Ma ri n es p ace is sti ll gro s s ly underrepre s en ted in the gl obal pro tecteda reas net work (on ly abo ut 0.5% of the su rf ace area of t h eoceans is curren t ly pro tected ; Ch a pe et al. 2 0 0 5 ) , a fact thatadds urgency to the need for tools to su pport the scaling upof ef fective , repre s en t a tive marine con s erva ti on . The key ide au n derlying the term “repre s en t a tive” is the intent to pro tecta full ra n ge of bi od ivers i ty worl dwi de — gen e s , s pec i e s , a n d
Ma rk D. Spalding (e-mail: mspa l d i n g @ tn c . o rg ) , Za ch A . Ferd a ñ a , Jen n i fer Mol n a r, and James Robert son are co n serva tion sci en tists in The Na tu re Co n serva n c y ’s
Co n serva tion Stra tegies Grou p, Arl i n g to n , VA 22203. Hel en E. Fox and Al Lo m bana are marine bi ol o gists in the Co n serva tion Sci en ce Pro gra m , Wo rld Wi l dl i fe Fu n d – U S ,
Wa s h i n g to n , DC 20037. Gerald R. All en is a re se a rch asso ci a te at the We s tern Au s tralian Mu seu m , Pert h , We s tern Au s tralia 6986, Au s tra l i a . Ni ck Davi d son is the depu ty
se cret a ry gen eral of the Ramsar Co nven tion Secret a ri a t , CH-1196 Gl a n d , Swi t zerl a n d . Max Fi n l ayson is a mem ber and fo rm er chair of Ra m s a r ’s Sci en tific and
Te chnical Revi ew Pa n el and pri n ci pal re se a rch er in wetland ecol o gy at the In tern a tional Wa ter Ma n a gem ent In s ti tu te , Col o m b o, Sri La n k a . Ben jamin S. Ha l pern is
proje ct coo rd i n a tor for eco s ys tem - ba sed managem ent of coastal marine sys tems at the Na tional Cen ter for Ecol o gical An a lysis and Syn t h e s i s , Santa Ba rba ra , CA 93101.
Mi g u el A . Jo rge is depu ty dire ctor of WWF In tern a ti o n a l ’s Gl obal Ma rine Pro gra m m e , CH-1196 Gl a n d , Swi t zerl a n d . S a ra A . Lou rie is a re se a rch asso ci a te at the
Red path Mu seu m , Mc G i ll Un ivers i ty, Mo n tre a l , Q u ebec H3A 2K6, C a n a d a . Ki rs ten D. Ma rtin was a marine pro gram of f i cer with IUCN (Wo rld Co n serva tion Un i o n )
wh en this arti cle was prepa red and is curren t ly wo rking as a fre el a n ce co n sultant for the Cen sus of Ma rine Li fe In i ti a tive , 1205 Gen eva , Swi t zerl a n d . Ed mund Mc Ma nu s
is a senior pro gram of f i cer in the UNEP (Un i ted Na tions Envi ro n m ent Pro gramme) Wo rld Co n serva tion Mo n i to ring Cen tre , C a m b ri d ge CB3 0DL, Un i ted
MARK D. SPA L D I N G , HELEN E. FOX , GERALD R. ALLEN, NICK DAV I D S O N , Z ACH A. FERDA Ñ A , MAX FINLAY S O N ,
BENJAMIN S. HALPERN, MIGUEL A. JORGE, AL LOMBANA, SARA A. LOURIE, KIRSTEN D. MARTIN, E D M U N D
MCM A N U S , JENNIFER MOLNAR, CHERI A. RECCHIA, AND JAMES RO B E R T S O N
The co n serva tion and su s t a i n a ble use of m a rine re sou rces is a high l i gh ted goal on a growing nu m ber of n a tional and intern a tional policy agen d a s .Un fo rtu n a tely, ef fo rts to assess pro gre s s , as well as to stra tegi c a lly plan and pri o ri ti ze new marine co n serva tion measu re s , h ave be en hampered by thel a ck of a det a i l ed , co m preh en s ive bi o ge o graphic sys tem to classify the oce a n s . Here we repo rt on a new gl obal sys tem for coastal and shel f a reas: theMa rine Eco regions of the Wo rl d , or MEOW, a nested sys tem of 12 re a l m s , 62 provi n ce s , and 232 eco regi o n s . This sys tem provides co n s i d era bly bet ters pa tial re sol u tion than earl i er gl obal sys tem s , yet it pre serves many common el em ents and can be cro s s - referen ced to many regional bi o ge o graphic cl a s s i f i c a ti o n s . The design a tion of terre s trial eco regions has revol u ti o n i zed pri o ri ty set ting and planning for terre s trial co n serva tion; we anti ci pa te similar ben efits from the use of a coh erent and cred i ble marine sys tem .
Key wo rds: eco regi o n s , m a rine bi o ge o gra p hy, m a pp i n g , m a rine prote cted are a s , repre sen t a tive co n serva ti o n
h i gh er taxa, a l ong with the com mu n i ti e s , evo luti on a ry p a t tern s , and eco l ogical processes that sustain this divers i ty.Bi ogeogra phic cl a s s i f i c a ti ons provi de a crucial fo u n d a ti on forthe assessment of repre s en t a tiveness (Ol s on and Di n ers tei n2 0 0 2 , Lo u rie and Vi n cent 2004).
The growing com m i tm ent by govern m ents and the Un i tedNa ti ons (UN; e . g. , the UN Law of the Se a , the UN Fish Stock sAgreem ent) to implem ent com preh en s ive arra n gem ents for ocean govern a n ce provi des an ad d i ti onal arena in wh i chm a rine bi ogeogra phic cl a s s i f i c a ti ons are needed . Bi ogeo-gra phic regi ons are natu ral fra m eworks for marine zon i n g,wh i ch is a tool incre a s i n gly used by regi onal fisheries man-a gem ent or ga n i z a ti on s .
