The Global 200: Priority Ecoregions for Global Conservation

Author(s): David M. Olson and Eric Dinerstein

Source: Annals of the Missouri Botanical Garden, Vol. 89, No. 2 (Spring, 2002), pp. 199-224
Published by: Missouri Botanical Garden Press
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THE GLOBAL200: PRIORITY DavidM. Olson23 and Eric Dinerstein2
ECOREGIONSFOR GLOBAL
CONSERVATION1

ABSTRACT

A globalstrategyto conservebiodiversity mustaimto protectrepresentative examplesof all of theworld'secosystems,

as well as those areasthatcontainexceptionalconcentrations of species and endemics.Althoughlackingthe richness
of tropicalforests,deserts,tropicallakes, and subpolarseas all containdistinctspecies, communities,and ecological
phenomena. Weanalyzedglobalpatternsof biodiversityto identifya set of the Earth'sterrestrial,
freshwater,
andmarine
ecoregionsthatharborexceptionalbiodiversityandare representative of its ecosystems.Weplacedeach of the Earth's
ecoregionswithina systemof 30 biomesand biogeographic realmsto facilitatea representation analysis.Biodiversity
featureswerecomparedamongecoregionsto assess their irreplaceability or distinctiveness.Thesefeaturesincluded
speciesrichness,endemicspecies,unusualhighertaxa,unusualecologicalor evolutionary phenomena,andthe global
rarityof habitats.This processyielded238 ecoregions the Global200 comprisedof 142 terrestrial,53 freshwater,
and 43 marinepriorityecoregions.Effectiveconservationin this set of ecoregionswouldhelp conservethe most
outstanding and representativehabitatsfor biodiversityon this planet.
Keywords: biodiversity, conservation,
ecoregions,endemism,global,phenomena,priority-setting, representation.

Tropicalrainforestsrightfullyreceivemuchcon- or those with unusual ecological or evolutionary

servationattentionas they maycontainhalf of the phenomena.Wemustalso targetrepresentativeex-
world'sspecies. A comprehensivestrategyfor con- amplesof all of the world'sbiomeswithineach bio-
servingglobalbiodiversity,however,muststriveto geographicrealm where they occur (Fig. 1). Be-
save the other50 percentof species and the dis- cause of distinct biogeographichistories, similar
tinctiveecosystemsthatsupportthem.Forexample, kinds of ecosystemsfoundon differentcontinents
while they may not supportthe rich communities or in differentoceanbasinssupportuniqueassem-
seen in tropicalrainforestsor coralreefs,tropical blages of species and highertaxa.For this reason,
dry forests,tundra,polarseas, and mangrovesall globalstrategiesshouldstriveto conserveexamples
larzor unlque specles, communltles,acaptatlons, of every biome in each realmwhere it occurs for
. . . . .

and phenomena.Some of these biomes, such as terrestrial,freshwater,and marinebiodiversity(01-

tropical dry forests and Mediterranean-climate son 8 Dinerstein,1998; Udvardy,1975; Dasmann,
shrublands,are morethreatenedthan are tropical 1974). Here we present the Global 200 an at-
ralnforestsand requlrelmmedlateconservatlon ac- temptto identifya set of ecoregionswhoseconser-
. . . 1 . .

tion. To lose examplesof these assemblageswould vationwould achieve this goal of saving a broad
representan enormousloss of globalbiodiversity. diversityof the Earth'secosystems(Figs.2, 3). This
Limitedfundingcompelsthe conservationcom- paperexpandsand updatesan earlieranalysisby
munityto be strategicand earmarkthe greatest Olson and Dinerstein(1998). Several additional
amountof resourcesto protectthe mostoutstanding ecoregionshavebeen identifiedthroughongoingre-
andrepresentative Ona glob- gionalanalyses(e.g., Wikramanayake
areasforbiodiversity. et al., 2001)
al scale, this requiresidentifyinglargeregionswith and the marine Global 200 have been reduced,
exceptionallevels of speciesrichnessorendemism, largely due to combiningseveral adjacentareas

1We thankthe regionalexperts,biologists,and conservationists who contributedtheirtime and knowledgeto the

conservationanalysesthat went into the Global200. J. Leapeand C. Hails have providedcriticalsupportfor this
effort.The staff of WWFcontributedgreatlyto the regionalassessmentsfromwhichthe map is derived.We thank
WorldWildlifeFund'sConservation ScienceProgram fortheircontribution
to the analysisandpreparationof the Global
200, specificallyR. Abell,T.Allnutt,C. Carpenter,
J. D'Amico,P. Hurley,K. Kassem,H. Strand,M. Taye,M.Thieme,
W.Wettengel,E. Underwood, E. Wikramanayake,I. Itoua,C. Loucks,T. Ricketts,S. Walters,P. Hedao,M. McKnight,
Y. Kura,J. Morrison,and G. Powell.J. Martin-Jones and U. Laglerhelpedin manywaysto facilitatethe completion
of this project.Wethankthe staffof the WWFNetwork,includingall of the nationalorganizations,variousfieldoffices
and programs, and associates,fortheirreviewand commentson earlierdrafts.
2 Conservation Science Program,WorldWildlifeFund-US,1250 24th Street,NW,Washington, D.C. 20037, U.S.A.
dolson@wes.org, eric.dinerstein@wwfus.org.
3 WCS SouthPacificProgram,P.O.Box3080, Lami,Fiji.
ANN.MISSOURI
BOT.GARD.89: 199-224. 2002.

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 Annals of the
200
Missouri Botanical Garden

Antarctic
_ Tropical
and Subtropical
MoistBroadleafForests g Temperate Grasslands, Savannas, and Shrublands
_ Tropical
and Subtropical
Dw BroadleafForests Flooded Grasslands and Savannas
w Tropical
and Subtropical
ConiferousForests _ Montane Grasslands and Shrublands
- TemperateBroadleafand MixedForests X Tundra
-

_ _
TemperateConiferousForests _ MediterraneanForests, Woodlands, and Scrub
1|1| BorealForests/Taiga 1 | Deserts and Xeric Shrublands
_ Tropical
and Subtropical
Grasslands,Savannas _ Mangroves
and Shrublands
Figure1. Terrestrialbiomes,terrestrialand freshwaterbiogeographic
realms,and marinebiogeographic
realms
(sensuDasmann,1974; Udvardy,1975).

into largerunits.In addition,the conservationsta- terrestrial,53 freshwater, and43 marineecoregions

tus of all ecoregionshas been estimated. nestedwithin30 biomesand8 terrestrialandfresh-
Ecoregionsare regional-scale(continental-scale) waterbiogeographic realmsand 13 marinebiogeo-
unitsof biodiversity.Wedefineecoregionsas a rel- graphicsubdivisions(Table1).
atively large area of land or water containinga
characteristic set of naturalcommunitiesthatshare
a largemajorityof theirspecies,ecologicaldynam- DELINEATION OF ECOREGIONS
ics, andenvironmental conditions(Dinersteinet al.,
1995;Groveset al., 2000). Theyfunctioneffective- TERRESTRIAL ECoREGIoNS
ly as coarse-scaleconservationunits becausethey
encompass similar biological communities,and
their extent roughlycoincides with the area over Dasmann(1974) and Udvardy(1975) were the
which key ecological processes interact most firstto conducta globalrepresentation analysisfor
strongly(Orians,1993). terrestrialconservation.Dasmann'ssystemof 198
Foreach of the Earth's30 terrestrial,freshwater, bioticprovincesandUdvardy's 193 unitsarenested
and marinebiomes (formerlyreferredto as major within 7 biogeographicrealms and 13 terrestrial
habitattypes in our previousanalysis),we com- biomes and 1 freshwaterbiome. Both these geo-
pared the biodiversityof each constituentecore- graphic models serve as logical frameworksfor
gion.Thoseecoregionswhoselevels of biodiversity analysesof globalrepresentation.
were consideredexceptional(that is, highly dis- Therelativecoarsenessof Dasmann's andUdvar-
tinctive or irreplaceable;see Dinersteinet al., dy'sbioticprovinces,however,limitstheirutilityas
1995; Pressey et al., 1994) for their biome, or regionalconservationplanningtools as manydis-
whichwereconsideredthe best exampleof a biome tinctive biotas may remainunrecognized.We be-
withina realm(evenif noneof the candidateshar- lieved a morefinely resolvedmap of biodiversity
boredexceptionalbiodiversity),were identifiedas patternswas required,one thatmappeddistinctive
areasof particularimportancefor achievingglobal biotaswithinsingle,continuousbiomes.Thiscalled
conservationgoals. This prioritizationyielded238 for intensiveregionalanalysesof biodiversitypat-
ecoregions the Global 200- comprisedof 142 ternsacrossfivecontinentsby synthesizingexisting

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 _ Tropicaland SubtropicalMoistBroadleafForests g TemperateGrasslands,Savannas,a
_ Tropicaland SubtropicalDryBroadleafForests _ FloodedGrasslandsand Savannas
_ Tropicaland SubtropicalConiferousForests _ MontaneGrasslandsand Shrublan
_ TemperateBroadleafand MixedForests m Tundra
_ TemperateConiferousForests _ MediterraneanForests,Woodlands
_ Borealhrests/Taiga _ Desertsand XericShrublands
_ Tropicaland SubtropicalGrasslands,Savannas, _ Mangroves
and Shrublands
Figure2 The terrestrialGlobal200 ecoregionstargetsecoregionswith outstandingbiodiversityfeaturesand representative
value.Th

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202 Annals of the
Missouri Botanical Garden

Table 1. Global 200 ecoregions organized by biomes Table1. Continued.

