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Journal of Biogeography (J. Biogeogr.

) (2012) 39, 12–30


A realignment of marine biogeographic
provinces with particular reference to fish
John C. Briggs1* and Brian W. Bowen2


Department of Fisheries and Wildlife, Oregon
State University, Corvallis, OR 97331, USA,
Hawaii Institute of Marine Biology,
University of Hawaii, Kaneohe, HI 96744,


*Correspondence and present address: John C.
Briggs, 43939 Spiaggia Place, Indio, CA 92203,

Endemism, evolution, fishes, marine biogeography, phylogeography, provinces,
regions, speciation, zoogeography.

Marine provinces, founded on contrasting floras or faunas, have been recognized
for more than 150 years but were not consistently defined by endemism until
1974. At that time, provinces were based on at least a 10% endemism and nested
within biogeographic regions that covered large geographic areas with contrasting
biotic characteristics. Over time, some minor adjustments were made but the
overall arrangement remained essentially unaltered. In many provinces, data on
endemism were still not available, or were available only for the most widely
studied vertebrates (fishes), a problem that is ongoing. In this report we propose a
realignment for three reasons. First, recent works have provided new information
to modify or redefine the various divisions and to describe new ones, including
the Mid-Atlantic Ridge, Southern Ocean, Tropical East Pacific and Northeast
Pacific. Second, phylogeographic studies have demonstrated genetic subdivisions
within and between species that generally corroborated provinces based on taxonomic partitions, with a notable exception at the Indian–Pacific oceanic
boundary. Third, the original separation of the warm-temperate provinces from
the adjoining tropical ones has distracted from their close phylogenetic relationships. Here we propose uniting warm-temperate and tropical regions into a
single warm region within each ocean basin, while still recognizing provinces
within the warm-temperate and tropical zones. These biogeographic subdivisions
are based primarily on fish distribution but utilize other marine groups for
comparison. They are intended to demonstrate the evolutionary relationships of
the living marine biota, and to serve as a framework for the establishment of
smaller ecological units in a conservation context.

Biogeographic patterns are most useful when they identify
those parts of the world that host the more unique biotas, that
is, areas of evolutionary innovation or refuges where an older
biota persists. Edward Forbes, in his posthumous work The
Natural History of European Seas (Forbes, 1859), made three
observations of lasting value: (1) each zoogeographic province
is an area where there was a special manifestation of creative
power, and the animals originally formed there are apt to
become mixed with emigrants from other provinces; (2) each
species was created only once, and individuals tend to migrate
outwards from their centre of origin; and (3) provinces must,
to be understood, be traced back like species to their origins in

past time. Sven Ekman undertook the huge task of analysing all
the pertinent literature on marine animal distribution, leading
to the publication of his book Tiergeographie des Meeres
(Ekman, 1935). This was followed by a revised English edition,
Zoogeography of the Sea (Ekman, 1953).
Ekman (1953) considered the marine world to be composed
of a series of large regions or subregions. For the continental
shelf, he described regions located in warm, temperate, and
polar waters; their separation by zoogeographic barriers; and
their endemism. Later, Briggs (1974) divided the continental
shelf into a series of large biogeographic regions that, in turn,
contained smaller provinces. Provinces were defined on the
basis of endemism, and it was observed that the greater the
proportion of endemic biota, the greater the evolutionary
ª 2011 Blackwell Publishing Ltd

