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Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas

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 Articles

Marine Ecoregions of the World:

A Bioregionalization of Coastal
and Shelf Areas
MARK D. SPALDING, HELEN E. FOX, GERALD R. ALLEN, NICK DAVIDSON, ZACH A. FERDAÑA, MAX FINLAYSON,
BENJAMIN S. HALPERN, MIGUEL A. JORGE, AL LOMBANA, SARA A. LOURIE, KIRSTEN D. MARTIN, EDMUND
M C MANUS, JENNIFER MOLNAR, CHERI A. RECCHIA, AND JAMES ROBERTSON

The conservation and sustainable use of marine resources is a highlighted goal on a growing number of national and international policy agendas.
Unfortunately, efforts to assess progress, as well as to strategically plan and prioritize new marine conservation measures, have been hampered by the
lack of a detailed, comprehensive biogeographic system to classify the oceans. Here we report on a new global system for coastal and shelf areas: the
Marine Ecoregions of the World, or MEOW, a nested system of 12 realms, 62 provinces, and 232 ecoregions. This system provides considerably better
spatial resolution than earlier global systems, yet it preserves many common elements and can be cross-referenced to many regional biogeographic
classifications. The designation of terrestrial ecoregions has revolutionized priority setting and planning for terrestrial conservation; we anticipate
similar benefits from the use of a coherent and credible marine system.

Keywords: ecoregions, marine biogeography, mapping, marine protected areas, representative conservation

M apped classifications of patterns in biodiversity

have long been an important tool in fields from
evolutionary studies to conservation planning (Forbes 1856,
In the absence of compelling global coverage, numerous
regional classifications have been created to meet regional
planning needs. This, of course, does not satisfy the need for
Wallace 1876, Spellerberg and Sawyer 1999, Lourie and a global system that is consistent across the many marine
Vincent 2004). The use of such systems (notably, the widely realms and coastal zones.
cited system developed by Olson et al. [2001]) in broadscale Biogeographic classifications are essential for developing
conservation, however, has largely been restricted to terres- ecologically representative systems of protected areas, as re-
trial studies (Chape et al. 2003, Hazen and Anthamatten quired by international agreements such as the Convention
2004, Hoekstra et al. 2005, Burgess et al. 2006, Lamoreux et on Biological Diversity’s Programme of Work on Protected
al. 2006). In the marine environment, existing global classi- Areas and the Ramsar Convention on Wetlands. Marine
fication systems remain limited in their spatial resolution. space is still grossly underrepresented in the global protected
Some are inconsistent in their spatial coverage or method- areas network (only about 0.5% of the surface area of the
ological approach. The few publications that have attempted oceans is currently protected; Chape et al. 2005), a fact that
to use biogeographic regionalization in global marine adds urgency to the need for tools to support the scaling up
conservation planning (e.g., Kelleher et al. 1995, Olson and of effective, representative marine conservation. The key idea
Dinerstein 2002) have been qualitative, and have expressed underlying the term “representative” is the intent to protect
concern about the lack of an adequate global classification. a full range of biodiversity worldwide—genes, species, and

Mark D. Spalding (e-mail: [email protected]), Zach A. Ferdaña, Jennifer Molnar, and James Robertson are conservation scientists in The Nature Conservancy’s
Conservation Strategies Group, Arlington, VA 22203. Helen E. Fox and Al Lombana are marine biologists in the Conservation Science Program, World Wildlife Fund–US,
Washington, DC 20037. Gerald R. Allen is a research associate at the Western Australian Museum, Perth, Western Australia 6986, Australia. Nick Davidson is the deputy
secretary general of the Ramsar Convention Secretariat, CH-1196 Gland, Switzerland. Max Finlayson is a member and former chair of Ramsar’s Scientific and
Technical Review Panel and principal researcher in wetland ecology at the International Water Management Institute, Colombo, Sri Lanka. Benjamin S. Halpern is
project coordinator for ecosystem-based management of coastal marine systems at the National Center for Ecological Analysis and Synthesis, Santa Barbara, CA 93101.
Miguel A. Jorge is deputy director of WWF International’s Global Marine Programme, CH-1196 Gland, Switzerland. Sara A. Lourie is a research associate at the
Redpath Museum, McGill University, Montreal, Quebec H3A 2K6, Canada. Kirsten D. Martin was a marine program officer with IUCN (World Conservation Union)
when this article was prepared and is currently working as a freelance consultant for the Census of Marine Life Initiative, 1205 Geneva, Switzerland. Edmund McManus
is a senior program officer in the UNEP (United Nations Environment Programme) World Conservation Monitoring Centre, Cambridge CB3 0DL, United
Kingdom. Cheri A. Recchia is marine program director at the Wildlife Conservation Society, New York, NY 10461. © 2007 American Institute of Biological Sciences.