and biogeographic realms. the estimated conservation sta-
tus for each ecoregion is noted as follows: CE for critical 46. Central Andean Yungas
or endangered, V for vulnerable, and RS for relatively 47. SouthwesternAmazonian Moist Forests CE
stable or intact. Ecoregions marked by asterisks (8) rep- 48. Atlantic Forests RS
resent new areas presently under review for elevation to Oceania CE
49. South Pacific Island Forests
Global 200 status based on their biodiversity features and
50. Hawaii Moist Forests CE
representationvalue.
CE
TROPICAL AND SUBTROPICAL DRY BROADLEAF FORESTS
TERRESTRIAL
REALM Afrotropical
51. MadagascarDry Forests E
C1E
TRoPICAIANDSUBTRoPICAL MoISTBRoADLEAF FoRESTS Australasia
Afrotropical 52. Nusa TenggaraDry Forests E
C1E
1. Guinean Moist Forest CE 53. New Caledonia Dry Forests C1E
2. Congolian Coastal Forests CE
3. CameroonHighland Forests CE Indo-Malayan
4. NortheasternCongo Basin Moist Forests V 54. IndochinaDryForests
5. Central Congo Basin Moist Forests RS 55. Chhota-Nagpur
DryForests CE
6. Western Congo Basin Moist Forests V Neotropical CE
7. Albertine Rift Montane Forests CE 56. MexicandryForests
8. East African Coastal Forests CE 57. Tumbesian-AndeanValleysDryForests CE
9. Eastern Arc MontaneForests CE 58. ChiquitanoDryForests CE
10. MadagascarForests and Shrublands CE 59. AtlanticDryForests CE
11. Seychelle and Mascarene Moist Forests CE Oceania CE
Australasian 60. HawaiiDryForests
12. Sulawesi Moist Forests CE CE
13. Moluccas Moist Forests V TROPICAL AND SUBTROPICAL CONIFEROUS FORESTS
14. Southern New Guinea Lowland Forests V
Nearctic
15. New Guinea MontaneForests RS
61. SierraMadreOrientaland OccidentalPine-
16. Solomons-Vanuatu-BismarekMoist Forests V
OakForests
17. Queensland Tropical Forests V
Neotropical CE
18. New Caledonia Moist Forests CE
62. GreaterAntilleanPine Forests
19. Lord Howe-Norfolk Islands Forests CE
63. Mesoamerican Pine-OakForests CE
Indo-Malayan
20. SouthwesternGhats Moist Forests CE CE
TEMPERATE BROADLEAF AND MIXED FORESTS
21. Sri Lankan Moist Forests CE
22. Northern Indochina Subtropical Moist For- Australasia
ests V 64. EasternAustraliaTemperateForests CE
23. Southeast China-Hainan Moist Forests CE 65. TasmanianTemperate Rain Forests V
24. Taiwan Montane Forests V 66. New ZealandTemperateForests V
25. Annamite Range Moist Forests V Indo-Malayan
26. SumatranIslands Lowlandand MontaneFor- 67. Eastern HimalayanBroadleafand Conifer
ests CE Forests V
27. Philippine Moist Forests CE 68. WesternHimalayanTemperate Forests CE
28. Palawan Moist Forests CE Nearctic
29. Kayah-Karen/TenasserimMoist Forests V 69. Appalachianand MixedMesophyticForests V
30. Peninsular Malaysian Lowland and Montane Palearctic
Forests V 70. SouthwestChinaTemperateForests V
31. Borneo Lowland and Montane Forests CE 71. RussianFarEastTemperate Forests V
32. Nansei Shoto Archipelago Forests CE
33. Eastern Deccan Plateau Moist Forests CE TEMPERATE CONIFEROUS FORESTS
34. Naga-Manupuri-ChinHills Moist Forests V Nearctic
35. CardamomMountains Moist Forests RS 72. PacificTemperate
Rainforests
36. Western Java Montane Forests CE 73. Klamath-Siskiyou
ConiferousForests CE
Neotropical 74. SierraNevadaConiferousForests CE
37. GreaterAntillean Moist Forests CE 75. SoutheasternConiferousand BroadleafFor- CE
8 (Lesser Antillean Moist Forests) CE ests
38. Talamancan-IsthmianPacific Forests RS Neotropical CE
39. Choco-Darien Moist Forests RS 76. ValdivianTemperateRainforests/Juan
Fer-
40. NorthernAndean Montane Forests CE nandezIslands
41. Coastal Venezuela Montane Forests V 8 (JuanFernandez
IslandsandDesventuradasCE
42. Guianan Moist Forests RS Islands)
43. Napo Moist Forests V Palearctic CE
44. Rio Negro-JuruaMoist Forests CE 77. European-Mediterranean
MontaneMixed
45. Guayanan Highland Moist Forests RS Forests
CE

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Volume 89, Number2 Olson & Dinerstein 203
2002 The Global 200

Table1. Continued. Table1. Continued.

78. Caucasus-Anatolian-Hyrcanian
Temperate 111. Middle Asian MontaneSteppe and Wood-
Forests CE lands
79. Altai-SayanMontaneForests V 112. EasternHimalayanAlpineMeadows v
80. HengduanShanConiferousForests RS RS
TUNDRA
BOREAL FORESTS/TAIGA Nearctic
Nearctic 113. AlaskanNorthSlopeCoastalTundra
81. Muskwa/Slave
LakeBorealForests RS 114. CanadianLowArcticTundra RS
82. CanadianBorealForests RS Palearctic RS
Palearctic
115. Fenno-ScandiaAlpineTundraand Taiga
83. UralMountainsTaiga V 116. Taimyrand SiberianCoastalTundra V
84. EasternSiberianTaiga RS 117. ChukoteCoastalTundra RS
85. Kamchatka Taigaand Grasslands RS RS
TROPICAL AND SUBTROPICAL GRASSLANDS, SAVANNAS, AND MEDITERRANEAN FORESTS, WOODLANDS, AND SCRUB
SHRUBLANDS Afrotropical
Afrotropical 118. Fynbos
86. Hornof AfricaAcaciaSavannas V Australasia CE
87. East AfricanAcaciaSavannas V 119. Southwestern
AustraliaForestsand Scrub
88. Centraland EasternMiomboWoodlands V 120. SouthernAustraliaMalleeand Woodlands CE
89. SudanianSavannas CE Nearctic CE
Australasia 121. CaliforniaChaparral
and Woodlands
90. NorthernAustraliaand Trans-FlySavannas RS Neotropical CE
Indo-Malayan 122. ChileanMatorral
91. Terai-Duar
Savannasand Grasslands CE CE
Neotropical Palearctic
92. LlanosSavannas V 123. MediterraneanForests, Woodlands,and
93. CerradoWoodlands
and Savannas V Scrub
CE
TEMPERATF GRASSI ANI)S, SAVANNAS, AND .SEIRUBIANDS 1)ESERTS ANI) XERIC SHRUBI ANI)S
Nearctic Afrotropical
94. NorthernPrairie CE 124. Namib-Karoo-KaokoveldDeserts
* Tallgrassprairies 125. Madagasear SpinyThifket V

Neotropical 126. SocotraIslandDesert CE

95. PatagonianSteppe CE 127. Arabian HighlandWoodlandsand Shrub- CE
Palearctic lands
96. DaurianSteppe V Australasia V

128. CarnavonXericScrub
FLOODED GRASSLANDS AND SAVANNAS 129. GreatSandy-Tanami Deserts CE
Afrotropical Nearctic RS
97. Sudd-SahelianFloodedGrasslandsand Sa- 130. Sonoran-BajaDeserts
vannas CE 131. Chihuahuan-Tehuacan Deserts RS
98. ZambezianFloodedSavannas v Neotropical V
Indo-Malayan
99. Rannof KutchFloodedGrasslands CE 132. GalapagosIslandsDeserts
133. Atacama-Sechura Scrub V
Neotropical V
100. EvergladesFloodedGrassland Palearctic
101. PantanalFloodedSavannas v 134. CentralAsian Deserts
CE CE
MONTANE GRASSLANDS AND SHRUBLANDS MANGROVES
Afrotropical Afrotropical
Atlantic
102. EthiopianHighlands CE 135. Gulfof GuineaMangroves
103. SouthernRiftMontaneWoodlands CE Afrotropical
Indian CE
104. East AfricanMoorlands RS 136. East AfricanMangroves
105. Drakensberg MontaneShrublands
andWood- 137. Madagascar Mangroves CE
lands CE Australasia CE
Australasia 138. New GuineaMangroves
106. CentralRangeSubalpineGrasslands RS Indo-Malayan Indo-Pacific RS
Indo-Malayan 139. Sundarbans Mangroves
107. KinabaluMontaneShrublands RS 140. GreaterSundasMangroves CE
Neotropical Neotropical
Atlantic CE
108. NorthernAndeanParamo RS 141. Guianan-Amazon Mangroves
109. CentralAndeanDryPuna V Neotropical
Pacific RS
Paleartic 142. PanamaBightMangroves
110. TibetanPlateauSteppe V RS

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 Annals of the
204 Missouri Botanical Garden

Table1. Continued. Table1. Continued.

FRESHWATER
REALM 179. GreaterAntilleanFreshwater CE
8 (SouthernConeFreshwater,especiallyVal-
LARGE
RIVERS
Afrotropical
divianregion) V
143. CongoRiverand FloodedForests RS 8 (AtlanticCoastriversof SE Brazil,Uruguay)V
Palearctic
Indo-Malayan
180. BalkanRiversand Streams CE
144. MekongRiver
Nearctic
V 8 (expansionto Mediterranean regionin gen-
145. ColoradoRiver CE eral including
181. Russian western
FarEast North
Rivers Africa)
and Wetlands RS
146. LowerMississippiRiver CE 8 (AralSea Basin,particularlySyr-andAmu-
Neotropical
147. AmazonRiverand FloodedForests RC Dar'yaRivers) CE
148. OrinocoRiverand FloodedForests RS LARGE LAKES
Palearctic
Afrotropical
149. YangtzeRiverand Lakes CE 182. Rift Valley Lakes
Neotropical CE
LAR6 RIVF,R HEADWATERS 183. High Andean Lakes
Afrotropical
Palearctic CE
150. CongoBasinPiedmontRiversand Streams 184. Lake Baikal
RS
Nearctic
185. Lake Biwa V
151. MississippiPiedmontRiversand Streams
CE CE
Neotropical
152. UpperAmazonRiversand Streams RS SMALL LAKES
153. UpperParanaRiversand Streams CE Jrotropzeat
186. Cameroon
CraterLakes
154. Brazilian Shield AmazonianRivers and CE
Streams V Australasia
187. LakesKutubuand Sentani
188. CentralSulawesiLakes RS
LARGE
RIVe§t
DELTAS Indo-Malayan V

Afrotropical 189. PhilippinesFreshwater

155. NigerRiverDelta CE 190. LakeInle CE
Indo-Malay(ln V
191. YunnanLakesand Streams
156. IndusRiverDelta CE Neotropical CE
Palearctic 192. MexicanHighlandLakes
157. VolgaRiverDelta rvn CE
158. Mesopotamian
Deltaand Marshes CE XERIC BASINS
159. DanubeRiverDelta CE Australasia
160. LenaRiverDelta RS 193. CentralAustralianFreshwater
V
Nearctic
SMALL
RIVeRS 194. Chihuahuan
Freshwater
CE
Afrotropical Palearctic
161. UpperGuineaRiversand Streams CE
195. AnatolianFreshwater
CE
162. MadagascarFreshwater CE
163. Gulfof GuineaRiversand Streams MARINE
REALM
V
164. CapeRiversand Streams CE
Australasia POLAR
165. New GuineaRiversand Streams RS
Antarctic
166. New CaledoniaRiversand Streams CE
196. AntarcticPeninsulaand WeddellSea
RS
167. KimberleyRiversand Streams Arctic
RS
168. SouthwestAustraliaRiversand Streams CE
197. BeringSea
V
169. EasternAustraliaRiversand Streams CE
198. Barents-Kara
Seas
CE
* (NewZealandRiversand Streams) V
Indo-Malayan TEMPERATE SHELF AND SEAS
170. Xi JiangRiversand Streams Mediterranean
CE
171. WesternGhatsRiversand Streams CE
199. Mediterranean
Sea
CE
172. SouthwesternSri LankaRiversand Streams V North TemperateAtlantic
173. SalweenRiver 200. NortheastAtlanticShelfMarine
V CE
174. SundalandRiversand Swamps 201. GrandBanks
V CE
Nearctic 202. ChesapeakeBay
V
175. Southeastern
Riversand Streams CE
North TemperateIndo-Pacific
176. PacificCoastalRiversand Streams CE
203. YellowSea
CE
177. Gulfof AlaskaCoastalRiversand Streams RS
204. OkhotskSea
RS
Neotropical
178. GuiananFreshwater RS

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Volume 89, Number 2 Olson & Dinerstein 205
2002 The Global 200