Hawaiian. Aral. Within each zone. In recent years. whereas few extend into the cold-temperate regions (Briggs. 2006a. MARINE BIOGEOGRAPHY With regard to the continental shelves. Easter Island. Auckland. plants and even microbes) receives greater attention.Marine biogeographic provinces significance. there is a high concordance between levels of endemism in fishes. An objective standard was considered to be necessary in order that provinces could be recognized within the larger regions. the 10% criterion has the advantages of stability and functionality: areas that possess > 10% endemism have proved to be locations of unusual evolutionary interest. We have endeavoured to bring in other taxa where information is available and find that. Black Sea. Caspian. At the same time. a separation into different regions eventually proved to be inappropriate. the fish fauna of Hawaii is 25% endemic (Randall. 3. a value of 10% was chosen for an area to qualify as a distinct province (Briggs. Eastern Atlantic Region Provinces: Lusitania. 2010). St Helena. Red Sea. warm-temperate. 5. The 10% criterion is also conservative. Tristan–Gough. Caribbean. while other advances have improved our concept of how speciation and dispersal operate in the marine environment. 2005a). Journal of Biogeography 39. The provinces described herein are in coastal and shallow habitats. Indo-Polynesian. fishes that are very unlikely candidates for endemism on the scale of provinces.. Fishes in shallow areas. 2. a realignment is proposed here in which expanded regions (Fig. Therefore. For example. after an examination of endemism rates in numerous areas. However. the four temperature zones of the world’s oceans have usually been identified as tropical. provinces closely linked to those of Briggs (1995) were recently subdivided into ecoregions to address the appropriate scale for conservation efforts (Spalding et al. Western Atlantic Region Provinces: Carolina. this minimum value would admit most of the areas that were previously recognized as provinces on the basis of less formal criteria. the upsurge of phylogeography has produced many useful studies with biogeographic connotations. In the Caribbean Province. It is now possible to provide more accurate reconstructions of evolutionary relationships in several of the large oceanic regions. 2000) and corals 37% (Veron. Endemism rates in many areas and many taxa are still poorly known and are likely to change as the marine biota (especially invertebrates. 2007). Marquesas. Western Pacific Region Provinces (warm-temperate): Sino-Japanese. Considering that there is a very close relationship between each warm-temperate province and its adjacent tropical equivalent (Vermeij. cold-temperate and cold. usually defined as < 200 m depth (Randall. the red algae 25% (Abbott. if no change is required. Tropical Indo-West Pacific Region Provinces: Western Indian Ocean. provinces will remain as described in Briggs (1995). molluscs and other biotas. In order to keep abreast of continuing research.. the reef fishes are 33% endemic (Floeter et al. If new research has indicated that provinces need to be altered. 1980). Drew et al. Kermadec. Some of the research advances need to be reflected in the arrangement of regions and provinces. Many families and genera span the tropical and warm-temperate regions within each ocean basin. 1999). In the absence of compelling reasons to the contrary. So. and the molluscan fauna 20% (Kay. Eastern Pacific Region 13 . and provinces were located within the regions (Briggs. palaeontological research has produced discoveries about fossils. providing a fascinating foundation for further study. 2007). often are the only groups with sufficient information for biogeographic inference. Otherwise. and based largely on the distributions of fishes. but some of these disparities in endemism may be cases where one or two taxonomic groups are much better known than others. Benguela. 2007. because it is typically based on species lists that include oceanic wanderers such as tunas (Randall. this limitation also indicates the state of knowledge. earth movements and sealevel changes that are critical to historical biogeography. Various biotic areas had previously been called provinces but there was no agreement as to the qualifications necessary for provincial status. While both the geographic and taxonomic frameworks are admittedly aligned with the authors’ field of study. but discoveries made during the next 20 years required the changes published in Briggs (1995). Furthermore. 2008). we choose to retain the 10% criterion. while recognizing that this is not an absolute limit but a guidepost for recognizing unique biotic assemblages. where the fauna is relatively well known. COMPLETE OUTLINE OF SHELF REGIONS AND PROVINCES WARM REGIONS (tropical and warm-temperate waters) 1. a primary criticism of this arrangement was the placement of warm-temperate provinces in different regions from tropical ones. Brazilian.. as phylogeographic studies are revealing unrecognized endemic species (Bowen et al. Tropical Eastern Atlantic. Notably. The 10% criterion may often be an underestimate. decapod crustaceans 32% (Boschi. Argentinian. 1974). they are illustrated and references are provided. 12–30 ª 2011 Blackwell Publishing Ltd The regions and provinces that were defined in 1974 proved to be useful. Southeastern Australian. 1995. a series of biogeographic regions was recognized. which may have little evolutionary significance. 2000). Grant et al. The definition of provinces by 10% endemism has been generally accepted for the past 35 years. additional modifications are required. 1) will encompass provinces in both temperature zones. 1974). Amsterdam–St Paul. 2007).. especially to combine depauperate outposts of larger provinces. 2007). Southwestern Australian. Good arguments can be made for a higher criterion (15% or 20% endemism). Ascension. 2010).. Over time. 4. Exceptions to this concordance may become apparent.