www.biosciencemag.org July/August 2007 / Vol. 57 No. 7 • BioScience 573

Articles

higher taxa, along with the communities, evolutionary Sherman et al. 2005). Unlike the systems of Briggs and
patterns, and ecological processes that sustain this diversity. Longhurst, LMEs represent an expert-derived system with-
Biogeographic classifications provide a crucial foundation for out a rigorous, replicable core definition. LMEs are “rela-
the assessment of representativeness (Olson and Dinerstein tively large regions on the order of 200,000 km2 or greater,
2002, Lourie and Vincent 2004). characterized by distinct: (1) bathymetry, (2) hydrography, (3)
The growing commitment by governments and the United productivity, and (4) trophically dependent populations”
Nations (UN; e.g., the UN Law of the Sea, the UN Fish Stocks (www.lme.noaa.gov/Portal/). LMEs are largely conceived as
Agreement) to implement comprehensive arrangements units for the practical application of transboundary man-
for ocean governance provides an additional arena in which agement issues (fish and fisheries, pollution, habitat restora-
marine biogeographic classifications are needed. Biogeo- tion, productivity, socioeconomics, and governance). The
graphic regions are natural frameworks for marine zoning, LME system focuses on productivity and oceanographic
which is a tool increasingly used by regional fisheries man- processes, and in its present form omits substantial areas of
agement organizations. islands in the Pacific and the Indian oceans.
In this article, we present a new biogeographic classifica- These and other global systems continue to play an im-
tion for the world’s coastal and shelf areas, which draws heav- portant role in developing our understanding of marine bio-
ily on the existing global and regional literature. We believe geography and in practical issues of natural resource
that this classification will be of critical importance in sup- management. However, improvements are clearly possible and
porting analyses of patterns in marine biodiversity, in un- desirable. An ideal system would be hierarchical and nested,
derstanding processes, and, perhaps most important, in and would allow for multiscale analyses. Each level of the
directing future efforts in marine resource management and hierarchy would be relevant for conservation planning or
conservation. management interventions, from the global to the local, al-
though it is beyond the scope of the present effort to classify
Approaches for defining boundaries individual habitats or smaller features, such as individual es-
Observations of global biogeographic patterns in the marine
tuaries or seagrass meadows.
environment include early works by Forbes (1856), Ekman
We focus here on coastal and shelf waters, combining ben-
(1953, first published in German in 1935), and Hedgpeth
thic and shelf pelagic (neritic) biotas. These waters represent
(1957a), and more recent publications by Briggs (1974, 1995),
the areas in which most marine biodiversity is confined,
Hayden and colleagues (1984), Bailey (1998), and Longhurst
where human interest and attention are greatest, and where
(1998). These authors used a variety of definitions and cri-
there is often a complex synergy of threats far greater than in
teria for drawing biogeographic divisions. For example, Briggs
offshore waters (UNEP 2006). From a biodiversity perspec-
(1974, 1995) focused on a system of coastal and shelf provinces
tive, it is not simply that coastal and shelf waters have greater
defined by their degree of endemism (> 10%). This strong tax-
species numbers and higher productivity, but also that they
onomic focus and clear definition have led to relatively wide-
spread adoption of Briggs’s system, including its use by are biogeographically distinct from the adjacent high seas and
Hayden and colleagues (1984), with minor amendments, as deep benthic environments (Ekman 1953, Hedgpeth 1957a,
a part of their “classification of the coastal and marine envi- Briggs 1974).
ronments.” Adey and Steneck (2001) provided independent Our intention was to develop a hierarchical system based
verification of many of Briggs’s subdivisions in a study that on taxonomic configurations, influenced by evolutionary
modeled “thermogeographic”regions of evolutionary stability. history, patterns of dispersal, and isolation. We drew up ini-
Another important systematic approach, aimed mainly at tial guidelines on definitions and nomenclature to guide the
pelagic systems, is the two-tier system devised by Longhurst first data-gathering phase, then reviewed and refined them
(1998), which focuses on biomes and biogeochemical iteratively on the basis of the available data.
provinces. These subdivisions were based on a detailed array We reviewed over 230 works in journals, NGO (non-
of oceanographic factors, tested and modified using a large governmental organization) reports, government publica-
global database of chlorophyll profiles. The results represent tions, and other sources. For each of these, we looked at the
one of the most comprehensive partitionings of the pelagic underlying data and at the process of identification and de-
biota, but the scheme is of limited utility in the complex sys- finition of biogeographic units; we also considered the ob-
tems of coastal waters, a fact acknowledged by the author, who jectives of the classifications. To facilitate comparisons, we used
has recommended combining his open ocean system with oth- digital mapped versions of many of the existing biogeo-
ers for coastal and shelf waters (Watson et al. 2003; Alan R. graphic units. More than 40 independent experts provided fur-
Longhurst, Galerie l’Academie, Cajarc, France, personal com- ther advice (see the acknowledgments section). We refined a
munication, 2 November 2004). draft classification scheme through an assessment and review
The system of large marine ecosystems (LMEs) was de- process that involved a three-day workshop. In arriving at our
veloped over many years by a number of regional experts, with classification scheme, we adhered to three principles for our
considerable input from fisheries scientist Ken Sherman (e.g., classification: that it should have a strong biogeographic ba-
Sherman and Alexander 1989, Hempel and Sherman 2003, sis, offer practical utility, and be characterized by parsimony.