Table1. Continued. classificationsfromfiner scales.4Furthermore, de-

lineations were conducted in collaborationwith
SouthernOcean hundredsof regionalexpertsand included exten-
205. PatagonianSouthwestAtlantic v
206. SouthernAustralianMarine RS
sive literaturereviews.The resultis a digitalmap
207. NewZealandMarine V of 867 terrestrialecoregions,classifiedwithinbi-
omes and realms, to be used for priority-setting
TEMPERATE UPWELLING analyses(Olsonet al., 2001). This map providesa
North TemperateIndo-Pacific muchmoredetailedpictureof how species assem-
208. CaliforniaCurrent RS blages are distributedacross the world. The in-
South TemperateAtlantic
209. BenguelaCurrent V
creasedresolutionis most apparentin the tropics
South TemperateIndo-Pacific whereDasmann(1974) and Udvardy(1975) iden-
210. HumboldtCurrent V tified 115 and 117 provincesrespectively,com-
211. AgulhasCurrent RS paredto 463 terrestrialecoregions.
Dasmannand Udvardyboth went on to assess
TROPICAL UPWELLING
CentralIndo-PaciJ%c
how well existingprotectedareas representedthe
212. WesternAustraliaMarine RS
Earth'sterrestrialbiomes and realms.Biotic prov-
Eastern Indo-PaciJ%c inces with little or no protectionwereidentifiedas
213. PanamaBight V priorities.The Global200 analysisframesthe goal
214. Gulfof California CE of prioritization differently:We ask which regions
215. GalapagosMarine V
Eastern TropicalAtlantic should be a priorityfor conservationaction (e.g.,
216. CanaryCurrent CE designatingand strengthening protectedareas)be-
cause of their outstandingbiodiversityfeaturesor
TROPICAL CORAL theirrepresentation value Wealso applythis ques-
(DentralIndo-PaciJ%c tion to the terrestrial,freshwater,and marine
217. NanseiShoto CE
218. Sulu-SulawesiSeas realms.
CE
219. Bismarek-Solomon Seas RS
220. Banda-FloresSea V FRESHWATF.R ECORF.GIONS
221. New CaledoniaBarrierReef RS
222. GreatBarrierReef RS Separateanalyses of freshwaterand terrestrial
223. LordHowe-Norfolk IslandsMarine RS ecoregionswereconductedbecausethe distribution
224. PalauMarine V of freshwaterbiodiversitydivergesfromterrestrial
225. AndamanSea V
Eastern Indo-PaciJ%c patterns.Freshwaterecoregionswerebasedon sev-
226. Societies/ Marquesas/Tuamotus Marine V
eral regionalanalyses and consultationswith re-
227. HawaiianMarine V gionalexperts. Currently,the Global200 analysis
228. RapaNui Marine RS effectivelytargetsthe majorityof freshwaterprior-
229. Fiji BarrierReef & Marine RS ities. Sometargets,however,maychangeas we near
WesternIndo-PaciJ%c
230. Maldives,Chagos,Lakshadweep Atolls V
completionof a globalmapof freshwater ecoregions
231. Red Sea V that is based on a standardlevel of biogeographic
232. ArabianSea CE resolutionand relevantbiomes.
233. East AfricanMarine N
234. WestMadagascar Marine V
* (The MascareneIslandsare underconsid- 4Victor(1955)! Freitag(1971), Zohary(1973), Miya-
erationdue to highnumbersof endemicreef waki (1975), Yim (1977), ChineseVegetationMapCom-
fish) pilationCommittee(1979), Wiken(1986), New Zealand
* (The Maldivesare underconsiderationfor Departmentof Conservation(1987), Noirfalise(1987),
extensionto includeSriLankaandsouthern ChangehunInstituteof Geography and ChineseAcademy
Indiancoast) of Sciences(1990),Kurnaev(1990), Bohn(1994),Krever
WesternTropicalAtlantic et al. (1994), Dinersteinet al. (1995),EcologicalStratifi-
235. Mesoamerican Reef CE cationWorking Group(1995),Gallantet al. (1995),Hilbig
236. GreaterAntilleanMarine CE (1995), Omernik(1995), Thackway& Cresswell(1995),
237. SouthernCaribbeanSea V MongolianMinistryforNatureandthe Environment et al.
238. NortheastBrazilShelfMarine V (1996), EuropeanTopic Centreon NatureConservation
(2000), Rickettset al. (1999), WWF/IUCN(1994, 1995,
1997), Bohn & Katenina(1996), Wikramanayake et al.
(2001),S. Gon(pers.comm.).
5 Hocutt & Wiley (1986), Frest & Johannes(1991),

WCMC(1992), Maxwellet al. (1995), Oberdorffet al.

(1995), Kottelat& Whitten(1996), Olson et al. (1999),
Abell et al. (2000),Thiemeet al. (in press).

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 00
::0
:t ff f; f:f:E 0f:
X X X 0S S0 it - :C:L 00 ; : :L ::000
: :: --
t0 : {f :: :::: 0s 0

t- - 3
Freshwater Marine
_ Largelake _ Temperateshelf and sea
_ Largeriver - Coastaltemperateupwelling
- Largeriverdelta _ Coastaltropicalupwelling
- Largeriverheadwater _ Coastaltropicalcoral
- Smalllake - Polar
- Smallriverbasin
S Xericbasin
Figure3. The freshwater
and marineGlobal200 ecoregions.The numberscorrespond
to the ecoregionslisted in Table1.

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Volume 89, Number 2 Olson & Dinerstein 207
2002 The Global 200

MARINE ECOREGIONS versityconductedoverthe last severalyears(Krev-

Relativeto mostterrestrialandfreshwater ecore- er et al., 1994; Dinersteinet al., 1995; Olsonet al.,
gions, marineecologicaland biogeographicunits 1999;Rickettset al., 1999;Abell et al.,2000; Wik-
ramanayakeet al., 2001; Burgesset al., in press;
are morespatiallyand temporallydynamic(Sher-
man et al., 1990) and thereforemorechallenging Thieme et al., in press). Withineach biome and
to delineate.Marineecoregionsdelineatedby the biogeographicrealm, the relative importanceof
Global200 are derivedfroma synthesisof global ecoregionswas classifiedat one of fourlevels:glob-
and regionalspatialschemas,reviewof the avail- ally outstanding,regionallyoutstanding(e.g., Neo-
able literature6andconsultationswithexperts.Kel- tropics,AtlanticOcean),bioregionallyoutstanding
leher et al. (1995), Shermanet al. (1990), Lon- (e.g., Caribbean),or locally important.The criteria
ghurst (1998), and Bailey (1998) served as the used to prioritizeecoregionsforthe Global200 are
primarysourcesfor the Global200. Ourbase map the same as those used for the regionalassess-
ments.Wechose the set of ecoregionswithineach
does notcoverdeep waterecosystems(i.e., pelagic,
abyssal,or hadal)nor are its biogeographicunits biome that were consideredto harborbiodiversity
that was globally outstandingor regionallyout-
as finely resolvedas the maps used in the fresh-
standingbasedon the parametersdescribedbelow.
wateror terrestrialanalyses.We believe that sev-
These parameterswere weightedand measured
eral forthcomingand detailed analysesof marine
in the regionalanalysesas illustratedin Appendix
biodiversityaround the world (CallumRoberts,
1. The weightassignedto the differentparameters
pers. comm. 2001, Gerald Allen, pers. comm.
variedby biometo betteraddressspecificpatterns
2002) will be useful in testingand improvingthe
of biodiversityand ecologicaldynamics.
accuracyof our results.As in the land-basedanal-
yses, the delineationof marineecoregionsis in- SPECIES RICHNESS AND ENDEMISM
tended to highlightgeneralregionswithin which
characteristicanimals,plants, ecologicalinterac- Richnessvalueswerefirstcorrectedforarea.We
tions, and biophysicalprocessesoccur. then dividedthe rangeof valuesforthe set of ecore-
gions sharingthe same biomeand realminto four
sIo(izoGRAPHIc RESoI UTIoN categoriesbased on naturalbreaks.Globallyout-
standing ecoregions were comparedwith those
Themajorityof Global200 regionsarecomposed identifiedfor otherrealmsto ensureconsistency.In
of an aggregationof continental-scaleecoregions. general,widelyrecognizedglobalandregionalcen-
This reflectsthe coarserlevel of biogeographicres- ters of richness and endemismwere selected for
olution applied on a global scale. For example, Global200 status.The precisionof the datavaried
whereas12 terrestrialunitsweredelineatedforthe considerablyas illustratedby richness and ende-
islandof NewGuineain the regionalanalysis(Wik- mism values for vascularplants in temperateco-
ramanayake et al., 2001), only 5 Global200 units niferand tundrabiomes(Tables2, 3).
are recognized.The ecoregionsthatwerecombined
are adjacent,relatedby habitattype, and are bio- HIGHER TAXONOMIC UNIQUENESS
geographicallysimilarat a global scale. Appoxi- The presenceof an endemichighertaxon(genus
matelya thirdof the ecoregionsusedin the regional or family)wouldcontributemoreto an ecoregion's
analysescorresponddirectlyto Global200 ecore- biotic distinctivenessthanwouldan endemicspe-
gions. The specific location and configurationof cies. Some ecoregionsare notablefor biotas that
boundariesof Global200 ecoregionsdo notpresent containuniquetaxaat highertaxonomiclevels than
exact target areas for detailed planning.Rather, species (Vane-Wright et al., 1991; Williamset al.,
Global200 ecoregionsare primarilyintendedto 1991;Gaston& Williams,1993;Foreyet al., 1994;
spotlightregionsof exceptionalimportance forstra- Williams& Humphries,1994). For example,the
tegic decision-making. moist forests of northeasternAustralia,northern
New Zealand,and New Caledoniaare recognized
SELECTION CRITERIA FORTHEGLOBAL 200 as havinga numberof the mostprimitivelineages
Selectionof the Global200 drawsheavilyfrom of conifersandfloweringplantsin the world(WWF/
the resultsof intensiveregionalanalysesof biodi- IUCN,1994-1997).
UNIQUE ECOLOGICAL OR EVOLUTIONARY
6 Haydenet al. (1984),IUCN(1988),Sherman(1990), PHENOMENA
Croomet al. (1992), Ray & Hayden(1993), Kelleheret
al. (1995),Groombridge & Jenkins(1996),Ormondet al. Some ecoregionswere elevated to Global 200
(1997),Sullivan& Bustamante (1996),Longhurst
(1998). status because of their extraordinaryecological

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208 Annals of the
Missouri Botanical Garden

Table2. Estimatedrichnessand endemism(expressedas numberof species)of nativevascularplantspecies for

temperateconiferousforestecoregionsaroundthe world.Data for ecoregionsof the UnitedStatesand Canadaare
derivedfromthe Biotaof NorthAmericaProgramdatabasesdevelopedby KarteszandMeacham(1999).Theestimates
for Eurasianecoregionsmaybe comparatively higherthanvaluesfor the Americasbecausethe formertypicallyen-
compassbiogeographic areasthatare broaderin scope (i.e., they includemixed-conifer
and broadleafforesthabitats)
thanecoregionsdelineatedfor the Americas(WWF/IUCN, 1994-1997; Mittermeieret al., 1999).