and between cold and cold-temperate regions. Black Sea. Antarctic Region DISCUSSION Atlantic Warm Regions The reconstituted Eastern Atlantic Region (Fig. Western North Pacific Region Provinces: Oriental.. 2. Sub-Antarctic Region Provinces: South Georgia. Included within the two regions are 13 provinces. crustaceans and marine mammals show some population genetic separations. 12–30 ª 2011 Blackwell Publishing Ltd . earlier work was reviewed by Briggs (1974). The Lusitania Province includes the offshore islands of the Canaries. New Zealand–Australian Region Provinces: Tasmania. and cold-temperate regions are depicted in dark blue. and Benguela provinces in the East Atlantic (EA) and Mediterranean. Caspian. Azores and Madeira. C. 1). Warm-temperate provinces are depicted along the shore lines in medium blue. California and Peru-Chilean provinces in the East Pacific (EP). The Western Atlantic Region extends from Cape Hatteras and the northern Gulf of Mexico 14 southwards to the Valdes Peninsula on the South American east coast. no recent evaluations of endemism in these three provinces. the Tropical Eastern Atlantic (TEA) Province extends south to Mossamedes. The offshore islands of the Cape Verdes. including Carolina and Argentinian provinces in the West Atlantic (WA). A comprehensive treatise on tropical Atlantic biogeography and evolution has recently been published by Floeter et al. Eastern Atlantic Region 5. 2. Sa˜o Tome´ and Prı´ncipe are included. Oregon. Southern Argentina. Antipodes. to our knowledge. W. COLD-TEMPERATE AND POLAR NORTHERN HEMISPHERE 1. South American Region Provinces: Southern Chile. the warm-temperate Lusitania Province extends south to southern Morocco and eastwards through the Mediterranean (Fig. Auckland. Kurile. Panamanian. Peru–Chilean. where 28% of marine species are endemic (IUCN. Kerguelen. 1) now extends from the southern entrance to the English Channel southwards to the Cape of Good Hope. 2). Juan Ferna´ndez. Falkland Islands. Briggs and B. Sino-Japanese. (2008). Cortez. the demonstrated patterns and relationships have significance for many of the other phyla on the continental shelves. Farther to the east are the Black Sea.J. 3. From southern Morocco. 3. Kermadec. Bowen cold-temperate provinces warm-temperate provinces warm oceans 120° 60° 0° 60° 120° 60° 60° 30° 30° WA EP 0° 0° EA IWP 30° 30° 60° 60° 0 km 120° 60° 0° 60° 3500 7000 120° Figure 1 The realignment of marine biogeographic provinces eliminates the distinction between tropical and warm-temperate regions. Bouvet. The Straits of Gibraltar are often assumed to be a natural barrier between Mediterranean and Atlantic segments of the Lusitania Province. See text for precise geographic boundaries and additional warm-temperate provinces at oceanic islands. 2007). Angola (Fig. In the TEA there are about 388 Journal of Biogeography 39. Eastern Atlantic Region From its northern boundary at the southern British Isles. Caspian and Aral provinces. 4. Prince Edward. Southeastern Australian and Southwestern Australian provinces in the Indo-West Pacific (IWP). at Cap Juby. There are. New Zealand. it is the source of much of the new information utilized in this section. Western Atlantic Region 4. Arctic Region B. Macquarie. Tierra del Fuego. Cold regions (Arctic and Antarctic) are depicted in white. there is no consistent pattern of evolutionary partitions at the Straits (reviewed in Paternello et al. Provinces: California. Aral. Crozet. The endemism in this province is concentrated within the Mediterranean itself. Although this work is based on reef fishes. COLD-TEMPERATE AND POLAR SOUTHERN HEMISPHERE 1. molluscs. Although phylogeographic studies of fishes. Galapagos. 2010). Lusitania. Okhotsk. Eastern North Pacific Region Provinces: Aleutian. COOL REGIONS (cold-temperate and polar waters) A.