In this arti cl e , we pre s ent a new bi ogeogra phic cl a s s i f i c a-ti on for the worl d ’s coastal and shel f a re a s , wh i ch draws heav-i ly on the ex i s ting gl obal and regi onal litera tu re . We bel i evethat this cl a s s i f i c a ti on wi ll be of c ri tical import a n ce in su p-porting analyses of p a t terns in marine bi od ivers i ty, in un-derstanding proce s s e s , a n d , perhaps most import a n t , i nd i recting futu re ef forts in marine re s o u rce managem ent andcon s erva ti on .
A p p roaches for defining boundaries
Ob s erva ti ons of gl obal bi ogeogra phic patterns in the mari n eenvi ron m ent inclu de early works by Forbes (1856), E k m a n( 1 9 5 3 , f i rst publ i s h ed in German in 1935), and Hed gpet h( 1 9 5 7 a ) , and more recent publ i c a ti ons by Bri ggs (1974, 1 9 9 5 ) ,Hayden and co lleagues (1984), Ba i l ey (1998), and Lon ghu rs t( 1 9 9 8 ) . These aut h ors used a va ri ety of def i n i ti ons and cri-teria for drawing bi ogeogra phic divi s i on s . For ex a m p l e , Bri ggs( 1 9 7 4 , 1995) foc u s ed on a sys tem of coastal and shel f provi n ce sdef i n ed by their degree of en demism (> 10%). This strong tax-on omic focus and clear def i n i ti on have led to rel a tively wi de-s pre ad adopti on of Bri ggs’s sys tem , i n cluding its use byHayden and co lleagues (1984), with minor amen d m en t s , a sa part of t h eir “cl a s s i f i c a ti on of the coastal and marine envi-ron m en t s .” Adey and Sten eck (2001) provi ded indepen den tveri f i c a ti on of m a ny of Bri ggs’s su b d ivi s i ons in a stu dy thatm odel ed “t h erm ogeogra ph i c”regi ons of evo luti on a ry stabi l i ty.
An o t h er important sys tem a tic approach , a i m ed mainly atpel a gic sys tem s , is the two - ti er sys tem devi s ed by Lon ghu rs t( 1 9 9 8 ) , wh i ch focuses on bi omes and bi ogeoch em i c a lprovi n ce s . These su b d ivi s i ons were based on a det a i l ed arrayof oce a n ogra phic factors , te s ted and mod i f i ed using a largegl obal database of ch l orophyll prof i l e s . The re sults repre s en tone of the most com preh en s ive parti ti on i n gs of the pel a gi cbi o t a , but the sch eme is of l i m i ted uti l i ty in the com p l ex sys-tems of coastal waters , a fact ack n owl ed ged by the aut h or, wh ohas recom m en ded com bining his open ocean sys tem with oth-ers for coastal and shel f w a ters (Wa t s on et al. 2 0 0 3 ; Alan R.Lon ghu rs t , G a l erie l’Ac adem i e , Ca ja rc , Fra n ce , pers onal com-mu n i c a ti on , 2 Novem ber 2004).
The sys tem of l a r ge marine eco s ys tems (LMEs) was de-vel oped over many ye a rs by a nu m ber of regi onal ex pert s , wi t hcon s i dera ble input from fisheries scien tist Ken Sherman (e.g. ,S h erman and Al ex a n der 1989, Hem pel and Sherman 2003,
S h erman et al. 2 0 0 5 ) . Un l i ke the sys tems of Bri ggs andLon ghu rs t , LMEs repre s ent an ex pert - derived sys tem wi t h-o ut a ri goro u s , rep l i c a ble core def i n i ti on . LMEs are “rel a-tively large regi ons on the order of 200,000 km2 or gre a ter,ch a racteri zed by disti n ct : (1) bathym etry, (2) hyd rogra phy, ( 3 )produ ctivi ty, and (4) troph i c a lly depen dent pop u l a ti on s”(w w w. l m e . n oa a . gov / Po rt a l /) . LMEs are largely con ceived asunits for the practical app l i c a ti on of tra n s bo u n d a ry man-a gem ent issues (fish and fisheri e s , po lluti on , h a bitat re s tora-ti on , produ ctivi ty, s oc i oecon om i c s , and govern a n ce ) . Th eLME sys tem focuses on produ ctivi ty and oce a n ogra ph i cproce s s e s , and in its pre s ent form omits su b s t a n tial areas ofislands in the Pacific and the Indian oce a n s .
These and other gl obal sys tems con ti nue to play an im-portant role in devel oping our understanding of m a rine bi o-geogra phy and in practical issues of n a tu ral re s o u rcem a n a gem en t . However, i m provem ents are cl e a rly po s s i ble andde s i ra bl e . An ideal sys tem would be hiera rchical and nested ,and would all ow for mu l tiscale analys e s . E ach level of the h i era rchy would be rel evant for con s erva ti on planning orm a n a gem ent interven ti on s , f rom the gl obal to the loc a l , a l-t h o u gh it is beyond the scope of the pre s ent ef fort to cl a s s i f yi n d ivi dual habitats or small er fe a tu re s , su ch as indivi dual es-tu a ries or seagrass meadows .