Ecoregion Speciesrichness Endemism

Nearctic
SoutheasternConiferForests 3095 >201
SierraNevadaForests 2373 51-75
ArizonaMountainsForests 2204 76-110
SouthCentralRockiesForests 1933 51-75
Klamath-SiskiyouForests 1859 111-151
PineyWoodsForests 1729 4-10
NorthCentralRockiesForests 1695 21-50
ColoradoRockiesForests 1626 76-110
MiddleAtlanticCoastalForests 1488 11-20
OkanoganForests 1355 1-3
CascadeMountainLeewardForests 1328 11-20
NorthCascadesForests 1325 4-10
Centraland SouthernCascadesForests 1296 21-50
EasternCascadeForests 1224 21-50
NorthernCaliforniaCoastalForests 1212 11-20
Blue MountainForests 1134 21-50
Wasatchand UintaMontaneForests 1109 51-75
CentralPacificCoastalForests 1109 11-20
PugetLowlandsForests 1100 1-3
GreatBasinMontaneForests 1043 21-50
FraserPlateauand BasinComplex 1012 0
FloridaSandPine Scrub 951 21-50
NorthernBritishColumbiaMountainForests 909 0
NorthernTransitionalAlpineForests 876 0
Alberta/British
ColumbiaFoothillForests 740 1-3
AlbertaMountainForests 660 1-3
NorthernPacificCoastalForests 615 1-3
QueenCharlotteIslands 459 1-3
AtlanticCoastalPine Barrens 632 1-3
Neotropics
ValdivianTemperate
Rainforests 463 >33
Palearctic
CaucasusMountains 6300 1600
MiddleAsia Mountainse 5500 1500
Pyrenees 3500 200
Balkan-RhodopeMountains 3000 900
Alps 3000 350
Carpathians 2000 100
CentralChinaMixed-Conifer
Forests 1900 ?
EasternHimalayanTemperate
ConiferForests 1500 ?
8 Kopetdag,Tienshan,Pamiro-Alai,
Pamir,Dzhungarian
Alatau.

phenomena,Ecoregionsthat containextensivein- ests of the Amazon (varzea forests) (Goulding,

tact habitatsandlargevertebrateassemblageswere 1980; Gouldinget al,, 1996), Such phenomena
recognized,Also consideredwerethe long-distance wereonce widespreadbut are nowraredue to the
migrationsof largerterrestrialvertebratessuch as prevalenceof humandisturbancearoundthe world,
caribouorwildebeest,andthe tremendousseasonal This is the only situationwherewe considerglobal
fish migrationsandfish frugivoryin the floodedfor- patternswithin the context of threats,Otherwise,

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Volume 89, Number 2 Olson & Dinerstein 209
2002 The Global 200

Table3. Actualor estimatedvascularplantspeciesrichnessandendemism(expressedas speciesnumber)of some

tundraecoregionsor regionsbasedon datafromWWF/IUCN(1994), Rickettset al. (1999), and J. T. Kartesz(pers.
comm.).
E.
Speciesrichness Endemism
.

coreglon or reglon

Nearctic
Pacific Coastal MountainIcefields 792 o
Alaska/St. Elias Range Tundra 747 1 10
Interior Yukon/AlaskaAlpine Tundra 617 4-10
Brooks/BritishRange Tundra 593 1-3
Ogilvie/MacKenzie Alpine Tundra 589 1 10
Arctic Foothills Tundra 580 o
Beringia Lowland Tundra 553 o
Arctic Coastal Tundra 539 1-3
Beringia Upland Tundra 538 1-3
Low Arctic Tundra 497 o
Aleutian Islands Tundra 388 1 10
Middle Arctic Tundra 371 1-3
TorngatMountainTundra 286 o
High Arctic Tundra 245 o
David Highlands Tundra 216 o
Baffin Coastal Tundra 135 o
Palearctic
Chukotsky Peninsula 939 -50
TaimyrPeninsula 240 5

the Global 200 emphasizeshiodiversityfeatures rallyoecurringrarity,althoughhuman-in(lucedrar-

thatwerein plaee [)riorto majorhumanimpaetsof ity ix an importantconditionto assexxwhendevel-
naturalhatitats and speeies populations. opingc onservationstrategiex.
Bothecologicaland evolutionary phenomenaare
a critical, but widely overlooked,aspect of bio(li- IN'l'A(v'l'N >,%S
versity conservation.Unusual evolutionaryphe-
nomenasuch as the extraordinary adaptiveradia- For ecoregionsin the same biomethat were as-
tions seen in Hawaiianplants,birds, and insects, xessed at a similarlevel of biologi(al importance,
the radiationof Galapagosfinches,the radiationof we xelectedthe ecoregionsthathad relativelymore
cichlidsin RiftValleylakes of Africa,also elevated intacthabitatsand biotas (see conservationstatus
some ecoregionsto the Global200. While evolu- below).
tionaryor ecological phenomenaoccur in every
14FItItESENTA'rl()N
ecoregion,we highlightthosethatarerecognizedas
exceptionalin globalcomparisons.
Ecoregionswerealso elevatedto Global200 sta-
GLOBAIJ RARITY
tus if they were the best exampleof their biome
withina realmin situationswhereno otherecore-
All ecoregionsin globallyrarebiomeswerecon- gion had been selected due to its outstandingbio-
sideredpriorities.Weelevatedecoregionsto Global diversity.In this selection we emphasizedthose
200 statusif theirbiomeor majorhabitattypewas ecoregionsthat harboredthe richest or most en-
representedin fewer than eight distinct regions demic biotas, or had the most intact naturaleco-
aroundthe world.Examplesof rarebiomesinclude systemsif biologicalimportancewas similaramong
the six Mediterraneanwoodlands,forests, and candidates.
scrub,all of limitedarea.Temperate
rainforesteco- The Global200 focuses on biologicalvalues as
systems(a majorhabitat)occurin seven relatively the criticalfirststep in settingglobalconservation
localizedareas aroundthe world.Paramos,or wet priorities.Therearemanyotherfactorsthatmaybe
tropicalalpine shrublands,occurin only a few ar- used in the prioritization process.Wepurposefully
eas of the northernAndes and CentralAmerica,a did not use ecologicalfunction,conservationfea-
few East African mountainranges, and in New sibility (i.e., political, social, economic, cultural
Guinea.For this criterion,we countedonly natu- factors),or humanutilityas discriminators to iden-

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 Annals of the
210 Missouri Botanical Garden

tify the Global200 as these featuresare eitherdif- size of remaininghabitatblocks, degree of frag-
ficultto measureor are highlyfluid.The develop- mentation,degree of degradation,and degree of
ment and implementationof ecoregionstrategies, protection(see Appendix2). Weightingsforfactors
however,require careful attention to ecological variedby biomefor freshwaterand marineecore-
functionand non-biologicalfactors. gions.
The recognitionof remainingwild animalmigra-
tions and othercontemporary ecologicalphenome- THEGLOBAL
200 ECOREGIONS
na is the onlycriterionwherehumanimpactsto the
environmentare recognized,because areas of ex- Weidentified238 ecoregionswhosebiodiversity
tinguishedphenomenaare ignored.Otherwise,the and representationvalues are outstandingor sig-
Global 200 emphasizesbiodiversityfeaturesthat nificanton a globalscale (Table1). Theyrepresent
werein place priorto majorhumanimpactson nat- the terrestrial,freshwater,and marinerealms,and
ural habitatsand species populations. the 30 biomesnested withinthese realms.Among
the three realms, 142 (60%) are terrestrial,53
CONSERVATI()N
STATUS
OF THEGLOBAL
200 (22Wo) are freshwaterecoregions,and43 (18to)are
ECOREGIONS marine.Terrestrialecoregionsoutnumberthose of
the otherrealmslargelybecause thereis morelo-
Ecoregionsvarygreatlynot only in theirbiolog- calizedendemismin terrestrialthan in marinebi-
ical distinctiveness,but also in their conservation otas. Gaps in biogeographicinformation for fresh-
status. Conservationstatusrepresentsan estimate waterandmarinebiodiversityalso accountforsome
of the ability of an ecoregionto maintainviable of the variation.
species populations,to sustainecologicalprocess-
es, and to })e responsiveto short-and long-term TERRESTRIAL
REALM
environmental changes.Conservation statusassess-
mentsof the Global200 ecoregionswerebased on Tropicaland subtropical
moistforests
landscapeor aquascape-levelcriteria,such as total Among the 14 terrestrialbiomes, the largest
habitat losse the degree of fragmentation,water numberof Global200 ecoregionsfalls withinthe
quality,and estimatesof futurethreat.Froma prac- tropicaland subtropicalmoist forests biome (50
tical perspeetive, a measureof conservationstatus ecoregionsor 35to of all terrestrialecoregions)(Ta-
can dictate the urgency,kinds of conservationac- ble 1). The high numberof ecoregionsreflectsthe
tivities,andlevel of effortneededamongecoregions biologicalrichnessandcomplexityof tropicalmoist
or biomes.Conservation statuscan also indicatear- forests.Althoughthereare moretropicalmoistfor-
eas withrelativelyhighopportunity forfar-reaching est ecoregionsin the Indo-Malayan Biogeographic
conservationmeasures. realm(17)thanin the Neotropics(12),this is partly
We estimatedthe conservationstatus of ecore- due to the archipelagicdistributionsof Asiantrop-
gions specificallyto enable us to make decisions ical moist forests and their characteristicbiotas
about elevatingecoregionswhen the similarityof (Whitmore,1986, 1990; Whitten et al., 1987a,
their biodiversityfeatures made discrimination 1987b, 1996; Wikramanayake et al., 2001). Four
challenging.Conservationstatuswas also used to of the Asian tropicalmoistforextsare smallisland
assess broadtrendsin threatsamongdifferentre- systems,and the originalextentof all of the Asian
gionsand biomes.Again,we drewheavilyfromre- ecoregionsfits easily within the area coveredby
glona conservahonassessmentsto estlmatecon-
. . .

westernAmazonianmoistforestx.
servationstatus.7Forthe Global200, we classified The most diverseterrestriale( oregionsoccurin
ecoregionsinto one of threebroadcategories:crit- the WesternArc forestsof the AmazonBasin,with
ical/endangered,vulnerable,or relatively stable/ close rivalsin the AtlanticForestecoregionof Bra-
relativelyintactoverthe next 40 years.Forterres- zil, the Choc6-Darienecoregion of northwestern
trial ecoregions,the mostprominentcontributor to SouthAmerica,Sumatra,and PeninsularMalaysia
conservationstatusis habitatloss, followedby the and northernBorneoforest ecoregions.The mon-
tane forest biotas of the NorthernAndes are re-
t IUCN(1991, 1992), Kreveret al. (1994), BSP et al. markablefor their globallyhigh rates of beta-di-
(1995), Dinersteinet al. (1995), Harcourtet al. (1996), versityand extraordinary local endemism(Terborgh
MacKinnon& Bunting(1996), Bryantet al. (1997), Di- & Winter,1983;ICBP,1992;Hamiltonet al., 1995;
nersteinet al. (1995), Dobsonet al. (1997), Rickettset
al. (1999),Abell et al. (2000),Bryantet al. (2000), Con- Wege & Long, 1995; WWF/IUCN,1994-1997).
servationInternational(2000), Wikramanayake et al. The forestsof the Guayananregionand Cubaare
(2001), Burgesset al. (in press). knownfortheirpronouncedendemismandunusual

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Volume 89, Number 2 Olson & Dinerstein 211
2002 The Global 200

biogeographicrelationships(Whitmore& Prance, (Gentry,1993; Parkeret al., 1993; Bullocket al.,