formerly constituted the St Helena–Ascension Province (Briggs. Garcı´a & Bertsch (2009) reported 37% endemism for opisthobranch gastropods in the Atlantic section. 1890. indicating that this island may be a stepping stone for colonization into the wider Atlantic.. 1990). separate St Helena and Ascension provinces (Fig. the tropical Western Atlantic was subdivided into three provinces (Briggs. there are 551 reef fishes with 64% endemism. Gaither et al. Edwards. Unfortunately.. The Atlantic section is located between Cape Hatteras and Cape Canaveral. Easter. G. However. A. 1995). Ascension has a higher affiliation with the Brazilian Province (29% shared species) than with the TEA (6% shared species). Carlin et al. Although the two islands have many trans-Atlantic species in common. 1995).. 15% TEA. Within the Gulf. (2008). 1974): Caribbean. Bermuda. The more southerly St Helena also has molluscan and crustacean fauna shared with the Indian Ocean (Smith. Western Atlantic Region The warm-temperate Carolina Province exists in two parts (Fig. to the Amazon River. Phylogeographical studies show genetically distinct populations and perhaps cryptic species at Ascension (Muss et al. the East Pacific Region (orange). their faunal composition is otherwise quite different. but has faunal affinities with both the Indo-Polynesian Province and the Western Indian Ocean Province (Winterbottom & Anderson. each has sufficient endemism (St Helena 13% and Ascension 11%) to be considered a distinct province. The Chagos is depicted here as part of the Indo-Polynesian Province. Chace. 2002. Florida and Cape Rojo. the warm-temperate Southwest Africa Province is now called the Benguela Province (BP). Together the two islands harbour 111 fish species with 26% endemism. 1992). but observed only about 5% endemism. 2) should be recognized. while St Helena has nearly equal affiliations (16% Brazilian. Avise. E. the warmtemperate biota occupies the area north of the tropical boundaries between Cape Romano. in an earlier work. Hawaii. 1966). 1990. Chagos. and crosshatching indicates the Red Sea Province. In contrast. Florida. S. Phylogeographic studies demonstrate numerous genetic partitions between these areas in co-distributed species. The Caribbean Province (CA) extends from Bermuda and Cape Canaveral. Marquesas. For the Eastern Atlantic Journal of Biogeography 39. the Atlantic section had very little endemism and was considered a subset of the northern Gulf fauna. 1997. 1).. indicating isolation between the two segments of the Carolina Province (Bowen & Avise. The name was changed to the Mid-Atlantic Ridge Province (MAR) by Floeter et al. Of the two sections. and 124 genera with 31. we retain the Carolinian Province with two recognized sections.5% endemism. Hawaiian.Marine biogeographic provinces Figure 2 Tropical regions and provinces. 2001. and the Western Indian Ocean Province (green).. and. 2006b) but also corroborate the higher affinities with the Brazilian Province (Rocha et al. Bowen et al. 2009) and tunicate endemism about 31% (Naranjo et al. Boschi (2000) recognized the northern Gulf as a Texan Province based on decapod crustaceans. To the south. Brazilian and West Indian. Despite the genetic partitions and high endemism in gastropods. one in the northern Gulf of Mexico and the other on the Atlantic coast (Briggs. 2003). species of reef fishes with some 30% endemism (Floeter et al. Ascension and St Helena. 12–30 ª 2011 Blackwell Publishing Ltd Region as a whole. 2011). He regarded the Atlantic section as belonging to a separate ‘Carolinian’ Province but found little more than 1% endemism. including the Indo-Polynesian.. Previous 15 . Briggs (1974) noted that the fishes and invertebrates exhibited about 10% endemism. such comparisons are not available for St Helena. Selected islands and archipelagos are indicated with the following abbreviations: H. However. Mexico. St Helena. and they lie 1290 km apart. C. Vertical bars indicate the Brazilian Province. Considering that each island demonstrates significant evolutionary innovation.. Marquesas and Easter Island provinces (blue). the Gulf is the richer. 2008). 1998). opisthobranch gastropod endemism is about 36% (Garcı´a & Bertsch. B. if the islands are considered separately. the East and West Atlantic regions (yellow). Galapagos. The two isolated islands on the Mid-Atlantic Ridge. Ascension. M. Formerly.