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 Articles

A strong biogeographic basis. All spatial units were defined evolutionary history. Realms have high levels of
on a broadly comparable biogeographic basis. Existing sys- endemism, including unique taxa at generic and family
tems rely on a broad array of source information—range levels in some groups. Driving factors behind the devel-
discontinuities, dominant habitats, geomorphological fea- opment of such unique biotas include water tempera-
tures, currents, and temperatures, for example—to identify ture, historical and broadscale isolation, and the prox-
areas and boundaries. In many cases these divergent ap- imity of the benthos.
proaches are compatible, given the close links between bio-
diversity and the underlying abiotic drivers (see the This article, with its focus on coastal and shelf areas, does
comparisons below). We preferred to be informed by com- not consider realms in pelagic or deep benthic environments.
posite studies that combined multiple divergent taxa or mul- This is an area requiring further analysis and development.
tiple oceanographic drivers in the derivation of boundaries,
as these were more likely to capture robust or recurring pat- Provinces. Nested within the realms are provinces:
terns in overall biodiversity.
A number of systems we reviewed were broadly biogeo- Large areas defined by the presence of distinct biotas
graphic, but with some adjustments to fit political boundaries. that have at least some cohesion over evolutionary time
Where it was possible to discern the biogeographic elements frames. Provinces will hold some level of endemism,
from the political, these systems were still used to inform the principally at the level of species. Although historical
process. isolation will play a role, many of these distinct biotas
have arisen as a result of distinctive abiotic features
Practical utility. We sought to develop a nested system, op- that circumscribe their boundaries. These may include
erating globally at broadly consistent spatial scales and in- geomorphological features (isolated island and shelf
corporating the full spectrum of habitats found across shelves. systems, semienclosed seas); hydrographic features
We thus avoided very fine-resolution systems that separated (currents, upwellings, ice dynamics); or geochemical
coastal and shelf waters into constituent habitats. We chose
influences (broadest-scale elements of nutrient supply
not to try to define minimum or maximum spatial areas for
and salinity).
our bioregions, but in some cases we did seek out systems that
subdivided very large spatial units (such as Briggs’s Indo-
In ecological terms, provinces are cohesive units likely, for
Polynesian Province, which covers more than 20% of the
example, to encompass the broader life history of many con-
world’s shallow shelf areas) or that amalgamated fine-scale
stituent taxa, including mobile and dispersive species. In
units such as single large estuaries or sounds.
many areas, the scale at which provinces may be conceived is
similar to that of the detailed spatial units used in global sys-
Parsimony. There are a number of respected and widely uti-
tems such as Briggs’s provinces, Longhurst’s biogeochemical
lized global and regional systems, and lack of agreement be-
tween such systems can be problematic. In developing a new provinces, and LMEs.
system, we sought to minimize further divergence from ex-
isting systems, yet still to obtain a truly global classification Ecoregions. Ecoregions are the smallest-scale units in the
system. We did this by adopting a nested hierarchy that (a) uti- Marine Ecoregions of the World (MEOW) system and are
lized systems that are already widely adopted (e.g., the Nature defined as follows:
Conservancy’s system in much of the Americas and the In-
terim Marine and Coastal Regionalisation for Australia) and Areas of relatively homogeneous species composition,
(b) fitted closely within broader-scale systems or alongside clearly distinct from adjacent systems. The species com-
other regional systems. position is likely to be determined by the predominance
of a small number of ecosystems and/or a distinct suite
Definitions of oceanographic or topographic features. The domi-
After the review process, we arrived at a set of critical work- nant biogeographic forcing agents defining the eco-
ing definitions. regions vary from location to location but may include
isolation, upwelling, nutrient inputs, freshwater influx,
Realms. The system’s largest spatial units are based on the ter- temperature regimes, ice regimes, exposure, sediments,
restrial concept of realms, described by Udvardy (1975) as currents, and bathymetric or coastal complexity.
“continent or subcontinent-sized areas with unifying fea-
tures of geography and fauna/flora/vegetation.” From our In ecological terms, these are strongly cohesive units, suf-
marine perspective, realms are defined as follows: ficiently large to encompass ecological or life history processes
for most sedentary species. Although some marine ecoregions
Very large regions of coastal, benthic, or pelagic ocean may have important levels of endemism, this is not a key
across which biotas are internally coherent at higher determinant in ecoregion identification, as it has been in ter-
taxonomic levels, as a result of a shared and unique restrial ecoregions.