1987; Borhidi,1991; Dinersteinet al., 1995; Stey- 1996). The dryforestsof the PacificCoastof north-
ermarket al., 1995; Hedges, 1996). The forestsof westernSouthAmericasupporta wealthof unique
the GreaterAntilles also are notablefor a number species due to theirisolation(Parker& Carr,1992;
of relict mammals,such as solenodonsand hutias. WWF/IUCN,1994; Bullocket al., 1996). The sub-
The Congoliancoastal forestsare likely the most tropicalforestsof Maputaland-Pondoland in south-
diversein the Afrotropics,althoughdiversityinfor- eastern Africa are diverse and support many
mationis scarcefor severalecoregionsin the cen- endemics(Cowling& Hilton-Taylor, 1994; WWF/
tral Congo Basin (Oates, 1996; Kingdon,1997; IUCN,1994). The dry forestsof centralIndiaand
Burgesset al., in press).The Guineanmoistforests Indochinaare notablefor their diverse large ver-
supportmanyspecies not foundin the CentralAf- tebratefaunas(Corbett& Hill, 1992; Stewart-Cox,
rican region (IUCN/UNEP,1986a; IUCN, 1990; 1995). Dry forests of Madagascarand New Cale-
Martin,1991; IUCN, 1992; Mittermeieret al., doniaare globallydistinctivebecauseof theirhigh
1999). The AlbertineRift montaneforestsare ex- numberof relictual taxa and extremeendemism
tremelyrichfor sometaxa,such as birds,and have (IUCN/UNEP/WWF, 1987; Preston-Mafham, 1991;
a highdegreeof endemism(Collar& Stuart,1988; WWF/IUCN,1994; Wikramanayake et al., 2001).
Kingdon,1989; WWF/IUCN,1994). The distinc-
tivenessof the EasternArc montaneand East Af- Tropicaland subtropicalconiferous
forests
ricancoastalforestsis attributable to theirgreatage
and isolation (Hamilton& Bensted-Smith,1989; Mexico harborsthe world'srichest and most
Lovett& Wasser,1993; Hamiltonet al., 1995;Bur- complex subtropical coniferous forests (Perry,
gess et al., in press).Madagascar forestsandshrub- 1991; Petersonet al., 1993; Ramamoorthy et al.,
lands are also highly distinctiveon global scales, 1993; WWF/IUCN,1994). The conifer forests of
especially at higher taxonomiclevels (Nicoll & the GreaterAntilles containmany endemics and
Langrand,1989; Preston-Mafham, 1991; WWF/ relictualtaxa (Borhidi,1991). Subtropicalconifer
IUCN,1994). Tropitalmoistforestsof NewGuinea forestsof Indo(hinaare in(orporatedinto the dry
are highly distinetive (Brookse1987; Flannerye and moistforextsof the region.
1990, 1994; WWF/IUCN,1994; Mittermeier et al.,
1996; Wikramanayake et al., 2001), althoughAus- Temper(lte l)roa(lle(lJ an(l m,i.xe(lforests

tralianmoist forestsdo share manyaffinitieswith Tempelatebroa(lleafan(l mixedforestsare rich-

New Guinea The long-ixolatedforestsof New Cal- est in central China and eastern NorthAmerica,
edoniaareexceptionallyunusual,withso manyen- withothergloballydistinctiveecoregionsoeeurring
demic and relict highertaxa and species that the in the Cau(asus,the Himalayas,southernEurope,
islandis consideredthe 'Madagascar of the Pacific.' and the Russian Far East (Table2) (Zhaoet al.,
The forestsof Sulawesiare notedfor their region- 1990; Martin et al., 1993; Oosterbroeke1994;
ally high degreeof endemismin a rangeof taxa,a WWF/IUCN,1994; MacKinnon& Hicks, 1996;
phenomenonalso seen in the Philippinemoistfor- Rickettset a1., 1999).
ests and in the LesserSundassemi-evergreen for-
ests (IUCN/UNEP,1986b; IUCN, 1991; ICBP, Temperate
coniferous
forests
1992; Stattersfieldet al., 1998; Wikramanayake et
al., 2001).The WesternGhatsandsouthwestern Sri
Temperaterain forests only occur in seven re-
Lankanmoist forests are distinctivedue to their
gionsaroundthe world the PacificNorthwest,the
isolationand stabilityof conditionsovermillionsof
Validivianforests of southwesternSouthAmerica,
years.Tropicalmoistforestson oceanicislandsarethe rainforestsof New Zealandand Tasmania,the
oftenhighlydistinctivedue to high rates of ende-
NortheasternAtlantic (small, isolated pockets in
mism,extraordinary radiationsof taxaandadaptive
Ireland,Scotland,and Iceland),southwesternJa-
radiation,andrelictualor uniquehighertaxa(Dahl,
pan, and those of the easternBlack Sea (Kellogg
1986; Mitchell, 1989; Johnson &t Stattersfield,
et al., 1992; WWF/IUCN,1994). Forest commu-
1990; Flannery,1994; WWF/IUCN,1994; Wagner nities dominatedby huge trees (e.g., giantsequoia,
& Funk,1995). Sequoiadendron gigantea(Lindl.)J. Buchholz;red-
wood,Sequoiasempervirens (D. Don)Endl.;moun-
Tropicaland subtropicaldryforests tain ash, EucalyptusregnansF. Muell.)areunusual
ecologicalphenomenathat are foundonly in west-
The mostdiversedry forestsin the worldoccur ern NorthAmerica,southwesternSouth America,
in southernMexicoand in the Bolivianlowlands and in the Australasianregion in such areas as

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212 Annals of the
Missouri Botanical Garden

southeasternAustraliaand northernNew Zealand. unusuallyhigh levels of endemismand betadiver-

The Klamath-Siskiyou ecoregionof westernNorth sity in plants for tropical savannas.The tropical
Americaharborsdiverseand unusualassemblages savannasof northernAustraliaand southernNew
and displays notableendemismfor a numberof Guineasupportdistinctivecommunitieswith sev-
plant and animal taxa. The Valdivianforests of eral pocketsof endemismfor a rangeof taxa(Stat-
Chile are notablefor theirdiversityof tree genera, tersfieldet al., 1998).
manyof which are monotypicand have Gondwan-
aland origins. These long-isolatedforests have Temperate
grasslands,savannas,and shrublands
manyotherunusualtaxaand uniquecommunities.
The vastexpansesof grassin NorthAmericaand
Borealforestsand t(liga Eurasiaonce sustainedvastmigrations of largever-
tebratessuch as buffalo (Bison bison)and saiga
Low species richnessand endemismare char- (Saiga tatarica). Such extraordinaryphenomena
acteristicof circumborealand circumpolarecore- now occuronly in isolatedpockets,such as on the
gions (USSRAcademyof Sciences, 1988), thusthe DaurianSteppe(Kreveret al., 1994; Hilbig,1995;
presence of intact ecologicalphenomenadenoted Finch, 1996).The extraordinary
floralcommunities
outstandingecoregions.Large-scalemigrationsof of the Eurasiansteppes and the NorthAmerican
caribou,or reindeer(Rangifertarandus),andintact GreatPlains have been largelyextirpatedthrough
predatorassemblagescan still be found in some conversion to agriculture.Nearly 300 different
regions.For example,the NorthernCordillerabo- plantspecies can occuron a few hectaresof North
real forests of Canadahave been called the Ser- Americantallgrassprairie.The Patagoniansteppe
engetiof the FarNorthdue to theirabundanceand and grasslandsare notableforendemichighertaxa
diversityof largevertebrates(Rickettset al., 1999). for mammals.
Extensivetractsof borealforestandtaigastill exist
in the northernNearcticand Palearctic,the largest Floodedgrasslandsand savannas
expanses being in central and eastern Russia
(Stewart,1992; Kreveret al., 1994). This biome Somegloballyoutstandingfloodedsavannasand
also enjoysrelativelyunalterednaturaldisturbance grasslandsoccur in the Everglades,Pantanal,Sa-
regimes,an increasinglyraresituationin otherbi- helianfloodedsavannas,Zambezianfloodedsavan-
omes. nas (includingthe OkavangoDelta),andthe Sudd.
The Evergladesare the world'slargest rain-fed
Tropicaland subtropical
grasslands,savannas,and flooded grasslandon a limestonesubstrate.The
shrublands floodedsavannasand grasslandsselectedare gen-
erallythe largestcomplexesin each region.Anoth-
In manypartsof the tropicslarge mammalfau- er extraordinaryinlanddelta,the Mamberamo Riv-
nas have evolvedto take advantageof the produc- er inland delta, is capturedwithin the montane
tive grassesandbrowsetypicalof this biome.These forestsof the New Guineaecoregion.
largemammalfaunasare richestin Africansavan-
nas and grasslands.Presentlythe most intact as- Montanegrasslandsand shrublands
semblagesoccur in East Africanacacia savannas
and Zambeziansavannascomprisedof mosaicsof The paramosof the northernAndesarethe most
miombo,mopane,and otherhabitats(McClanahan extensiveexamplesof this biome.Paramoecosys-
& Young,1996). Large-scalemigrationof tropical temsoccurin onlya few otherlocalitiesin the trop-
savannaherbivores,such as wildebeest (Conno- ics. The heathlandsand moorlandsof East Africa
chaetestaurinus)and zebra(Equuszebra),are con- (e.g., Mt. Kilimanjaro,Mt. Kenya,RwenzoriMts.,
tinuing to decline throughhabitatalterationand EthiopianHighlands),Mt.Kinabaluof Borneo,and
hunting.Onlyin EastAfrica,the centralZambezian the CentralRangeof New Guineaareall limitedin
region,and in the Suddregion(Ugandakob orKo- extent, extremelyisolated,and supporthighlyen-
buskob)do sizablemigrationsstill persist.Manyof demic plantsand animals.A characteristic feature
the extraordinarymigrationsof the GuineanandSa- of manytropicalparamosis the presenceof large
helian savannashave disappeared.Sahelianecore- rosetteplantsfroma varietyof plantgenera,such
gions supporta large numberof endemic rodent as Lobelia(Africa),Puya (SouthAmerica),Cyathea
species, while the Somalianbushlandand thickets (NewGuinea),andArgyroxiphium (Hawaii) these
harbora concentration of endemicmammals,from plantformscan reach elevationsof 4500-4600 m
rodentsto antelopes.Boththe Cerradoand the Lla- above sea level. Drier,yet distinctive,subtropical
nos are noted for complexityof habitatsand the montanegrasslands,savannas,and woodlandsin-