We focus here on coastal and shel f w a ters ,com bining ben-thic and shel f pel a gic (neri tic) bi o t a s . These waters repre s en tthe areas in wh i ch most marine bi od ivers i ty is con f i n ed ,wh ere human interest and atten ti on are gre a te s t , and wh eret h ere is of ten a com p l ex syner gy of t h reats far gre a ter than inof fs h ore waters (UNEP 2006). From a bi od ivers i ty pers pec-tive , it is not simply that coastal and shel f w a ters have gre a ters pecies nu m bers and high er produ ctivi ty, but also that theya re bi ogeogra ph i c a lly disti n ct from the ad jacent high seas anddeep benthic envi ron m ents (Ekman 1953, Hed gpeth 1957a,Bri ggs 1974).
Our inten ti on was to devel op a hiera rchical sys tem basedon taxon omic con f i g u ra ti on s , i n f lu en ced by evo luti on a ryh i s tory, p a t terns of d i s pers a l , and isolati on . We drew up ini-tial guidelines on def i n i ti ons and nom en cl a tu re to guide thef i rst data-ga t h ering ph a s e , t h en revi ewed and ref i n ed them i tera tively on the basis of the ava i l a ble data.
We revi ewed over 230 works in journ a l s , NGO (non-govern m ental or ga n i z a ti on) report s , govern m ent publ i c a-ti on s , and other source s . For each of t h e s e , we loo ked at theu n derlying data and at the process of i den ti f i c a ti on and de-f i n i ti on of bi ogeogra phic units; we also con s i dered the ob-j ectives of the cl a s s i f i c a ti on s . To fac i l i t a te com p a ri s on s , we usedd i gital mapped vers i ons of m a ny of the ex i s ting bi ogeo-gra phic units. More than 40 indepen dent ex perts provi ded fur-t h er advi ce (see the ack n owl ed gm ents secti on ) . We ref i n ed ad raft cl a s s i f i c a ti on sch eme thro u gh an assessment and revi ewprocess that invo lved a three - d ay work s h op. In arriving at ourcl a s s i f i c a ti on sch em e , we ad h ered to three principles for ourcl a s s i f i c a ti on : that it should have a strong bi ogeogra phic ba-s i s , of fer practical uti l i ty, and be ch a racteri zed by pars i m ony.
A strong biogeographic basis. All spatial units were def i n edon a broadly com p a ra ble bi ogeogra phic basis. Ex i s ting sys-tems rely on a broad array of s o u rce inform a ti on — ra n ged i s con ti nu i ti e s , dominant habi t a t s , geom orph o l ogical fe a-tu re s , c u rren t s , and tem pera tu re s , for ex a m p l e — to iden ti f ya reas and bo u n d a ri e s . In many cases these diver gent ap-proaches are com p a ti bl e , given the close links bet ween bi o-d ivers i ty and the underlying abi o tic drivers (see thecom p a ri s ons bel ow ) . We preferred to be inform ed by com-po s i te studies that com bi n ed mu l tiple diver gent taxa or mu l-tiple oce a n ogra phic drivers in the deriva ti on of bo u n d a ri e s ,as these were more likely to captu re robust or rec u rring pat-terns in overa ll bi od ivers i ty.
A nu m ber of s ys tems we revi ewed were broadly bi ogeo-gra ph i c , but with some ad ju s tm ents to fit po l i tical bo u n d a ri e s .Wh ere it was po s s i ble to discern the bi ogeogra phic el em en t sf rom the po l i ti c a l , these sys tems were sti ll used to inform theproce s s .
Practical utility. We sought to devel op a nested sys tem , op-era ting gl ob a lly at broadly con s i s tent spatial scales and in-corpora ting the full spectrum of h a bitats found ac ross shelve s .We thus avoi ded very fine-re s o luti on sys tems that sep a ra tedcoastal and shel f w a ters into con s ti tu ent habi t a t s . We ch o s enot to try to define minimum or maximum spatial areas forour bi oregi on s , but in some cases we did seek out sys tems thatsu b d ivi ded very large spatial units (su ch as Bri ggs’s In do -Po lynesian Provi n ce , wh i ch covers more than 20% of t h eworl d ’s shall ow shel f a reas) or that amalga m a ted fine-scaleunits su ch as single large estu a ries or sounds.
P a r s i m o n y. Th ere are a nu m ber of re s pected and wi dely uti-l i zed gl obal and regi onal sys tem s , and lack of a greem ent be-t ween su ch sys tems can be probl em a ti c . In devel oping a news ys tem , we sought to minimize furt h er diver gen ce from ex-i s ting sys tem s , yet sti ll to obtain a tru ly gl obal cl a s s i f i c a ti ons ys tem .We did this by adopting a nested hiera rchy that (a) uti-l i zed sys tems that are alre ady wi dely adopted (e.g. , the Na tu reCon s erva n c y ’s sys tem in mu ch of the Am ericas and the In-terim Ma rine and Coastal Regi on a l i s a ti on for Au s tralia) and(b) fitted cl o s ely within broader-scale sys tems or alon gs i deo t h er regi onal sys tem s .
D e f i n i t i o n s
Af ter the revi ew proce s s , we arrived at a set of c ri tical work-ing def i n i ti on s .
R e a l m s . The sys tem’s largest spatial units are based on the ter-re s trial con cept of re a l m s , de s c ri bed by Udva rdy (1975) as“con ti n ent or su bcon ti n en t - s i zed areas with unifying fe a-tu res of geogra phy and fauna/flora / veget a ti on .” From ourm a rine pers pective , realms are def i n ed as fo ll ows :
Very large regi ons of coa s t a l , ben t h i c , or pel a gic oce a nac ross wh i ch biotas are intern a lly co h erent at high ert a xon omic level s , as a re sult of a shared and uniqu e
evo luti on a ry history. Realms have high levels ofen dem i s m , i n cluding unique taxa at gen eric and familyl evels in some gro u p s . D riving factors behind the devel-opm ent of su ch unique biotas inclu de water tem pera-tu re , h i s torical and broadscale isolati on , and the prox-i m i ty of the ben t h o s .