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Articles

We suggest that the most appropriate outer boundary for with a stronger biogeographic basis than the current LME de-
these coastal and shelf realms, provinces, and ecoregions is the lineations. Both the Briggs and Hayden systems and the
200-meter (m) isobath, which is a widely used proxy for the LMEs show considerable variation in the size of their spatial
shelf edge and often corresponds to a dramatic ecotone units; the Briggs approach of using 10% endemism distin-
(Forbes 1856, Hedgpeth 1957b, Briggs 1974). Such a sharp guishes many isolated communities around oceanic islands,
boundary can only be indicative: Shelf breaks are not always but fails to disaggregate vast areas with gradual faunal changes,
clear; the bathymetric location of an “equivalent” biotic tran- even where the incremental effects of such changes are very
sition is highly variable; and there is considerable overlap large indeed (e.g., the Indo-Pacific). The large spatial units in
and influence between shelf, slope, and adjacent pelagic bio- all of these systems clearly encompass significant levels of in-
tas. At the same time, most of the classifications that we re- ternal biogeographic heterogeneity, which we were keen to dis-
viewed have been heavily influenced by data from nearshore aggregate through a more detailed system of ecoregions.
and intertidal biotas, and data from deeper water typically had
We found regional systems for almost all coastal and shelf
decreasing influence on boundary definitions. We believe
waters, although many are described only in the gray litera-
that beyond 200 m, other biogeographic patterns will in-
ture. Notable exceptions were the Russian Arctic and the
creasingly predominate, altering or hiding the patterns rep-
continental coasts of much of South, Southeast, and East
resented by the system proposed here.
Asia. For these areas, we relied heavily on global data sets and
A global, nested system unpublished expert opinion, using more focused biogeo-
We propose a nested system of 12 realms, 62 provinces, and graphic publications (where available) for refining individ-
232 ecoregions covering all coastal and shelf waters of the ual boundaries.
world. Figure 1 depicts the review process, showing four biogeo-
As the MEOW system is based on existing classifications, graphic schemes: Briggs’s system of provinces (1974, 1995);
variation and mismatch among systems led to challenges an expert-derived system combining biotic and abiotic fea-
and compromises. The global coastal classifications of Briggs tures for South America (Sullivan Sealey and Bustamante
and Hayden, for example, do not show great congruence 1999); the current LMEs; and a regional classification based
with the LMEs. The Briggs and related Hayden systems on a single taxonomic grouping (decapod crustaceans; Boschi
appeared to be more closely allied to our need for a system 2000). Despite their different origins, these systems show a re-

Figure 1. Reconciliation of differing boundary systems for South America. The map on the left illustrates four
biogeographic systems: (A) Briggs’s provinces, (B) Sullivan Sealey and Bustamante’s provinces, (C) large
marine ecosystems, and (D) Boschi’s provinces. System similarities are exemplified in three inset maps:
northern Peru (inset 1), Cabo Frio (inset 2), and Chiloé Island (inset 3). The map on the right shows the
Marine Ecoregions of the World provinces (labeled) and their ecoregion subdivision boundaries.

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 Articles

markable congruence at a number of key biogeographic [1998], Fernandez et al. [2000], Ojeda et al. [2000], and
boundaries. Camus [2001] for data concerning the Chilean coast).
Thus, it was possible to adopt a single system as a pri- Although the boundaries in other regions were not as
mary source, and the MEOW provinces (figure 1, right) were simple to resolve as those along the South American coast,
based almost entirely on Sullivan Sealey and Bustamante we applied the same approaches. The section that follows
(1999), while remaining well aligned with the other systems. gives some information on the key sources used in drawing
At a finer resolution, the ecoregions for South America are de- boundaries.
rived almost entirely from the same publication (Sullivan
Sealey and Bustamante 1999), this being the only compre- Marine Ecoregions of the World
hensive system for these coasts. Even at this scale, however, Box 1 and figures 2 and 3 give a summary of the entire
efforts were made to locate independent verification of MEOW system, which covers all coastal and shelf waters
boundaries, and it is reassuring to note that these more de- shallower than 200 m. The shaded area of each map (figures
tailed subdivisions were often supported by data from other 2, 3) extends 370 kilometers (200 nautical miles) offshore
oceanographic and ecological literature (see, e.g., Strub et al. (or to the 200-m isobath, where this lies further offshore),

Figure 2. Final biogeographic framework: Realms and provinces. (a) Biogeographic realms with ecoregion
boundaries outlined. (b) Provinces with ecoregions outlined. Provinces are numbered and listed in box 1.

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Box 1. Marine Ecoregions of the World.