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Volume 89, Number 2 Olson & Dinerstein
2002 The Global 200 213

clude the Ethiopian Highlands,the Zambezian of the Earth'splantspecies (Cody,1986; KalinAr-

montanegrasslandsand woodlands,and the mon- royoet al., 1995; Picker& Samways,1995). Phy-
tane habitatsof southeasternAfrica(Werger,1978; togeographersconsiderthe Fynbos as a separate
White, 1983; Huntley,1989, 1994; Timberlake& floralkingdombecause 68% of the 8600 vascular
Muller,1994; WWF/IUCN,1994). The montane plant species crowdedinto its 90,000 km2are en-
grasslandsof the TibetanPlateaustill supportrel- demic and highly distinctiveat severaltaxonomic
ativelyintactmigrationsof Tibetanantelope(Pan- levels (Cowlinget al., 1989, 1996; Cowling& Hil-
tholopshodgsoni)and kiang,the Tibetanwild ass ton-Taylor, 1994).In termsof speciesdensities,this
(Equushemionus).The punagrasslandsof the high is equivalentto about40 percentof the plantspe-
Andes supportover30 species of endemicrodents cies of the United States and Canadacombined,
(45 totalspecies). foundwithinan areathe size of the stateof Indiana
(N. Myers,pers. comm.).The Fynbosand South-
Tundra west Australiashrublandshave florasthat are sig-
Tundraecoregionswere selected primarilybe- nificantlymorediversethan the otherecoregions,
cause of extraordinary seasonalconcentrationsof althoughany Mediterranean shrublandis still rich
breedingwaterfowland shorebirds,and caribou in species andendemicsrelativeto othernon-forest
(Stewart,1992; Kreveret al., 1994;Rickettset al., ecoregions (Cowling et al., 1996; Oosterbroek,
1999). Relatively intact tundra ecoregionswere 1994).
chosen,whereverpossible.The Chukotskytundra
ecoregionis unusualwithnearly50 endemicplant Mangroves
species (Knystautas,1987; USSRAcademyof Sci-
ences, 1988; WWF/IUCN,1994). The diversityof mangrovesin the Indo-WestPa-
cific (IWP)regionis muchgreaterthanthoseof the
Desertsatld xeric shrublan(l.s Atlantic-Caribbean-East Pa(ific (ACEP)region-
the formersupporting17 generaand40-42 species
The Namib-Karoo desertsof southwesternAfrica of truemangrovesand the latterhavingonly4 gen-
supportthe world'srichest(lesertfloras(Cowling& era and 7 species (Ma(Nae,1968; I,a(erda,1993;
Hilton-rraylor,1994; Maggset al., 1994; WWF/ Olsonet al.! 1996; Spal(linget al., 1997; Rickleffs
IUCNe1994) while the ChihuahuanDesert an(l 8z I atham,199.3).A single site in the ACEPtypi-
c entralMexicandesertsare a close seeon(lan(tare c ally ( ontains3 or 4 true mangrovespeeies, while
the richest Neotropie al deserts (Cowlinget al., 30 see ies havebeen re(or(tedfromone loealityin
1989; Hernandez& Bareenas, 1995; Rieketts et the IWP region(Ricklefs& I,atham,1993). Man-
al., 1999). Australiandeserts supportthe ri(hest groveforests on the western(oast of Madagascar
reptilefaunas.The CarnavonXeric Scrubof west- supporta numberof endemicbirdspecies thatare
ern Australiais a regionalcenterof endemismfor endangered.The mangroveswampsand forestsof
a rangeof taxa. Unusualdesertcommunities dom- the Indo-Malayan and Australasianrealmsare the
inatedby giantcolumnarcactioccurin the Sonoran world'smost extensive.South and SoutheastAsia
and BajaDesertsof NorthAmerica(Brown,1994), alone eontain42Woof the total area of the world's
while the spiny thicketsof southwestern Madagas- mangroves(Spaldinget al., 1997). The Sundarbans
car are globallyuniquein termsof structureand are the largestcontiguousmangroveforest in the
taxa. Some Baja Californiacommunitiesare par- world.Thevastfloodplainsof NewGuineaalso sup-
tially convergentin structurewith the Madagascar port extensive mangroveswampsunrivaledelse-
thickets.The AtacamaDesertecoregionof western wherein the world.
South America(includingthe adjacenttransition If all of the marine,freshwater,and terrestrial
area of the Monte/Puna/Yungas) and the Horn of species that occur in mangrovesare considered,
Africadesertswererecognizedas someof the more these seeminglysimple forestscan be considered
outstandingregionalcenters of richness and en- as one of the morediverseecosystemsin the trop-
demism.The CentralAsian deserts,while not as ics. Mangrovesare keystonehabitatsin the sense
richas Afrotropical or Neotropicaldeserts,arerep- that they have an inordinatelystronginfluenceon
resentativeof the region'sdesertswithdiverserep- species populationsand ecosystemswell beyond
tile and mammalfaunas. their limitedarea. In additionto providinghabitat
and resourcesto a wide rangeof species, mangrove
Mediterranean forests,woodlands,and scrub forestsand swampsalso protectinlandhabitatsand
All five Mediterranean-climate ecoregionsare shorelinesfromdamageby dampingstormwaves
highlydistinctive,collectivelyharboring20 percent andtidalaction.Mangroves filtersilt andpollutants

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214 Annals of the
Missouri Botanical Garden

fromterrestrialrunoffthatwouldotherwisedamage one of the Earth'srichesttemperatefreshwaterbi-

seagrassbeds and coralreefs. otas (Hocutt& Wiley,1986; Hackneyet al., 1992;
Abell et al., 2000). The headwaterstreamsandriv-
FRESHWATER REALM ers of the YangtzeRiverin centralChinaare also
extremelydiverse(recognizedas a large riverbi-
Largerivers ome in this analysis)(Mori,1936; Nichols, 1943;
Faunasadaptedto high-flowregimesof largeriv- Taki,1975). Secondarycentersof temperatediver-
ers are uncommonandbest developedin the Yang- sity occurin the riversand streamsof southeastern
tze, Colorado,and lowerCongoRivers.A relatively NorthAmerica,the westerncoastof NorthAmerica,
smallareaof rapidsin the latterregionsupports22 and the RussianFar East (Zhadin& Gerd,1961;
endemic species of fish that are rapid specialists Lee et al., 1980; Hocutt& Wiley, 1986; Groom-
(Lowe-McConnell, 1987). The Mekong,Congo,Pa- bridge& Jenkins,1998;Abell et al., 2000). Several
rana,and Amazon-Orinoco Riversharborthe four freshwaterbiotason islands are highlydistinctive,
greatlarge tropicalriverfish faunas (Mori,1936; includingthose of Madagascar,New Guinea,the
Roberts, 1975; Hocutt & Wiley, 1986; Lowe- GreaterSundas,the GreaterAntilles, Sri Lanka,
McConnell,1987; Kottelat& Whitten,1996). The and New Caledonia (IUCN/UNEP/WWF, 1987;
watersof the LowerYangtzeandMississippiRivers Zakaria-Ismail, 1987, 1994; Allen, 1991; Preston-
containoutstandingexamplesof large-riverfishes, Mafham,1991; Oberdorffet al., 1995). The South-
amphibians,reptiles, and invertebrates,including west AustralianRiversand streamsecoregionis a
relicts and manyendemics(Abellet al., 2000). centerof endemism,while also harboringa number
of primitivehighertaxa and several species with
Largeriverheadwaters highlyunusualfreshwaterlife histories(McDowall,
1996; Stateof the Environment AdvisoryCouncil,
Speciese assemblages,and processes in head- 1996). Riversandstreamsalongthe Gulfof Guinea
waterareas are distinctfromthose of their larger harborsome of the richest and most endemicriv-
mainstems.The MississippiPiedmont,Guayanan erine freshwaterbiotas in Africa(King(ton,1989;
highlandseUpperAmazon,UpperParana,Brazilian Levequeet al., 1992; Leveque,1997; Thiemeet
Shield, and CongoBasin Piedmontharbora tre- al., in press).The SalweenRiverof SoutheastAsia
mendousarrayof species, includingnumerousen- is recognizedfor its rich and endemicfreshwater
demics adaptedto life in these waters. In turn, fish fauna(WCMC,1992). The riversand streams
these riversystemsultimatelyfeed a numberof the of New Guinea,includingthe inland delta of the
world'slargestand richestrivers(Hocutt& Wiley, Mamberamo Riverof New Guinea,supporta large
1986; Kottelat& Whitten,1996; Thiemeet al., in numberof unusualandendemicspeciesandhigher
press).The mostdiversevertebrateassemblageson taxa.
Earthoccurin freshwatercommunitiesof the Am-
azonand the OrinocoRiverbasins.Over3000 spe- Largelakes
cies of fish are estimatedto occur in the Amazon
Basin alone (Goulding,1980). The Global200 also identifiesthe mostoutstand-
ing examplesof diverse and endemiefreshwater
Largeriverdeltas faunasin largelakes foundin temperateand trop-
ical regions,manydisplayingextraordinary species
Delta complexesof severallarge temperateand flocks and adaptiveradiationsin fish taxa. Some
polar riversare identified,includingthe Mesopo- particularly notablelake biotasincludethoseof the
tamian,Volga,and Lena River deltas. The Niger African Rift Lakes and Lake Tana in EthiopiaS
Riverdelta,the mostextensiveriverdeltain Africa, Lake Baikal, Lake Biwa of southernJapan the
is characterized by highspecies richness(Wetlands high-altitudelakes of the Andes,and the highland
Internationaland The WorldBankS1996; Thieme lakes of Mexico(Myers,1960; Roberts,1975; Ho-
et al., in press).Theextensivedeltasof the Orinoco cutt & Wiley, 1986; Allen, 1991; Stiassnyet al.,
and AmazonRivers are encompassedin their re- 1992; WCMC, 1992; Nagelkerkeet al., 1995; Kot-
spectivelarge-riverecoregions(see above). telat & Whittert,1996; Olsonet al., 1999;Thieme
et al., in press).
Small riverbasins
Small lakes
The MississippiRiver embayment^, the Mobile
Riverbasin,andnumerouscoastalstreamsandriv- Similarly,a numberof smallerlakes aroundthe
ers of southeasternNorthAmericatogethersupport worldhost extraordinary expressionsof freshwater

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Volume 89, Number 2 Olson & Dinerstein
2002 The Global 200 215

biodiversity.LakeKutubuandLakeSentaniof New ated with isolatedislands and enclosed seas tend

Guinea, Yunnan Lakes and Streams, Mexican to displaypronouncedendemism(Kelleheret al.,
HighlandLakes,the CameroonCraterLakes,Lake 1995; Groombridge & Jenkins,1996).
Lanaoof the Philippines,Lake Inle in Myanmar Wecategorizedthe marinerealminto 10 biomes.
(Burma),andthe CentralSulawesiLakeshavebeen Pelagic (tradesand westerlies),abyssal,and hadal
selected for their globallyoutstandingbiodiversity biomes,however,were not assessed for the Global
features. 200 marineanalysisbecause of the large scale of
these units comparedto other Global200 ecore-
gions,the lack of consensuson theirclassification,
. , .