15% in echinoderms. 33% in crustaceans. Tropical Indo-West Pacific Region In contrast to the case in the Atlantic. W. (2008) also examined decapod distributions. because distribution patterns along the Chinese coast are not well known (at least in the Western literature). (2008) subsequently recognized a ‘Greater’ Caribbean Province that included all the northern Western Atlantic tropics (Fig. fish distribution along the coasts of Taiwan has been well studied (Shao et al. The region also reaches northern New Zealand. The larger Caribbean Province contains 814 species of reef fishes with about 33% being endemic. On the Chinese coast. or an area roughly the size of China. 2).J. the Mascarenes. A West Indian Province.000 km2. many Red Sea fishes (and possibly other fauna) extend into the Gulf of Aden. 1). Allen (2008) documented an average range of 9. now makes it possible to define biogeographic subdivisions with increased confidence. 2000). who recognized a southern boundary at Cape Frio. so that area needs to be added to the Red Sea Province (Fig. 2000). The northern Indian Ocean is included and so is the East African coast to the Cape of Good Hope. being under the influence of the Kuroshio Current. including the Auckland Peninsula eastwards to East Cape and 16 the offshore Kermadec Islands. Journal of Biogeography 39. the vast multitude of species that range from the Central Indian Ocean to the eastern limits of the Western Pacific give the impression of one homogeneous fauna. However. was originally recognized on the basis of considerable endemism in the fishes and several invertebrate groups. it begins at about Wenchou and extends southwards to Hong Kong (Fig. and 158 genera with about 35% endemism. but not including. to the Valdez Peninsula. while the south-eastern coast is purely tropical. 2000). However. with the exception of those occupying the shallows around tiny oceanic islands. the regional fauna begins at the Korean Peninsula and the south entrance to the Sea of Japan. The Red Sea Province is distinguished by 14% endemism in fishes (Goren & Dor. 1968). Floeter et al. comprising the islands extending from Bermuda in the north to Grenada in the south.. with 14. Included are the offshore islands of Atol das Rocas. the region extends to Robe in south-eastern Australia and to Bermagui on the south-western coast. In general. 2000).8% of the 3919 IWP species occupy areas smaller than 120. many of the putative island endemics were found along the mainland shores of the Caribbean. To the south of these boundaries. including Madagascar. although this may not be true for some of the invertebrates. 12–30 ª 2011 Blackwell Publishing Ltd . found 572 species with 11% endemism. The latter two boundaries are suggested primarily on the basis of sea surface temperature.2% endemism. the decapod crustaceans include 1058 species with about 32% endemism (Boschi. even limited-range endemics occupy much larger areas than their terrestrial counterparts. On the mainland side. The warm-temperate Argentinian Province extends from Santa Catarina. Randall. as well as many of the echinoderms and molluscs. 2). 1994. and up to 25% in corals (Cox & Moore. ending at the Taitao Peninsula (Fig.. Fernando de Noronha. 1994). Only about 10. About 25% of the coral species are endemic (Veron. Information on the reef fishes of the IWP. The tropical Brazilian Province was modified by Floeter et al. The north-western coast of the island exhibits an affinity with the warm-temperate mainland coast. 2). It may also be found along the Chinese coast and Taiwan as far as Hong Kong (Briggs. with about 1000 fish species and 142 endemics. about 19% of the West Indian fishes appeared to be endemics (Bo¨hlke & Chaplin. Briggs and B. Bowen to that subdivision.5% endemism in the Brazilian Province. the Seychelles and the Comoros. as more work was devoted to the fishes. The new Eastern Pacific Region begins at Los Angeles on the California coast and extends southwards to southern Chile. The West Indian fauna is not as distinct as it first appeared (Burgess et al. the entire region had been considered to be occupied by a homogeneous fauna. Brazil (Fig. However.070 km2 for reef fishes. However.357. Argentina. Indo-West Pacific (IWP) tropical marine provinces are characterized by prodigious numbers of wide-ranging species. 2000). In total. (2008) and now extends from the mouth of the Amazon River south to Santa Catarina. For the decapods. 1995). it begins at the tip of the Korean Peninsula and at Hamada on the lower part of the Sea of Japan. Brazil. and this observation is generally supported by phylogeographic studies (Shulman & Bermingham. The Western Indian Ocean (WIO) Province (Fig. reporting 12. 1). may be considered distinct. 1). St Paul’s Rocks and Trindade. Along the mainland shores. 2). from Cape Inubo southwards to. the Western Atlantic Region has about 1023 reef fish species with 86% endemism. The latter are considered as having restricted distributions and may merit special conservation consideration. Western Pacific Region The warm-temperate Sino-Japanese Province extends. Indo-Pacific Warm Regions The newly expanded Western Pacific Region begins in the north at Cape Inubo on the Pacific coast of Japan (Fig. 2008 and updated information from him). the Amami Islands. and the coral species have about 37% endemism (Veron. (2005) and Taylor & Hellberg (2005). 1999). There are about 471 fish species with 25% endemism. At the time of the original subdivision. C. Briggs (2005) suggested that the southern Caribbean had the richer fauna. we caution that some boundaries have not been tested or supported with data from invertebrates (see Veron. but it now appears that the fishes of the Greater Caribbean represent a homogeneous assemblage. Coelho et al. but see Baums et al. as the result of recent surveys (Allen. Boschi (2000). The Southern Hemisphere warm-temperate provinces are discussed separately. However. 1974). on the oceanic side. 1994).