This arti cl e , with its focus on coastal and shel f a re a s , doe snot con s i der realms in pel a gic or deep benthic envi ron m en t s .This is an area requ i ring furt h er analysis and devel opm en t .
P ro v i n c e s . Ne s ted within the realms are provi n ce s :
L a r ge areas def i n ed by the pre s en ce of d i s ti n ct bi o t a sthat have at least some co h e s i on over evo luti on a ry ti m ef ra m e s . Provi n ces wi ll hold some level of en dem i s m ,pri n c i p a lly at the level of s pec i e s . Al t h o u gh histori c a li s o l a ti on wi ll play a ro l e , m a ny of these disti n ct bi o t a sh ave ari s en as a re sult of d i s ti n ctive abi o tic fe a tu res that circ u m s c ri be their bo u n d a ri e s . These may inclu degeom orph o l ogical fe a tu res (isolated island and shel fs ys tem s , s em i en cl o s ed seas); hyd rogra phic fe a tu res ( c u rren t s , u pwell i n gs , i ce dy n a m i c s ) ; or geoch em i c a li n f lu en ces (broadest-scale el em ents of nutri ent su pp lyand salinity ) .
In eco l ogical term s , provi n ces are co h e s ive units likely, forex a m p l e , to en compass the broader life history of m a ny con-s ti tu ent taxa, i n cluding mobile and dispers ive spec i e s . Inm a ny are a s , the scale at wh i ch provi n ces may be con ceived issimilar to that of the det a i l ed spatial units used in gl obal sys-tems su ch as Bri ggs’s provi n ce s , Lon ghu rs t’s bi ogeoch em i c a lprovi n ce s , and LMEs.
E c o re g i o n s . E coregi ons are the smallest-scale units in theMa rine Ecoregi ons of the World (MEOW) sys tem and are def i n ed as fo ll ows :
Areas of rel a tively hom ogen eous species com po s i ti on ,cl e a rly disti n ct from ad jacent sys tem s . The species com-po s i ti on is likely to be determ i n ed by the predom i n a n ceof a small nu m ber of eco s ys tems and/or a disti n ct su i teof oce a n ogra phic or topogra phic fe a tu re s . The dom i-nant bi ogeogra phic forcing agents defining the eco-regi ons va ry from loc a ti on to loc a ti on but may inclu dei s o l a ti on , u pwell i n g, nutri ent input s , f re s hw a ter influ x ,tem pera tu re regi m e s , i ce regi m e s , ex po su re , s ed i m en t s ,c u rren t s , and bathym etric or coastal com p l ex i ty.
In eco l ogical term s , these are stron gly co h e s ive units, su f-f i c i en t ly large to en compass eco l ogical or life history proce s s e sfor most seden t a ry spec i e s . Al t h o u gh some marine ecoregi on sm ay have important levels of en dem i s m , this is not a key determinant in ecoregi on iden ti f i c a ti on , as it has been in ter-re s trial ecoregi on s .
We su ggest that the most appropri a te outer bo u n d a ry forthese coastal and shel f re a l m s ,provi n ce s , and ecoregi ons is the2 0 0 - m eter (m) isob a t h , wh i ch is a wi dely used proxy for thes h el f ed ge and of ten corre s ponds to a dra m a tic eco ton e( Forbes 1856, Hed gpeth 1957b, Bri ggs 1974). Su ch a sharpbo u n d a ry can on ly be indicative : S h el f breaks are not alw ayscl e a r;the bathym etric loc a ti on of an “equ iva l en t” bi o tic tra n-s i ti on is high ly va ri a bl e ; and there is con s i dera ble overl a pand influ en ce bet ween shel f , s l ope , and ad jacent pel a gic bi o-t a s . At the same ti m e , most of the cl a s s i f i c a ti ons that we re-vi ewed have been heavi ly influ en ced by data from nears h oreand intertidal bi o t a s , and data from deeper water typ i c a lly haddec reasing influ en ce on bo u n d a ry def i n i ti on s . We bel i evethat beyond 200 m, o t h er bi ogeogra phic patterns wi ll in-c re a s i n gly predom i n a te , a l tering or hiding the patterns rep-re s en ted by the sys tem propo s ed here .
A global, nested system
We propose a nested sys tem of 12 re a l m s , 62 provi n ce s , a n d232 ecoregi ons covering all coastal and shel f w a ters of t h eworl d .
As the MEOW sys tem is based on ex i s ting cl a s s i f i c a ti on s ,va ri a ti on and mismatch among sys tems led to ch a ll en ge sand com prom i s e s . The gl obal coastal cl a s s i f i c a ti ons of Bri ggsand Hayden , for ex a m p l e , do not show great con gru en cewith the LMEs. The Bri ggs and rel a ted Hayden sys tems a ppe a red to be more cl o s ely all i ed to our need for a sys tem
with a stron ger bi ogeogra phic basis than the current LME de-l i n e a ti on s . Both the Bri ggs and Hayden sys tems and theLMEs show con s i dera ble va ri a ti on in the size of t h eir spati a lu n i t s ; the Bri ggs approach of using 10% en demism disti n-guishes many isolated com mu n i ties around oceanic islands,but fails to disaggrega te vast areas with gradual faunal ch a n ge s ,even wh ere the increm ental ef fects of su ch ch a n ges are veryl a r ge indeed (e.g. , the In do - Pac i f i c ) . The large spatial units ina ll of these sys tems cl e a rly en compass significant levels of i n-ternal bi ogeogra phic heterogen ei ty, wh i ch we were keen to dis-a ggrega te thro u gh a more det a i l ed sys tem of ecoregi on s .