Numbers for the provinces and ecoregions match those shown on the maps in figures 2b and 3. Realms are indicated in boldface, provinces (1–62) in
italics, and ecoregions (1–232) in roman type.
Arctic 54. Gulf of Alaska 23. Bay of Bengal
1. Arctic (no provinces identified) 55. North American Pacific Fijordland 107. Eastern India
1. North Greenland 56. Puget Trough/Georgia Basin 108. Northern Bay of Bengal
2. North and East Iceland 57. Oregon, Washington, Vancouver 24. Andaman
3. East Greenland Shelf Coast and Shelf 109. Andaman and Nicobar Islands
4. West Greenland Shelf 58. Northern California 110. Andaman Sea Coral Coast
5. Northern Grand Banks–Southern 11. Warm Temperate Northeast Pacific 111. Western Sumatra
Labrador 59. Southern California Bight Central Indo-Pacific
6. Northern Labrador 60. Cortezian
7. Baffin Bay–Davis Strait 61. Magdalena Transition 25. South China Sea
8. Hudson Complex 112. Gulf of Tonkin
Tropical Atlantic 113. Southern China
9. Lancaster Sound
10. High Arctic Archipelago 12. Tropical Northwestern Atlantic 114. South China Sea Oceanic Islands
11. Beaufort–Amundsen–Viscount 62. Bermuda 26. Sunda Shelf
Melville–Queen Maud 63. Bahamian 115. Gulf of Thailand
12. Beaufort Sea—continental coast 64. Eastern Caribbean 116. Southern Vietnam
and shelf 65. Greater Antilles 117. Sunda Shelf/Java Sea
13. Chukchi Sea 66. Southern Caribbean 118. Malacca Strait
14. Eastern Bering Sea 67. Southwestern Caribbean
68. Western Caribbean 27. Java Transitional
15. East Siberian Sea
69. Southern Gulf of Mexico 119. Southern Java
16. Laptev Sea
70. Floridian 120. Cocos-Keeling/Christmas Island
17. Kara Sea
18. North and East Barents Sea 13. North Brazil Shelf 28. South Kuroshio
19. White Sea 71. Guianan 121. South Kuroshio
Temperate Northern Atlantic 72. Amazonia 29. Tropical Northwestern Pacific
14. Tropical Southwestern Atlantic 122. Ogasawara Islands
2. Northern European Seas
73. Sao Pedro and Sao Paulo Islands 123. Mariana Islands
20. South and West Iceland
74. Fernando de Naronha and Atoll 124. East Caroline Islands
21. Faroe Plateau
das Rocas 125. West Caroline Islands
22. Southern Norway
23. Northern Norway and Finnmark 75. Northeastern Brazil 30. Western Coral Triangle
24. Baltic Sea 76. Eastern Brazil 126. Palawan/North Borneo
25. North Sea 77. Trindade and Martin Vaz Islands 127. Eastern Philippines
26. Celtic Seas 15. St. Helena and Ascension Islands 128. Sulawesi Sea/Makassar Strait
78. St. Helena and Ascension Islands 129. Halmahera
3. Lusitanian
130. Papua
27. South European Atlantic Shelf 16. West African Transition 131. Banda Sea
28. Saharan Upwelling 79. Cape Verde 132. Lesser Sunda
29. Azores Canaries Madeira 80. Sahelian Upwelling 133. Northeast Sulawesi
4. Mediterranean Sea 17. Gulf of Guinea 31. Eastern Coral Triangle
30. Adriatic Sea 81. Gulf of Guinea West 134. Bismarck Sea
31. Aegean Sea 82. Gulf of Guinea Upwelling 135. Solomon Archipelago
32. Levantine Sea 83. Gulf of Guinea Central 136. Solomon Sea
33. Tunisian Plateau/Gulf of Sidra 84. Gulf of Guinea Islands 137. Southeast Papua New Guinea
34. Ionian Sea 85. Gulf of Guinea South
35. Western Mediterranean 86. Angolan 32. Sahul Shelf
36. Alboran Sea 138. Gulf of Papua
Western Indo-Pacific 139. Arafura Sea
5. Cold Temperate Northwest Atlantic
18. Red Sea and Gulf of Aden 140. Arnhem Coast to Gulf of Carpenteria
37. Gulf of St. Lawrence–Eastern
87. Northern and Central Red Sea 141. Bonaparte Coast
Scotian Shelf
88. Southern Red Sea 33. Northeast Australian Shelf
38. Southern Grand Banks–South
89. Gulf of Aden 142. Torres Strait Northern Great
Newfoundland
39. Scotian Shelf 19. Somali/Arabian Barrier Reef
40. Gulf of Maine/Bay of Fundy 90. Arabian (Persian) Gulf 143. Central and Southern Great
41. Virginian 91. Gulf of Oman Barrier Reef
92. Western Arabian Sea 34. Northwest Australian Shelf
6. Warm Temperate Northwest Atlantic
93. Central Somali Coast 144. Exmouth to Broome
42. Carolinian
43. Northern Gulf of Mexico 20. Western Indian Ocean 145. Ningaloo
94. Northern Monsoon Current Coast 35. Tropical Southwestern Pacific
7. Black Sea
95. East African Coral Coast 146. Tonga Islands
44. Black Sea
96. Seychelles 147. Fiji Islands
Temperate Northern Pacific 97. Cargados Carajos/Tromelin Island 148. Vanuatu
8. Cold Temperate Northwest Pacific 98. Mascarene Islands 149. New Caledonia
45. Sea of Okhotsk 99. Southeast Madagascar 150. Coral Sea
46. Kamchatka Shelf and Coast 100. Western and Northern Madagascar
101. Bight of Sofala/Swamp Coast 36. Lord Howe and Norfolk Islands
47. Oyashio Current
102. Delagoa 151. Lord Howe and Norfolk Islands
48. Northeastern Honshu
49. Sea of Japan 21. West and South Indian Shelf Eastern Indo-Pacific
50. Yellow Sea 103. Western India 37. Hawaii
9. Warm Temperate Northwest Pacific 104. South India and Sri Lanka 152. Hawaii
51. Central Kuroshio Current 22. Central Indian Ocean Islands 38. Marshall, Gilbert, and Ellis Islands
52. East China Sea 105. Maldives 153. Marshall Islands
10. Cold Temperate Northeast Pacific 106. Chagos 154. Gilbert/Ellis Island
53. Aleutian Islands