within the Oregon Province. with Journal of Biogeography 39. 2000) (Fig. however. 2008). though. 2) was previously recognized as a provincial barrier (Briggs. Horn & Allen (1978) recognized a similar boundary for fishes at Monterey Bay. 2008) and reef echinoderms (Lessios et al. and the phylogeography of the rockfishes (genus Sebastes) (Sivasundar & Palumbi. but genetic breaks in some rockfish species were found at Cape Mendocino (Sivasundar & Palumbi. The IndoPolynesian Province (Fig. is now recognized between Monterey Bay and Los Angeles. a separation of the WIO and Red Sea provinces leaves the remainder of the Indian Ocean with too little endemism to distinguish it from the Western Pacific. (2008) use bathymetry profiles to demonstrate that dispersal between Australia and the Tuamotus (Polynesia) requires no deep-water traverse longer than 800 km. 2001). 2010). Laccadives. 22 short-range endemics (3. Although the Indian Ocean. This means that one can distinguish in that area the western extension of a huge biogeographic province that is larger than any of the regions in other parts of the world. two more tropical provinces were recognized: Lord Howe–Norfolk and Northwestern Australian. Mexico (Fig. The entrance to the Arabian Gulf (Fig. which lies only 650 km west of Valparaiso. 2006). This number. 1995). at the same time. 2003).. the tropical area to the east of the Philippines and Australia. need to be characteristic of and confined to the Pacific Plate. There are. including the Andaman Sea. 2006). in its entirety. Sumatra coast. with the California Province extending from the latter to Magdalena Bay. 2007. 1).2% endemism. In view of such information. 2007.. The horizontal measurement of the province extends halfway around the world: its latitudinal reach is from Sandy Cape and Shark Bay on the east and west coasts of Australia to the Amami Islands in southern Japan (Briggs. 17 .Marine biogeographic provinces The area between the Horn of Africa and the Arabian Gulf has been described as a major biogeographic barrier (Kemp. The barrier effect is demonstrated by the composition of the fish fauna of Oman (Randall. a California Transition Zone (CTZ). 2010). the Maldives and Chagos exhibits only 73 endemics. one cannot recognize a separate province for the Eastern Indian Ocean. Eastern Pacific Region A phylogenetic analysis involving the genetic structure of 40 taxa in coastal California (Dawson. 2004. In some cases. A peak in the southern range termini of cold-temperate fishes occurs at 33 N (Horn et al. and most of the others belong to the WIO Province. including 130 endemics. each area is now known to have < 10% endemism (Allen. possess sufficient endemism to qualify for provincial status (Briggs. 1995). 2010). 2000). Schultz et al.. which approximate the vicinities of Los Angeles and Monterey Bay. 2). with 415 species and 11. Pacific Plate endemics. The Pacific Plate concept was re-examined by Allen (2008). Cocos Keeling Islands. 2001. (1980) recognized clusters of geographic endpoints for northern algae species at Monterey Bay and endpoints for southern species near Los Angeles. Genetic surveys of dispersive reef organisms are consistent with the boundaries of the Indo-Polynesian Province.. 1995). This continuity of shallow habitat is doubtless a primary factor in shaping the cohesiveness of the IndoPolynesian Province. Sri Lanka. Springer (1982) identified the Pacific Plate.8%) off Oman. The Eastern Indian Ocean. located 3210 km west of the Chilean mainland. it appears that there is good reason for doing so in this case. 2010) leads to a reconsideration of the limits of the warmtemperate California Province (formerly the San Diego Province).6% endemism (Randall & Earle. and 1403 species were documented. The presence of short-range endemics has been noted in conjunction with other barriers. when compared with an approximate total of 1400 species results in 5. However... and (3) the Marquesas. This number. Sala y Gomez Island. Species endemic to a given island group are so characterized because they occur at that particular location and nowhere else. 1974).5% of them being endemic (Allen. phylogeographic studies of the Central and West Pacific show high connectivity in many reef fishes (Bay et al. 2) extends from the Arabian Gulf to the Tuamotu Archipelago (Polynesia). Christmas Island. Although transition zones within provinces have not previously been recognized (Briggs. 12–30 ª 2011 Blackwell Publishing Ltd 612 species and 25% endemics (Randall. A study of the inshore fishes of the US Line and Phoenix Islands revealed that 6. however. Therefore. While few studies extend to the Arabian Gulf. Veron (1995) regarded the Eastern Indian Ocean as continuous with the IWP. by definition. 1994). In fact. 1974). That vast area did not. south India. as a biogeographic region of major significance. Adjacent to the enormous Indo-Polynesian Province are three isolated locations whose relatively high endemism in reef fishes requires provincial status: (1) the Hawaiian Islands. Previously. also has a strong south-west Pacific component in its marine fauna (Pequen˜o & Sa´ez. Schultz et al. 2010). data on all California fishes (Allen et al. Furthermore. which is not enough to define a biogeographic province. a high incidence of short-range ‘edge-effect’ species occurs at the same two latitudes. They are not. Almost half are widespread Indo-Pacific species. includes about 2086 fish species with 532 or 25. The latter. The peaks in the range termini of the molluscs and marine algae occur between 33 and 34 N and between 36 and 37 N. 2007). Craig et al.. In terms of coral distribution.. the Juan Ferna´ndez Province. 1998). Many more species extend past Pt Conception at 34–35 N than terminate there. In earlier work (Briggs. 1995). with 169 species and 21.. this Central/West Pacific connectivity extends to the central Indian Ocean (Gaither et al.7% endemism (Randall & Cea. possesses a fish fauna with a strong Indo-Polynesian relationship so was considered to be an isolated outpost of this province (Parin. Horne et al. Murray et al. and there seems no reason for a change.3% were restricted to the Pacific Plate (Mundy et al. (2) Easter Island. included endemics specific to certain islands and archipelagos on the Pacific Plate as well as those that were widespread but confined to that area. 2008).