We found regi onal sys tems for almost all coastal and shel fw a ters , a l t h o u gh many are de s c ri bed on ly in the gray litera-tu re . No t a ble excepti ons were the Russian Arctic and thecon ti n ental coasts of mu ch of So ut h , So ut h e a s t , and EastAs i a . For these are a s , we rel i ed heavi ly on gl obal data sets andu n p u bl i s h ed ex pert op i n i on , using more foc u s ed bi ogeo-gra phic publ i c a ti ons (wh ere ava i l a ble) for refining indivi d-ual bo u n d a ri e s .
F i g u re 1 dep i cts the revi ew proce s s , s h owing four bi ogeo-gra phic sch em e s : Bri ggs’s sys tem of provi n ces (1974, 1 9 9 5 ) ;an ex pert - derived sys tem com bining bi o tic and abi o tic fe a-tu res for So uth Am erica (Su ll ivan Se a l ey and Bu s t a m a n te1 9 9 9 ) ; the current LMEs; and a regi onal cl a s s i f i c a ti on basedon a single taxon omic grouping (dec a pod cru s t ace a n s ; Bo s ch i2 0 0 0 ) . De s p i te their different ori gi n s , these sys tems show a re-
Fi g u re 1. Re co n ci l i a tion of d i f f ering bou n d a ry sys tems for South Am eri c a . The map on the left ill u s tra tes fou rbi ogeographic sys tems: (A) Bri ggs’s provi n ce s , (B) Su ll ivan Sea l ey and Bustamante’s provi n ce s , (C) large m a rine eco s ys tem s , and (D) Bo sch i ’s provi n ce s . Sys tem similari ties are ex em plified in three inset maps:n o rt h ern Peru (inset 1), Cabo Frio (inset 2), and Chiloé Island (inset 3). The map on the ri ght shows the Ma rine Eco regions of the Wo rld provi n ces (labeled) and their eco region su b d ivision bou n d a ri e s .
m a rk a ble con gru en ce at a nu m ber of key bi ogeogra ph i cbo u n d a ri e s .
Thu s , it was po s s i ble to adopt a single sys tem as a pri-m a ry source , and the MEOW provi n ces (figure 1, ri ght) wereb a s ed almost en ti rely on Su ll ivan Se a l ey and Bu s t a m a n te( 1 9 9 9 ) , while remaining well align ed with the other sys tem s .At a finer re s o luti on , the ecoregi ons for So uth Am erica are de-rived almost en ti rely from the same publ i c a ti on (Su ll iva nSe a l ey and Bu s t a m a n te 1999), this being the on ly com pre-h en s ive sys tem for these coa s t s . Even at this scale, h owever,ef forts were made to loc a te indepen dent veri f i c a ti on ofbo u n d a ri e s , and it is re a s su ring to note that these more de-t a i l ed su b d ivi s i ons were of ten su pported by data from otheroce a n ogra phic and eco l ogical litera tu re (see , e . g. , S trub et al.
[ 1 9 9 8 ] , Fern a n dez et al. [ 2 0 0 0 ] , O j eda et al. [ 2 0 0 0 ] , and Ca mus [2001] for data con cerning the Chilean coa s t ) .
Al t h o u gh the bo u n d a ries in other regi ons were not assimple to re s o lve as those along the So uth Am erican coa s t ,we app l i ed the same approach e s . The secti on that fo ll ows gives some inform a ti on on the key sources used in drawi n gbo u n d a ri e s .
Marine Ecoregions of the Wo r l d
Box 1 and figures 2 and 3 give a su m m a ry of the en ti reM E OW sys tem , wh i ch covers all coastal and shel f w a terss h a ll ower than 200 m. The shaded area of e ach map (figure s2 , 3) ex tends 370 kilom eters (200 nautical miles) of fs h ore ( or to the 200-m isob a t h , wh ere this lies furt h er of fs h ore ) ,
Fi g u re 2. Final bi ogeographic fra m ewo rk: Realms and provi n ce s . (a) Biogeographic realms with eco regi o nb ou n d a ries ou t l i n e d . (b) Provi n ces with eco regions ou t l i n e d . Provi n ces are nu m bered and listed in table 1.
1 3 9 . Arafura Sea1 4 0 . A rnhem Coast to Gulf of Carp e n t e r i a1 4 1 . B o n a p a rte Coast
3 3 . N o rtheast Australian Shelf
1 4 2 . To rres Strait Nort h e rn Great B a rrier Reef
1 4 3 . Central and Southern Great
B a rrier Reef
3 4 . N o rthwest Australian Shelf
1 4 4 . Exmouth to Broome1 4 5 . N i n g a l o o
3 5 . Tropical Southwestern Pa c i f i c
1 4 6 . Tonga Islands
1 4 7 . Fiji Islands1 4 8 . Va n u a t u1 4 9 . N ew Caledonia
1 5 0 . Coral Sea
3 6 . L o rd Howe and Norfolk Islands
1 5 1 . Lord Howe and Norfolk Islands
E a s t e rn Indo-Pa c i f i c
3 7 . H awa i i
1 5 2 . H awa i i
3 8 . M a rs h a l l , G i l b e rt , and Ellis Islands
1 5 3 . M a rshall Islands1 5 4 . G i l b e rt/Ellis Island
Box 1. Marine Ecoregions of the Wo rl d .
N u m b e rs for the provinces and ecoregions match those shown on the maps in figures 2b and 3. Realms are indicated in boldface, p r ovinces (1–62) ini t a l i c s , and ecoregions (1–232) in roman type.
but , as alre ady noted , we con s i der the principal focus of t h i scl a s s i f i c a ti on to be the benthos above 200 m and the overlyi n gw a ter co lu m n .