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Box 1. (continued)

Numbers for the provinces and ecoregions match those shown on the maps in figures 2b and 3. Realms are indicated in boldface, provinces (1–62) in
italics, and ecoregions (1–232) in roman type.
39. Central Polynesia 47. Warm Temperate Southwestern Atlantic 56. Southeast Australian Shelf
155. Line Islands 180. Southeastern Brazil 204. Cape Howe
156. Phoenix/Tokelau/Northern 181. Rio Grande 205. Bassian
Cook Islands 182. Rio de la Plata 206. Western Bassian
157. Samoa Islands 183. Uruguay–Buenos Aires Shelf 57. Southwest Australian Shelf
40. Southeast Polynesia 48. Magellanic 207. South Australian Gulfs
158. Tuamotus 184. North Patagonian Gulfs 208. Great Australian Bight
159. Rapa-Pitcairn 185. Patagonian Shelf 209. Leeuwin
160. Southern Cook/Austral Islands 186. Malvinas/Falklands 58. West Central Australian Shelf
161. Society Islands 187. Channels and Fjords of 210. Shark Bay
41. Marquesas Southern Chile 211. Houtman
162. Marquesas 188. Chiloense
Southern Ocean
42. Easter Island 49. Tristan Gough
189. Tristan Gough 59. Subantarctic Islands
163. Easter Island 212. Macquarie Island
Tropical Eastern Pacific Temperate Southern Africa 213. Heard and Macdonald Islands
43. Tropical East Pacific 50. Benguela 214. Kerguelen Islands
164. Revillagigedos 190. Namib 215. Crozet Islands
165. Clipperton 191. Namaqua 216. Prince Edward Islands
166. Mexican Tropical Pacific 51. Agulhas 217. Bouvet Island
167. Chiapas–Nicaragua 192. Agulhas Bank 218. Peter the First Island
168. Nicoya 193. Natal 60. Scotia Sea
169. Cocos Islands 52. Amsterdam–St Paul 219. South Sandwich Islands
170. Panama Bight 194. Amsterdam–St Paul 220. South Georgia
171. Guayaquil 221. South Orkney Islands
Temperate Australasia 222. South Shetland Islands
44. Galapagos
172. Northern Galapagos Islands 53. Northern New Zealand 223. Antarctic Peninsula
173. Eastern Galapagos Islands 195. Kermadec Island 61. Continental High Antarctic
174. Western Galapagos Islands 196. Northeastern New Zealand 224. East Antarctic Wilkes Land
197. Three Kings–North Cape 225. East Antarctic Enderby Land
Temperate South America
54. Southern New Zealand 226. East Antarctic Dronning Maud Land
45. Warm Temperate Southeastern Pacific 198. Chatham Island 227. Weddell Sea
175. Central Peru 199. Central New Zealand 228. Amundsen/Bellingshausen Sea
176. Humboldtian 200. South New Zealand 229. Ross Sea
177. Central Chile 201. Snares Island
178. Araucanian 62. Subantarctic New Zealand
55. East Central Australian Shelf 230. Bounty and Antipodes Islands
46. Juan Fernández and Desventuradas 202. Tweed-Moreton 231. Campbell Island
179. Juan Fernández and Desventuradas 203. Manning-Hawkesbury 232. Auckland Island