The tropical fauna of the Panamanian (Panamic) Province extends from the mouth of the Gulf of California south to the Gulf of Guayaquil.. The Galapagos Archipelago has 13. Within the Gulf. are examples of taxa that apparently have not changed since the formation of the Panamanian Isthmus. 1974). The contrast between the organisms occupying the warm-temperate versus cold-temperate environments is more extreme than that between the other temperature zones. Robertson & Cramer (2009) recognized a tropical Cortez Province extending from Magdalena Bay south around the tip of the Baja California Peninsula to include all of the Gulf of California.8%. A few species shared with the Caribbean. Therefore. In the extended Panamanian Province about 49% of the fish species are endemics. Cold-temperate surface temperatures for the coldest month generally range from 12 to 2 C. The other groups that retain strong faunal affinities with the Panamanian Province include the Revillagigedos. Warm-temperate waters were displaced into lower latitudes. The molluscan data from Valentine (1967) indicated a provincial endemism of about 21%. 1983) but also tend to be concentrated in cool. The difference is such that families and genera found in one usually do not appear in the other.. provincial status for the Galapagos should be retained. Several invertebrate groups have higher endemism. Cold-temperate regions A global cooling episode took place across the Eocene– Oligocene boundary c. resulting in a latitudinal restriction of the tropical seas. Around the rest of the world. Northern Hemisphere Most of the early work on the history and biogeography of the cold-temperate and cold waters of the north was carried out by Russian scientists (reviewed in Briggs. with 4. In the northern part of this range. 1974). has sufficient endemism to be considered a biogeographic province. in some groups more than 10%. This episode and subsequent cool periods resulted in cold-temperate sea surface conditions in the Arctic Ocean. 2006). but only one endemic species (Rimicola cabrilloi) (Dawson et al. that is. without regard to the origin of other species. and 18 Cocos. Their present global distributions are delineated in Fig. with 8. 1974). Hubbs (1960) determined provincial endemism at 32. the Galapagos.9%.R. North Pacific. Boschi (2000) found 38% endemism in the decapods. they occupied the circum-Antarctic region. this temperature barrier prevents the passage of many tropical species and allows speciation to take place in the adjoining warm-temperate provinces. W. with 5. The great majority of species in the Gulf range well into tropical waters. which brings more nutrients to the surface and enhances primary production. 2006).5%. Hastings. be considered endemic to a larger area. but the northern Gulf also contains about 20 California Province species with disjunct distributions (Dawson et al. a Mexican Province was previously recognized (Briggs. Owing to the high level of endemism.. the Cortez Province is retained as a warm-temperate unit within the Gulf of California. upwelling zones along the Baja coast. 2000). 2).6% endemism for shore fishes (McCosker & Rosenblatt. including the southern tips of Australia and South America. warmtemperate provinces not only are distinguished by significant endemism but also are separated from the tropics by the 20 C isotherm for the coldest month (Briggs. Boschi (2000) found 265 species of decapods in the northern part of the Gulf. The colder waters absorb more atmospheric oxygen. on the border between Ecuador and Peru (Fig. A recent Russian summary on marine biogeography (Golikov et al. and the formation of a new cold-temperate zone in each hemisphere. are trans-Pacific migrants.. The California Channel Islands have several fishes that demonstrate some genetic differentiation.6%. including decapods at 16% (Boschi. 12–30 ª 2011 Blackwell Publishing Ltd . about 163 out of 271 species (Horn et al. cold-temperate waters occupied the Arctic–North Atlantic and the North Pacific oceans at a time when the two areas were separated by the Bering land bridge.0% endemism among the shore fishes. pers. but that figure also may be too high owing to the smaller province.). The authors concluded that the initial formation of the coldtemperate faunas in the North Pacific took place coincident Journal of Biogeography 39. Robertson & Cramer (2009) placed all of the tropical offshore islands in a single Ocean Island Province. The new cold-temperate biotas were derived ultimately from tropical species that were able to adapt to the new environment (Krug et al. and their increased density stimulates thermohaline circulation. Panama. North Atlantic and the waters surrounding the Antarctic continent. 2009). comm. A significant number of species. but only one archipelago. 1990) paid particular attention to the Northern Hemisphere and reviewed the climatic history as well as the biogeographic subdivisions. This results in an increased upwelling. at the same time. As noted previously for the Pacific Plate.J. 2006).. Robertson. We suggest that this province should still be confined to the Gulf and be considered warm-temperate. and the other offshore islands should be regarded as outposts of the Panamanian Province. 1. with 2. Many of the northern species are usually found in relatively deep water (Eschmeyer et al. but that figure may be too large considering that the province now covers a smaller area. but the more recent information from Robertson & Cramer (2009) indicates that the section from the mouth of the Gulf of California to the Gulf of Tehuantepec does not demonstrate sufficient endemism. the barrier extends across the southern end of the Gulf of California approximately between La Paz and Topolobampo. In the Northern Hemisphere. Provincial recognition is given according to endemism. Smithsonian Research Institute. with 9% endemism.. 1974.. 2001). Malpelo. 2010) and has been continuously recognized as a separate province (Briggs. Clipperton. 2009). In this case. either exclusively or also with the Panamanian Province. species endemic to a particular island cannot. Bowen The California Province as reconstituted still contains large numbers of northern fishes. C. 35 Ma (Zachos et al. Briggs and B. slightly more than 10% of the fishes are endemic (D. In the south.