Key sources inclu ded the fo ll owi n g :
• Bi ogeogra phic assessments in the peer- revi ewed l i tera tu re , i n cluding the gl obal studies alre ady m en ti on ed and many regi onal publ i c a ti ons (e.g. ,Bu s t a m a n te and Bra n ch [1996] and Tu rpie et al. [ 2 0 0 0 ]for tem pera te sout h ern Af ri c a , Linse et al. [2006] for theSo ut h ern Oce a n )
• E coregi onal assessments con du cted by NGOs (e.g. ,Su ll ivan Se a l ey and Bu s t a m a n te [1999] for Latin Am eri c a , WWF [2004 and unpubl i s h ed reports] for mu ch of Af ri c a , Green and Mous [2006] for the Coral Tri a n gle provi n ce s )
• G overn m en t - derived or su pported sys tems (e.g. ,Th ack w ay and Cre s s well [1998] for Au s tra l i a ,Powles et al. [2004] for Ca n ad a )
• In p ut from several of the aut h ors of this arti cle anda s s e s s m ents com m i s s i on ed ex p l i c i t ly for the MEOW
process (e.g. , u n p u bl i s h ed reports by Jerry M. Kemp in2005 for the Mi d dle Eastern seas and by S. A . L . in 2006for the Andaman to Java coa s t s ) ; the sys tem for theIn do - Pacific oceanic islands was devel oped by one of u s( G . R . A.) on the basis of m a ny ye a rs of f i eld ex peri en ce ,ex pert revi ew, and net working with other scien ti s t sac ross the regi on
These sch emes were assessed alon gs i de other bi ogeogra ph i cl i tera tu re , and in some cases altera ti ons were made to bet terrepre s ent the arguments of bi ogeogra phy, uti l i ty, and pars i-m ony out l i n ed above . A full listing of the sources referen cedcan be found at w w w. n a tu re . o rg / M E OW or w w w. wo rl dwi l dl i fe .o rg / M E OW.
The propo s ed realms adopt the broad lati tudinal divi-s i ons of po l a r, tem pera te , and trop i c a l , with su b d ivi s i on sb a s ed on ocean basin (broadly fo ll owing the oceanic bi om e sof Lon ghu rst [1998]). In the tem pera te waters of the So ut h-ern Hem i s ph ere , we diver ge from this approach . We con s i derthe differen ces ac ross the oceans too su b s t a n ti a l , and thecon n ecti ons around the con ti n ental margins too gre a t , tosu pport ei t h er ocean basin su b d ivi s i ons or a single circ u m-gl obal realm (equ iva l ent to Lon ghu rs t’s An t a rctic We s terlyWinds Bi om e ) , and hen ce we have adopted con ti n ental
N u m b e rs for the provinces and ecoregions match those shown on the maps in figures 2b and 3. Realms are indicated in boldface, p r ovinces (1–62) ini t a l i c s , and ecoregions (1–232) in roman type.
m a r gin realms for tem pera te Au s tra l a s i a , s o ut h ern Af ri c a ,and So uth Am eri c a . The paucity of ex i s ting litera tu re dis-cussing these broadest-scale bi ogeogra phic units from agl obal pers pective pre s ents a stark con trast to the terre s tri a lbi ogeogra phic litera tu re .
The level of i n ternal heterogen ei ty of biotas within differ-ent realms is qu i te va ri ed . For some re a l m s , the differen ces inbiota at the provincial level are su b s t a n ti a l , i n cluding thew a rm tem pera te faunas on ei t h er side of the Tem pera te So ut hAm erica realm and the tropical faunas on ei t h er side of t h eTropical At l a n tic re a l m . By con tra s t , we have su b d ivi ded thewi dely used In do - Pacific “re a l m” i n to three units. This is theregi on of gre a test divers i ty, and it covers a vast are a .Ac ross thisregi on are clinal ch a n ges in taxa that lack clear bre a k s , but aresu f f i c i en t ly large that faunas at ei t h er end bear little re s em-bl a n ce to each other. Our In do - Pacific su b d ivi s i ons (wh i chit might be appropri a te to con s i der as su brealms) fo ll ow lesscl e a rly def i n ed bi ogeogra phic bo u n d a ries than other re a l m s ,but these divi s i ons produ ce spatial units that are more com p a ra ble to other realms in overa ll bi od ivers i ty, l evels ofen dem i s m , and spatial are a .
At broader scales, we undertook a simple spatial analys i sto ex p l ore the links or po s s i ble cro s s overs bet ween the MEOWs ys tem , L M E s , and Bri ggs’s provi n ce s . The incom p l ete cov-era ge of the LME sys tem is cl e a rly limiting for gl obal con-s erva ti on planning: 78 of our 232 ecoregi ons inclu de asu b s t a n tive area (gre a ter than 10% of t h eir total area) that isnot covered by any LME. O f the rem a i n der, s ome 49% ofLMEs show good con gru en ce (> 90% of s h el f a rea) with ei-t h er single ecoregi ons or ecoregi on com bi n a ti on s . (Th ebo u n d a ry of the Arctic LME has not been mapped , and so wasi gn ored in these calculati ons.) In com p a ri s on , 30 of Bri ggs’s53 provi n ces (57%) show good con gru en ce (> 90% of s h el fa rea) with single ecoregi ons or ecoregi on com bi n a ti on s . Th i sf i g u re rises to 39 (74%) if we inclu de con gru en ce at 85% ofthe shel f a re a .
We also used the MEOW sys tem to look at the covera ge ofthe marine and coastal net work of Ramsar site s . Con tracti n g
p a rties to the Ramsar Conven ti on have com m i t ted to ach i evea “co h erent and com preh en s ive nati onal and intern a ti onal net-work” ( Ramsar Conven ti on 1999), a l t h o u gh until now it hasnot been po s s i ble to assess the bi ogeogra phic covera ge ofm a rine and coastal Ramsar sites at the gl obal level . The re su l t sof this overl ay are pre s en ted in table 1.