but, as already noted, we consider the principal focus of this process (e.g., unpublished reports by Jerry M. Kemp in
classification to be the benthos above 200 m and the overlying 2005 for the Middle Eastern seas and by S. A. L. in 2006
water column. for the Andaman to Java coasts); the system for the
Key sources included the following: Indo-Pacific oceanic islands was developed by one of us
(G. R. A.) on the basis of many years of field experience,
• Biogeographic assessments in the peer-reviewed expert review, and networking with other scientists
literature, including the global studies already across the region
mentioned and many regional publications (e.g.,
Bustamante and Branch [1996] and Turpie et al. [2000] These schemes were assessed alongside other biogeographic
for temperate southern Africa, Linse et al. [2006] for the literature, and in some cases alterations were made to better
Southern Ocean) represent the arguments of biogeography, utility, and parsi-
mony outlined above. A full listing of the sources referenced
• Ecoregional assessments conducted by NGOs (e.g., can be found at www.nature.org/MEOW or www.worldwildlife.
Sullivan Sealey and Bustamante [1999] for Latin org/MEOW.
America, WWF [2004 and unpublished reports] The proposed realms adopt the broad latitudinal divi-
for much of Africa, Green and Mous [2006] for sions of polar, temperate, and tropical, with subdivisions
the Coral Triangle provinces) based on ocean basin (broadly following the oceanic biomes
of Longhurst [1998]). In the temperate waters of the South-
• Government-derived or supported systems (e.g., ern Hemisphere, we diverge from this approach. We consider
Thackway and Cresswell [1998] for Australia, the differences across the oceans too substantial, and the
Powles et al. [2004] for Canada) connections around the continental margins too great, to
support either ocean basin subdivisions or a single circum-
• Input from several of the authors of this article and global realm (equivalent to Longhurst’s Antarctic Westerly
assessments commissioned explicitly for the MEOW Winds Biome), and hence we have adopted continental

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580 BioScience • July/August 2007 / Vol. 57 No. 7

Figure 3. Final biogeographic framework, showing
ecoregions. Ecoregions are numbered and listed in box 1.

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margin realms for temperate Australasia, southern Africa, parties to the Ramsar Convention have committed to achieve
and South America. The paucity of existing literature dis- a “coherent and comprehensive national and international net-
cussing these broadest-scale biogeographic units from a work” (Ramsar Convention 1999), although until now it has
global perspective presents a stark contrast to the terrestrial not been possible to assess the biogeographic coverage of
biogeographic literature. marine and coastal Ramsar sites at the global level. The results
The level of internal heterogeneity of biotas within differ- of this overlay are presented in table 1.
ent realms is quite varied. For some realms, the differences in One value of biogeographic classifications is their use in un-
biota at the provincial level are substantial, including the covering inequities and dramatic gaps in conservation cov-
warm temperate faunas on either side of the Temperate South erage. Although a more thorough analysis would be required
America realm and the tropical faunas on either side of the to determine more clearly the degree of representation pro-
Tropical Atlantic realm. By contrast, we have subdivided the vided by the existing selection of Ramsar sites, some basic ob-
widely used Indo-Pacific “realm” into three units. This is the servations are immediately apparent. The Ramsar network is
region of greatest diversity, and it covers a vast area. Across this extensive, but it is dominated by sites in the temperate North
region are clinal changes in taxa that lack clear breaks, but are Atlantic and shows a striking paucity of sites in, for example,
sufficiently large that faunas at either end bear little resem- the eastern Indo-Pacific and the Southern Ocean. At finer hi-
blance to each other. Our Indo-Pacific subdivisions (which erarchical resolution, further gaps can be identified: While 92%
it might be appropriate to consider as subrealms) follow less of realms are represented, this translates to only 73% of
clearly defined biogeographic boundaries than other realms, provinces and 52% of ecoregions, leaving some 112 ecoregions
but these divisions produce spatial units that are more with no Ramsar representation. These gaps are widespread,
comparable to other realms in overall biodiversity, levels of including four ecoregions in the temperate North Atlantic.
endemism, and spatial area.
At broader scales, we undertook a simple spatial analysis Conclusions
to explore the links or possible crossovers between the MEOW The MEOW classification provides a critical tool for marine
system, LMEs, and Briggs’s provinces. The incomplete cov- conservation planning. It will enable gap analyses and
erage of the LME system is clearly limiting for global con- assessments of representativeness in a global framework. It
servation planning: 78 of our 232 ecoregions include a provides a level of detail that will support linkage to practi-
substantive area (greater than 10% of their total area) that is cal conservation interventions at the field level. For example,
not covered by any LME. Of the remainder, some 49% of two major international conservation organizations (the
LMEs show good congruence (> 90% of shelf area) with ei- Nature Conservancy and WWF) use ecoregions as planning
ther single ecoregions or ecoregion combinations. (The units. From a global standpoint, the MEOW system offers sim-
boundary of the Arctic LME has not been mapped, and so was ilar opportunities for the marine environment. It also provides
ignored in these calculations.) In comparison, 30 of Briggs’s a rational framework in which to analyze patterns and
53 provinces (57%) show good congruence (> 90% of shelf processes in coastal and shelf biodiversity.
area) with single ecoregions or ecoregion combinations. This The global and hierarchical nature of the MEOW can
figure rises to 39 (74%) if we include congruence at 85% of support analytical approaches that move between scales.
the shelf area. Using MEOW, global information can also be used to target
We also used the MEOW system to look at the coverage of action on the ground, while field-level information can be
the marine and coastal network of Ramsar sites. Contracting placed alongside information on adjacent or remote locations,

Table 1. The geographic spread of marine and coastal Ramsar sites within the Marine Ecoregions of the World
classification.
Ecoregions Provinces
Total Number with Percentage Number with Percentage
Ramsar Ramsar Total with Ramsar Ramsar Total with Ramsar
Realm sites sites number sites sites number sites