Xerzc oaszns
and the limited biodiversityinformationfor these
Ephemeralstreams,rivers,and lakes, and per-
ecosystems(see Gage& Tyler,1991;Grassle,1991;
manentspringscharacterizeecoregionsin this bi-
Grassle & Maciolek,1992). Largebiogeographic
ome. Lowrichnessand high endemismin fish and
units have been identifiedfor pelagic and abyssal
invertebrates(e.g., molluscs)is typicalof the Chi-
biotas(e.g., Brinton,1962;Angel,1993;Longhurst,
huahuan,Anatolian,and CentralAustralianfresh-
1998; Pierrot-Bults,1997; Vinogradova, 1997), but
water ecoregions(Hocutt& Wiley, 1986; Balik,
their scale is several ordersof magnitudegreater
1995; Abell et al., 2000). The CuatroCienegas
than most Global 200 ecoregions.These larger
springandpool complexin the Chihuahuan Desert
units may be biogeographically and dynamically
is globallyuniquein its highrichness,extremeen-
appropriate for open ocean environments.
The vast
demism, and unusual evolutionaryadaptations
size and dynamicnatureof these biomesprecluded
(Contreras-Balderas, 1978; Hocutt& Wiley,1986).
delineatingbiogeographicsubunitsat an appropri-
Freshwater habitatsin the Anatolianregionof Tur-
ate level of resolutionfor the Global200. Pelagic
key supportmanyendemicspecies (Balik,1995).
species are noted for widespreaddistributionse
while the few ocean trenchsurveysthat are avail-
MARINE REAI M
able suggest many spe(ies are endemic to single
The (listributionof marine biodiversityvaries trenches.The paucityof spe(ies dataforthese eco-
widelythroughoutoeean basins (Briggs,1974; E1- systemsalso reducesour (onfiden(e to undertake
der & Pernetta,1991; Angel, 1992! 1993; Clarke, comparativeanalyses.
1992; Kendall& As(hane 1993; Kelleheret al.,
1995; Groombridge & Jenkins, 1996; Ormon(let Polar
al., 1997). The abundan(ean(l diversityof most
taxatend to be highestnearcontinentaland islan(l The WeddellSea an(lPeninsularAntaretica were
marginsthatareless than2000 m deep (Ray,1991; identifie(las the mostproduetive an(ldiverseecore-
Johannes& Hatcher,1986; Gray,1997). These ar- gionsof the Antarcticlargemarineecosystem.The
eas experiencenutrientenrichmentfromupwelling Bering,Beaufort,and ChukehiSeas and Barents-
processesand terrestrialrunoff(Ray,1988; Norse, Kara Seas ecoregionsare arguablythe two most
1995).Areaswheresignificantupwellingoccursare diverse and productiveArctic marineecosystems
often extraordinarilyproductivein tropical,tem- (USSR Academy of Sciences, 1988; Reeves &
perate,and polar regions.Withinbiomes,species Leatherwood, 1994). Marine ecosystems near
richness and endemism also vary enormously southernGreenlandrequirefurtherevaluation.
aroundthe globe.
Currentbiogeographicdata suggestthat species Temperateshelf and seas
endemismtends to be less pronouncedin marine
ecosystemsthan in terrestrialor freshwaterecore- Someof the mostproductivemarineecosystems
gions,but severalregionalcentersof endemismare occur in the GrandBanks and New Zealandplus
recognized,includingthe southerncoastof Austra- the Patagoniaecoregions.The South Australian
lia, New Caledonia,Lord Howe and NorfolkIs- coastal watersare remarkablefor unusuallyhigh
lands,the northerncoastof SouthAmerica,the Yel- levels of endemism in invertebratesand some
low and East China Seas, the Red Sea, the groupsof fish, in additionto the diverse marine
Mediterranean Sea, the Sea of Cortez,the Great mammalassemblagefoundthere.Twoof the world's
BarrierReef, and tropicalPacificIslandssuch as largest temperateestuaries,the Chesapeakeand
Hawaii,the Marquesas,the Tuamotusand Socie- DelawareBays, and the NortheastAtlanticShelf
ties, and EasterIsland (Robbins,1991; Lieske & are elevated to the Global200 due to their size,
Myers,1996; Vernon,1995; Groombridge & Jen- productivity,
andhabitatdiversity.Someof the most
kins, 1996). In general,marineecoregionsassoci- distinctiveenclosed temperateseas, the Mediter-

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216 Annals of the
Missouri Botanical Garden

raneanSea and the Yellow-EastChina Seas, are omes, ecoregionsfalling within the tropicaland
recognizedin the Global200. subtropicaldry broadleafforests,temperategrass-
lands, Mediterraneanshrublands,and temperate
Temperateupwelling broadleafand mixed forests are the most threat-
ened. Virtuallyall biotason smallislandsare vul-
Highlyproductiveand diversecoastalupwelling
nerableor critical/endangered due, in largepart,to
areasoccuralongthe WestCoastof NorthAmerica
theirlimitedhabitatareaandextremesensitivityto
where the CaliforniaCurrentmoves southward.
anthropogenicdisturbanceand alien species (Ra-
Along the southwestcoast of Africathe Benguela
ven, 1988;Wilson,1988, 1992;WCMC,1992; Su-
Currentexhibitssimilardynamics.
jatnikaet al., 1995; Brookset al., 1997; Reaka-
Kudlaet al., 1997).Islandecoregionsareprojected
Tropicalupwelling
to experiencea wave of extinctionsover the next
The HumboldtCurrentalongthe West Coastof two decades given the fragilityof island ecosys-
SouthAmericaand the CanaryCurrentalong the tems, the sensitivityand endemicityof islandspe-
WestCoastof Africabringrich nutrientsto the sea cies, and the severe threats native island biotas
surfacewherethey supporthighlyproductivema- face worldwide.Mangrovehabitatsare threatened
rine systems.Important tropicalupwellingandcur- worldwidefroma rangeof threatsincludingclear-
rent areas also occur in the PanamaBight ecore- ing and channelization for shrimpponds,aquacul-
gions. ture, and agriculture,the extractionof timberand
fuelwood,pollution,and habitatloss due to urban
Tropicalcoral and industrialexpansion.
Assessmentof conservationstatusforfreshwater
SoutheastAsianseas supportmorethan450 spe-
ecoregionsin NorthAmericaand SouthAmerica
cies of hard(scleractinian)corals,the westernIn-
was based on existingregionalanalyses(Abell et
dianOeean around200, andthe Caribbeanonly50
al., 2000; Olsonet al., 1999).In AfricaandEurope,
species (Vernon,1995). Variationin reef fish and
analyses currentlyunder way (Thiemeet al., in
non-coralinvertebratediversityfollows a similar
press) providedthe basis for rankingspresented
biogeographicpattern (McAllisteret al., 1994;
here. In areas where no regionalassessmenthas
Lieske& Myers,1996).Overall,the coralreefcom-
been undertaken,reviewof relevantliteraturefa-
munitiesof the centralIndo-Pacificseas are the
cilitateddecisionson the levels of threatfaced by
mostdiversein the world,withthe Sulu, Sulawesi,
nativebiotas.Worldwide, freshwater organismsrep-
Banda,and CoralSea ecoregionsbeing the most
resent a disproportionate numberof endangered
diverseon Earth(Vernon,1995; Lieske & Myers,
specles; t aus, lt 1S not surprlslngt zat so many
. . . . .

1996). The largestbarrierreef in the worldis the

freshwaterecoregionsreceiveda criticalratingin
GreatBarrierReef. Otherworld-classbarrierreefs
the assessment.In particular,seasonallyflooded
includethe barrierreefsof NewCaledonia,the Me-
forests, cataracts,and freshwatercommunitiesin
soamericanand Bahamianbarrierreefs, and the
xeric areas,areendangeredworldwide(Gouldinget
large barrierreefs of Fiji. The largest coral atoll
al., 1996; Abell et al., 2000; Olson et al., 1999).
complexes occur in the Maldive-Lakshadweep
Moreover,most temperatefreshwaterbiotas are
ecoregionof the centralIndianOcean and in the
threatenedby invasionof exotics,pollution,dams,
Tuamotusof the centralPacific.
and habitatdegradation.Amongthe 53 freshwater
ecoregions31 (58Wo) were deemedto be criticalor
CC)NSERVATION STATUS OF ECOREGIONS
endangered,10 (l9Wo)wereassessedas vulnerable,
Amongall terrestrialGlobal200 ecoregions(142 and only 12 (23Wo) were assessedas relativelysta-
in total),75 ecoregions(53Wo) are consideredcrit- ble.
ical or endangered,39 ecoregions(27%)vulnera- The individualstatus of marineecoregionswas
ble, and 28 ecoregions(20Wo) relativelystable or estimatedthroughreviewof the literatureand con-
intact(Table1). Terrestrialecoregionboundariesdo sultationswith regionalspecialists.Twelvemarine
not reflectthe extensivehabitatloss, fragmentation, ecoregions(29Wo) were consideredrelativelystable
and degradationthathave occurredin manyof the or intact,while another12 (29Wo) wereconsidered
terrestrialecoregions.In ecoregionsthathavebeen criticalorendangered.In marinebiomes,upwelling
dramaticallyaltered, characteristicspecies and areasare heavilyoverfished,enclosedseas are de-
communitiessurvive only in the few remaining graded,andcoralreefs and mangrovesare severely
smallblocksof habitat(e.g., Collar& Stuart,1988; affected by habitatdestruction,degradation,and
Dinersteinet al., 1995). Amongthe terrestrialbi- overfishingaroundthe world(Shermanet al., 1990;

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Volume 89, Number 2 Olson & Dinerstein 217
2002 The Global 200

Suchanek,1994; Kelleheret al., 1995; Bryantet guidelines for biomes within realms. Biological
al., 1995; Olsonet al., 1996; Ormondet al., 1997). phenomenaare also importantcriteriaused in its
Increasinglyrising sea surfacetemperatures from selectionprotocol.The Global200 also emphasizes
globalwarmingmayendangerall coralreef ecore- freshwaterand marinebiodiversity.The Hotspots
gionswithinseveraldecades. analysis (Mittermeieret al., 1999; Myers et al.,
2000), for example,mostlytargetsvery large and
DEGREE
OFOVERLAP
OFTERRESTRIAL, threatenedterrestrialregionswithconcentrations of
FRESHWATER,
ANDMARINE
GLOBAL
200 range-restricted(locally endemic) species. The
ECOREGIONS Hotspotsare largelynested withinthe Global200
The linkages amongterrestrial,freshwater,and (> 90Wocongruence)because both analyses em-
marineconservationare often overlooked.Among phasizeexceptionallevels of endemismfor species
the Global 200, 33 (23Wo)of the 143 terrestrial and highertaxa. The Global200 can complement
ecoregions overlap extensively with freshwater hotspotanalysesby corroborating the vast majority
ecoregions(i.e., morethan 50%of the originalex- of their priorityareasand, in some cases, by pro-
tent of the terrestrialecoregionis covered by a viding a finer resolutionof the variationof biodi-
freshwaterunit).Thirty-four (23%)of the terrestrial versityfeatureswithin importantregions.For ex-
ecoregionsshare at least 50Woof their coastline ample,the Madagascar Hotspotidentifiedby Myers
witha marineecoregion.Ten(6%)of the terrestrial et al. (2000) correspondsto five separateGlobal
ecoregionsdo both,overlappingextensivelywith a 200 ecoregionsand the Indo-BurmaHotspotover-
freshwaterecoregionand sharingat least 50% of laps with 14 Global200 terrestrialand freshwater
theircoastlinewitha marineecoregion.The terres- ecoregions.The Global200 also encompassesdis-
trial ecoregionsof this thirdgroupare the Mada- tinct freshwaterand marine hotspotsand warm-
gascardryforests,Congoliancoastalforests,Great- spots, as well as ecoregionsimportantfor theirex-
er Antilles moist forests, Pacific temperaterain traordinaryecologicalor evolutionaryphenomena
forests of North America, Queenslandtropical andtheirrepresentation value.EndemicBirdAreas
moist forests, southeasternAustraliaEucaly,vtus- of the Worldhighlightsconcentrationsof birdspe-
Acaciaforests, New Caledoniamoistforests,New cies with restricted ranges (Stattersfieldet al.,
Caledoniadryforests,New Guinealowlandforests, 1998). I,ike hotspots,the majorityof the Endemic
Sulawesi moist forests, Philippinemoist forests, BirdAreasare nestedwithinthe Global200. Both
NortheastBorneo/Palawan moistforestseand Rus- Tropi(alForest WildernessAreas (Mittermeieret
sian FarEast temperateforests.Carefullydesigned al., 1999) and FrontierForests(Bryantet al., 1997)
conservationactivitiesin these 13 units could ul- map larger landscapes of relatively undisturbed
timatelyaffect39 ecoregions. natural forests around the world. Although the
Global200 does not specificallyemployforestwil-
THEGl OBAL200 AS A C()NSERVATION
T()OI
dernessas a discriminator, againthereis extensive
The Global200 is based on the best available overlapwith these wildernessareas because such
informationand biologicalinsights.As new inter- areasoftenharborrich assemblagesof species and
pretationsof biogeographyand better information endemics,and unusualphenomenasuch as intact
on the distributionof species and phenomenabe- predator-prey systems.
comeavailable,we expectto periodicallyrevisethe
Global200. The presentlist and map incorporate OTHER CONSERVATION TARGETS
a numberof changesfroman earlierversion(Olson
& Dinerstein,1998). For example,the highly un- Otherconservationtargets,such as species of
usual freshwaterbiotaof southwestern Australiais special concern, keystone species, habitats, and
now recognized,and the terrestrialecoregionsof phenomena, large-scale ecological phenomena
the AmazonBasinhave undergonemajorrevisions (e.g., bird, butterfly,caribou,cetacean, sea turtle
based on a recentbiogeographicanalysisby Silva
migrations),wildernessareas,amelioratingclimate
(1998).
change impacts,reducingtoxins, and maintaining
ecosystemswithlow impactsfromalien species are
EXPANDING CONSERVATION GOALS
also not directly addressedby the Global 200.
The Global 200 goes beyondthe conservation Again,effectiveconservationwithinpriorityecore-
ef- gionsandcoordinatedeffortsamongecoregionswill
targetsof otherprominentglobalpriority-setting
forts by explicitly incorporatingrepresentation help achieveconservationgoals for these targets.