One va lue of bi ogeogra phic cl a s s i f i c a ti ons is their use in un-covering inequ i ties and dra m a tic gaps in con s erva ti on cov-era ge .Al t h o u gh a more thoro u gh analysis would be requ i redto determine more cl e a rly the degree of repre s en t a ti on pro-vi ded by the ex i s ting sel ecti on of Ramsar site s , s ome basic ob-s erva ti ons are immed i a tely app a ren t . The Ramsar net work isex ten s ive , but it is dom i n a ted by sites in the tem pera te Nort hAt l a n tic and shows a striking paucity of s i tes in, for ex a m p l e ,the eastern In do - Pacific and the So ut h ern Oce a n . At finer hi-era rchical re s o luti on , f u rt h er gaps can be iden ti f i ed :While 92%of realms are repre s en ted , this tra n s l a tes to on ly 73% ofprovi n ces and 52% of ecoregi on s , l e aving some 112 ecoregi on swith no Ramsar repre s en t a ti on . These gaps are wi de s pre ad ,i n cluding four ecoregi ons in the tem pera te North At l a n ti c .
C o n c l u s i o n s
The MEOW cl a s s i f i c a ti on provi des a cri tical tool for mari n econ s erva ti on planning. It wi ll en a ble gap analyses and a s s e s s m ents of repre s en t a tiveness in a gl obal fra m ework . Itprovi des a level of detail that wi ll su pport linkage to practi-cal con s erva ti on interven ti ons at the field level . For ex a m p l e ,t wo major intern a ti onal con s erva ti on or ga n i z a ti ons (the Na tu re Con s ervancy and WWF) use ecoregi ons as planningu n i t s . From a gl obal standpoi n t , the MEOW sys tem of fers sim-ilar opportu n i ties for the marine envi ron m en t . It also provi de sa ra ti onal fra m ework in wh i ch to analy ze patterns andprocesses in coastal and shel f bi od ivers i ty.
The gl obal and hiera rchical natu re of the MEOW can su pport analytical approaches that move bet ween scales.Using MEOW, gl obal inform a ti on can also be used to targetacti on on the gro u n d , while fiel d - l evel inform a ti on can bep l aced alon gs i de inform a ti on on ad jacent or rem o te loc a ti on s ,
Ta ble 1. The geographic spread of m a rine and coastal Ramsar sites within the Ma rine Eco regions of the Wo rl dcl a s s i f i c a ti o n .
E c o r e g i o n s P r ov i n c e s
Total Number with Percentage Number with Percentage
Ramsar Ramsar Total with Ramsar Ramsar Total with Ramsar
R e a l m s i t e s s i t e s n u m b e r s i t e s s i t e s n u m b e r s i t e s
A r c t i c 2 6 1 0 1 9 5 3 1 1 1 0 0
Temperate Nort h e rn Atlantic 3 7 4 2 1 2 5 8 4 6 6 1 0 0
Temperate Nort h e rn Pa c i f i c 3 8 1 2 1 7 7 1 4 4 1 0 0
Tropical Atlantic 1 1 7 1 7 2 5 6 8 4 6 6 7
We s t e rn Indo-Pa c i f i c 4 1 1 4 2 5 5 6 7 7 1 0 0
Central Indo-Pa c i f i c 3 5 1 6 4 0 4 0 1 0 1 2 8 3
E a s t e rn Indo-Pa c i f i c 1 1 1 2 8 1 6 1 7
Tropical Eastern Pa c i f i c 2 9 8 1 1 7 3 2 2 1 0 0
Temperate South America 1 4 9 1 5 6 0 3 5 6 0
Temperate Southern Africa 9 3 5 6 0 2 3 6 7
Temperate Australasia 2 5 9 1 7 5 3 5 6 8 3
S o u t h e rn Ocean 0 0 2 1 0 0 4 0
To t a l 7 0 9 1 2 0 2 3 2 5 2 4 5 6 2 7 3
providing a wi der spatial pers pective . Roo ted in ex i s ting re-gi onal sys tem s , the base units of the MEOW alre ady under-pin con s erva ti on ef forts at regi onal level s , and a strong bodyof m a rine ecoregi onal planning litera tu re illu s tra tes howgl obal or regi onal con cerns can be converted into fiel d - b a s edcon s erva ti on acti on (Banks et al. 2 0 0 0 , Beck and Odaya 2001,L a rs en et al. 2 0 0 1 , Kra m er and Kra m er 2002, Ferdaña 2005).
The va lue of the MEOW sys tem ex tends beyond con s er-va ti on planning. Looking afresh at the broader-scale cl a s s e sand taking adva n t a ge of the improved re s o luti on of fered bythe MEOW sys tem , it is po s s i ble to revi ew wi der issues of bi o-d ivers i ty distri buti on and evo luti on .At the broadest scales, t h emost important el em ents of bi ogeogra phic su b d ivi s i on are theb a rri ers that have sep a ra ted su b s t a n tial areas over evo lu-ti on a ry timescales (Adey and Sten eck 2001). In the MEOWrealms (noting the special case of the In do - Pacific de s c ri beda bove ) , these barri ers consist of l a n d m a s s e s , wi de oce a nb a s i n s , and tem pera tu re grad i en t s .