Arctic 26 10 19 53 1 1 100
Temperate Northern Atlantic 374 21 25 84 6 6 100
Temperate Northern Pacific 38 12 17 71 4 4 100
Tropical Atlantic 117 17 25 68 4 6 67
Western Indo-Pacific 41 14 25 56 7 7 100
Central Indo-Pacific 35 16 40 40 10 12 83
Eastern Indo-Pacific 1 1 12 8 1 6 17
Tropical Eastern Pacific 29 8 11 73 2 2 100
Temperate South America 14 9 15 60 3 5 60
Temperate Southern Africa 9 3 5 60 2 3 67
Temperate Australasia 25 9 17 53 5 6 83
Southern Ocean 0 0 21 0 0 4 0
Total 709 120 232 52 45 62 73

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Articles

providing a wider spatial perspective. Rooted in existing re- and shelf areas, and the links between this system and other
gional systems, the base units of the MEOW already under- global and regional systems make it possible to adopt and use
pin conservation efforts at regional levels, and a strong body it with minimal disruption to existing data sets or analytical
of marine ecoregional planning literature illustrates how approaches. The unique collaboration of conservation orga-
global or regional concerns can be converted into field-based nizations in developing this system adds further value, and may
conservation action (Banks et al. 2000, Beck and Odaya 2001, reduce the duplication of effort that so often undermines
Larsen et al. 2001, Kramer and Kramer 2002, Ferdaña 2005). global conservation approaches (Mace et al. 2000). In short,
The value of the MEOW system extends beyond conser- the system proposed here is powerful and robust, and should
vation planning. Looking afresh at the broader-scale classes prove to be of great value in conservation planning and
and taking advantage of the improved resolution offered by broader biogeographic discussion. Two international con-
the MEOW system, it is possible to review wider issues of bio- servation agencies (the Nature Conservancy and WWF) have
diversity distribution and evolution. At the broadest scales, the already begun to use this system and expect to use it more
most important elements of biogeographic subdivision are the widely in the future. Similarly, members of the Scientific and
barriers that have separated substantial areas over evolu- Technical Review Panel of the Ramsar Convention who par-
tionary timescales (Adey and Steneck 2001). In the MEOW ticipated in developing this system are undertaking more
realms (noting the special case of the Indo-Pacific described detailed analyses to explore its utility to support the future
above), these barriers consist of landmasses, wide ocean identification and designation of coastal and marine Wetlands
basins, and temperature gradients. of International Importance.
Although there is variation in degree, the provinces can be
seen as finer-scale units of evolutionary isolation. They align Acknowledgments
with many of the more important factors driving recent and The Marine Ecosystems of the World system draws heavily on
contemporary evolutionary processes. Temperature, or lati- the work of others, including the hundreds of contributors
tude, continues to play an important role (separating warm to the publications, gray literature, and workshops that
and cold temperate provinces), but so does the further iso- created the many regional classifications. In addition, we
lation provided by deep water, narrow straits, or rapid changes would especially like to thank the following people, who have
in shelf conditions. Elsewhere, the connectivity provided by provided advice or commentary: Asa Andersson, Jeff Ardron,
ocean currents, such as the Antarctic Coastal Current and the Allison Arnold, Paul Barber, Mike Beck, Carlo Nike Bianchi,
Canaries Current, can be seen in the classifications, and the
John Bolton, George Branch, John Briggs, Georgina Busta-
importance of biological stepping-stones through various
mante, Rodrigo Bustamante, Jose Farina, Sergio Floeter,
island chains is clearly illustrated. Finally, the ecoregions,
Angus Gascoigne, Serge Gofas, Charlie Griffiths, Huw
which distinguish the MEOW system, reflect unique ecolog-
Griffiths, Randy Hagenstein, Jon Hoekstra, David John,
ical patterns that extend beyond the broad drivers of evo-
Peter Kareiva, Ken Kassem, Jerry Kemp, Phil Kramer, Katrin
lutionary processes.
Linse, Gilly Llewellyn, Stephan Lutter, Kasim Moosa, Alexis
Of course, as Wallace (1876) noted,“nothing like a perfect
Morgan, Dag Nagoda, Sergio Navarete, Kate Newman, (Bina)
zoological division of the earth is possible. The causes that have
Maya Paul, Sian Pullen, Callum Roberts, Rod Salm, Andrew
led to the present distribution of animal life are so varied, their
Smith, Jennifer Smith,Vassily Spiridonov,Victor Springer, Juan
action and reaction have been so complex, that anomalies and
Luis Suárez de Vivero, Marco Taviani, Charlie Veron, Eleni
irregularities are sure to exist which will mar the symmetry
of any rigid system” (p. 53). Consequently, the use of bio- Voultsiadou, Mohideen Wafar, Carden Wallace, Kathy Walls,
geographic data in a global classification is inevitably a process and David Woodland.
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