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 Annals of the
218
Missouri Botanical Garden

ANAMBITIOUS
BLUEPRINT
FORGLOBAL Borhidi,A. 1991. Phytogeography andVegetation Ecology
CONSERVATION of Cuba.AkademiaiKiado,Budapest,Hungary.
Briggs,J. C. 1974. MarineZoogeography. McGraw-Hill,
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Onetacticalconcernof the Global2()0 is thatit Brinton,E. 1962. Thedistributionof Pacificeuphausiids.
is ambitious,and that by focusingon 238 ecore- Bull. ScrippsInst. Oceanogr.8: 51-270.
gions rather than on a handful of conservalion Brooks,R. R. 1987.Serpentineand ItsVegetation: A Mul-
units, we run the risk of placingless emphasison tidisciplinaryApproach.Ecology,Phytogeography and
the most diverse and dIstinctecoregions.In re- PhysiologySeries,Vol. 1. DioscoridesPress,Portland,
Oregon.
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reachof the Global200 makesalmosteverynation estationpredictsthe numberof threatenedbirdsin in-
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globalperspectiveforlobbyingeffortsby local con- Coastlinesat Risk:An Indexof PotentialDevelopment-
sewationgroups.TheGlobal200 also can helpma- relatedThreatsto CoastalEcosystems.WRIIndicator
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not shrinkfromthis ambitiousbut necessaryagen- LatinAmericaandthe Caribbean.A ReportforUSAID.
da. The widespreaddestructionof the Earth'sbio- BiodiversitySupportProgram,Washington, D.C.
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essarilyambitioustemplatefor a globalconserva- Olson.In press.Terrestrial EcoregionsofAfrica:A Con-
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Volume 89, Number 2 Olson & Dinerstein
2002 The Global 200 223

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freshwaterfishes in Southand SoutheastAsia. Walla- werethen summedto derivea single richnessand ende-
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. 1994.Zoogeography andbiodiversity of thefresh- withineachbiomeandthe ( urvesbrokensubjectivelyinto
waterfishesof SoutheastArsia.Hydrobiologia 285: 4]- high,medium,andlows( ores.Globallyoutstanding s( ores
48. were determinedthrough( omparisonswith values for
Zhadin,V. 1. 8 S. V. Ger(l.1961.Faunaand Floraof the e(oregionswithinthe same })iOtilP foundthroughoutthe
Rivers,Lakes,an(l lleservoirsof the USSR.Smithson- worl(l.
ian Institutionan(lNationalS(ience Foun(lationWash-
ington,D.C. UNUSUAL ECOLOCICAL OR EVOLUTIONARY PHENOMENA
Zhao,Ji, ZhellgGuallt,nleig WangHua(long& Xu Jialin
(editors).]99t). The NatulalHistoryof (3hina.M(Graw- (^lol)ally outstanding 100
Hill, New York. l4egionally outstan(lillg
Zohary,M. 1973. Geobotalli( al Foun(lationsof the Mi(ldle No glofally 01 regiollally UllUSUdI phellolllena ()
East,Vols. 1, 2. GustavFis(hel Verlag,Stuttgart,Ger-
many. IP,xamplets of unusuale(ologi(al or evvolutiollary phe-
notllenaat glolal or regionals( ales il( lude relativelyin-
Al'l'l1N1)1X1. WHI(->HIIN(JAN1) Ml1AXURIN(}Bl()l2()(rlCA12
ta(t, large-.s(alemigrationsot large verte})rates su(h as
DlFrl'lNCrRlvENI<,S,% CRXrl'ERIA
(ari})ou,inta(tpre(latoras.Semblages, supera})undant con-
entrationksof breedingwaterfowl alld,shore})irdsgextraor-
Theweightingan(lmeasurement of theparameters useel dinarylevels of adaptiveradiationzs,rain-fedfloodedgrass-
to assess the liologieal distinetivenessof terrestrial
eeore- lands on limestone,and c onifer forests dominatedby
gions of NorthAmeritaare presentedhere to illustrate glgantletrees. . .

howdifferentbiodiversityfeatureswereevaluatedas con-
servationtargetsand howanalysesweretailoredto differ- (,I,()ISAI, ItAIllrrY 0t' HI()Mi,
ent biomes.Comparisons amongbiodiversityparameters
wereonly conductedwithinthe set of ecoregionssharing Global rarity 100
the samebiome. Regional rarity 5
Not rare at global scale o
SPECIESRICHNFSSE
Biomesor habitatsthat were consideredgloleallyrare
Globallyoutstanding 100 include Mediterranean-climate forests, woodlands,and
High 15 scrub,temperaterainforests,and paramo.
Medium 10
TOTAL SCORES FOR DETERMINING BIOLOGICAL
Low 5
DISTINCTIVENESS INDEX
tOnly nativespecies wereused in species counts.
The pointsfromeach criterionweresummedto arrive
ENDEMISM
at a finalscore.This scorewas then translatedinto a bi-
ologicaldistinctivenesscategoryas follows:
Globally outstanding 100 Globallyoutstanding 45, 50, or 55+ points
High 25 Regionallyoutstanding 30, 35 40
Medium 15 Bioregionally
outstanding 20, 25
Low 5 Locallyimportant 10, 15

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224 Annals of the
Missouri Botanical Garden

Ecoregionsidentifiedas globallyoutstanding
weresub- atedwithdifferentlandscapescenarios.Forexample,total
sequentlycomparedwith similarecoregionsaroundthe habitatloss scenarioswererelatedto pointsas follows:
worldto validatetheirrelativestatus. % Originalhabitat Heavilyaltered Altered
90-1009 40 20
APPENDIX
2. ASSESSING
CONSERVATION
STATUS
OF 75-89% 30 15
ECOREGIONS 50-74% 20 10
10-49% 10 5
Conservationstatus measureslandscapeand ecosys- 0-9% o o
tem-levelfeaturesand relatesthese to the ecologicalin-
tegrityof ecoregions,namely,howwithincreasinghabitat An ecoregionreceivesbotha heavilyalteredscoreand
loss, degradation,
andfragmentation, ecologicalprocesses an alteredhabitatscore,whichrepresentsthe amountof
cease to functionnaturally,or at all, resiliencyto distur- habitatin each categoryForexample,consideran ecore-
bancedeclines,andmajorcomponentsof biodiversityare gion with 35% heavilyalteredhabitat(10 points),55%
steadily eroded. We assess the conservationstatus of alteredhabitat(10 points),andtherefore10%intacthab-
ecoregionsin the traditionof IUCNRed DataBookcate- itat. By combiningthe two scores, the ecoregionwould
goriesforthreatenedandendangeredspecies:critical,en- receivea total score of 20 points.Differentquantitative
dangered,andvulnerable.Forecoregionswe used the fol- and qualitativebiodiversityand landscapeecologychar-
lowing conservation status categories: critical, acteristicsare used to defineintact,altered,and heavily
endangered,vulnerable,relativelystable, and relatively alteredstatestailoredto the specificpatternsanddynam-
intact.Throughout all of the regionalanalyses,the specific ics of differentbiomes.Totalscoresforeachof theparam-
parameters andthresholdsusedforassessingconservation etersare summedto give a totalconservation statusindex
score.
statuswere tailoredto the characteristic patternsof bio- Snapshotscores weresubsequentlymodifiedby a 20-
diversity,ecologicaldynamics,and responsesto distur- yearprojectedthreatanalysisto arriveat a finalconser-
banceof differentbiomes. vationstatus assessment.Ecoregionsthat wereassessed
as facinghighthreatwereelevatedto a moreseriouscon-
neRRESTRIAL
ECoe( IoNS servationstatus.The threatanalysisestimatedthe cumu-
lativeimpactsof all currentandprojectedthreatson hab-
Wepresentthe methodusedto assessconservation sta- itat conversion, habitat degradation, and wildlife
tus for the terrestrialecoregionsof NorthAmericato il- exploitationusinga pointsystemassociatedwithdifferent
lustratethe approa(h (Rickettset al., 1999).The relative qualitativeandquantitative impacts.Usingan indexof 0-
( ontributionsof differentparameterswere as follows: 100 points,pendingthreatswithinan ecoregionwereas-
4()% habitatloss, 25% numberand size of remaining sessed and point totals assignedfor each of the above
blocksof intacthabitat,20% degreeof habitatfragmen- categories.Conversionthreatswere consideredto be the
tation,and 15%{legree of protection.A snapshotcon- mostserious,andthus habitatloss comprisedhalf(50) of
servationstatus was estimatedusing currentlandscape all possiblepointsin the weightingof threats.Forexam-
and ecosystem-levelparameters, usinga pointrangeof 0 ple, 50 pointswereassignedto conversionthreatsif 25%
to 100, withhighervaluesdenotinga higherlevel of en- or more of remaininghabitatwould be categorizedas
dangerment. The pointthresholdsfor differentcategories heavilyalteredwithin20 years.Forconversionof between
of conservationstatus were as follows:critical 89-100 10%and 24% of remaininghabitat,a scoreof 20 points
points, endangered65-88, vulnerable3744, relatively was assigned.The remainingtwo threats,habitatdegra-
stable 7-37, and relativelyintact04. Totalpointvalues dationandwildlifeexploitation, wereassessedusingmax-
weredeterminedby summingpointsassignedforeachpa- imumpointtotalsof 30 and20 respectivelyusinga scale
rameter.Individualparameterpoint values were associ- basedon high,medium,or no threat.

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