Al t h o u gh there is va ri a ti on in degree , the provi n ces can bes een as finer-scale units of evo luti on a ry isolati on . Th ey alignwith many of the more important factors driving recent andcon tem pora ry evo luti on a ry proce s s e s . Tem pera tu re , or lati-tu de , con ti nues to play an important role (sep a ra ting warmand cold tem pera te provi n ce s ) , but so does the furt h er iso-l a ti on provi ded by deep water, n a rrow stra i t s , or rapid ch a n ge sin shel f con d i ti on s . E l s ewh ere , the con n ectivi ty provi ded byocean curren t s , su ch as the An t a rctic Coastal Cu rrent and theCa n a ries Cu rren t , can be seen in the cl a s s i f i c a ti on s , and thei m port a n ce of bi o l ogical stepp i n g - s tones thro u gh va ri o u sisland chains is cl e a rly illu s tra ted . F i n a lly, the ecoregi on s ,wh i ch distinguish the MEOW sys tem , ref l ect unique eco l og-ical patterns that ex tend beyond the broad drivers of evo-luti on a ry proce s s e s .
O f co u rs e , as Wa ll ace (1876) noted ,“nothing like a perfectzoo l ogical divi s i on of the earth is po s s i bl e . The causes that havel ed to the pre s ent distri buti on of animal life are so va ri ed , t h ei racti on and re acti on have been so com p l ex , that anomalies andi rreg u l a ri ties are su re to exist wh i ch wi ll mar the sym m etryof a ny ri gid sys tem” ( p. 5 3 ) . Con s equ en t ly, the use of bi o-geogra phic data in a gl obal cl a s s i f i c a ti on is inevi t a bly a proce s sof accom m od a ti on and pra gm a ti s m .The lines we have drawnshould be rega rded as indicative , m a rking approx i m a te lo-c a ti ons of rel a tively rapid ch a n ge in dominant habitats or com-mu n i ty com po s i ti on . O cean bo u n d a ries shift con ti nu o u s lywith we a t h er pattern s , with season s , and with lon ger or morera n dom flu ctu a ti ons in oce a n ogra phic con d i ti on s . In the fu-tu re , the impacts of cl i m a te ch a n ge wi ll add to the instabi l-i ty of m a ny bo u n d a ries in the ocean (Sa ga rin et al. 1 9 9 9 ,Be a u grand et al. 2 0 0 2 , Hi s cock et al. 2 0 0 4 ) .
The need for a com preh en s ive , det a i l ed , and gl ob a lly con-s i s tent marine bi ogeogra phy has been recogn i zed for manyye a rs in marine con s erva ti on . The requ i rem ents for repre-s en t a tive approaches to marine pro tected area de s i gn a ti on inva rious nati on a l , regi on a l , and gl obal planning com m i tm en t sand legal fra m eworks have given ad ded urgency to this need .The MEOW sys tem provi des a basis for planning for coa s t a l
and shel f a re a s , and the links bet ween this sys tem and othergl obal and regi onal sys tems make it po s s i ble to adopt and useit with minimal disru pti on to ex i s ting data sets or analyti c a la pproach e s . The unique co ll a bora ti on of con s erva ti on or ga-n i z a ti ons in devel oping this sys tem adds furt h er va lu e , and mayredu ce the du p l i c a ti on of ef fort that so of ten underm i n e sgl obal con s erva ti on approaches (Mace et al. 2 0 0 0 ) . In short ,the sys tem propo s ed here is powerful and robu s t , and shouldprove to be of great va lue in con s erva ti on planning andbroader bi ogeogra phic discussion . Two intern a ti onal con-s erva ti on agencies (the Na tu re Con s ervancy and WWF) havea l re ady begun to use this sys tem and ex pect to use it morewi dely in the futu re . Si m i l a rly, m em bers of the Scien tific andTechnical Revi ew Pa n el of the Ramsar Conven ti on who par-ti c i p a ted in devel oping this sys tem are undertaking moredet a i l ed analyses to ex p l ore its uti l i ty to su pport the futu rei den ti f i c a ti on and de s i gn a ti on of coastal and marine Wet l a n d sof In tern a ti onal Im port a n ce .
A c k n o w l e d g m e n t s
The Ma rine Eco s ys tems of the World sys tem draws heavi ly onthe work of o t h ers , i n cluding the hu n d reds of con tri butorsto the publ i c a ti on s , gray litera tu re , and work s h ops that c re a ted the many regi onal cl a s s i f i c a ti on s . In ad d i ti on , wewould espec i a lly like to thank the fo ll owing peop l e , who haveprovi ded advi ce or com m en t a ry: Asa An ders s on , Jef f Ard ron ,All i s on Arn o l d , Paul Ba rber, Mi ke Beck , Ca rlo Ni ke Bi a n ch i ,John Bo l ton , G eor ge Bra n ch , John Bri ggs , G eor gina Bu s t a-m en te , Rod ri go Bu s t a m en te , Jose Fa ri n a , Ser gio Floeter,Angus Gascoi gn e , Ser ge Gof a s , Ch a rlie Gri f f i t h s , Huw Gri f f i t h s , Ra n dy Ha gen s tei n , Jon Hoe k s tra , D avid Jo h n ,Peter Ka reiva , Ken Ka s s em , Jerry Kem p, Phil Kra m er, Ka tri nL i n s e , G i lly Llewelly n , S tephan Lut ter, Kasim Moo s a , Al ex i sMor ga n , Dag Na god a , Ser gio Nava rete , Ka te Newm a n , ( Bi n a )Maya Pa u l , Sian Pu ll en , Ca llum Robert s , Rod Sa l m , An d rewSm i t h , Jen n i fer Sm i t h ,Va s s i ly Spiri don ov,Vi ctor Spri n ger, Ju a nLuis Su á rez de Vivero, Ma rco Tavi a n i , Ch a rlie Veron , E l en iVo u l t s i ado u , Mo h i deen Wa f a r, Ca rden Wa ll ace , Ka t hy Wa ll s ,and David Woodl a n d .
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