In addition. 2004). 12–30 ª 2011 Blackwell Publishing Ltd southwards to the Strait of Belle Isle in the west and to the north-east beyond the Murmansk Peninsula. and (3) an Estuary–Arctic Interzonal Province was noted to occur along the shores of the Arctic Ocean. The northern (Aleutian) province has an endemism rate of about 24% in decapods and 23% in molluscs (Valentine. the Oregon Province. In the North Pacific. 1967. In each ocean. Horn et al. the northern coldtemperate biota may be much older than originally thought. From a world-wide perspective. 2004). 1978. At the time of the Great Interchange. often-called ‘boreal’. 2000).. More recently. In the North Atlantic. Global cooling during this (mid-Pliocene) interval was probably caused by four key tectonic events: (1) the isolation of Antarctica. the new cold-temperate.5 Ma. and (4) the uplift of the Panamanian isthmus (Crame. and a peak in range termination endpoints at about 50 N. Boschi. (2009) presented evidence for ice formation in the Arctic Ocean in the middle Eocene (47 Ma). The contemporary result is a distinct boreal region on each side of each ocean. Based on these findings. but a good part of the relationship between the two regions is caused by the large number of Pacific species that invaded in the mid-Pliocene. 3. In the Pacific. the A/B boundary was placed at the Bering Strait. fish (Merluccius spp. but by c. these southern extensions of Arctic water divided the original Pliocene boreal assemblage into eastern and western components. the Arctic Ocean had little ice. In contrast. however. the biotas occupied areas that had been previously outlined (Briggs. biotas evolved separately until the late Miocene when marine connections across the Bering land bridge began to develop. defined in terms of endemic species. 2000. the southern limit of the Oregon Province.Marine biogeographic provinces with a significant temperature fall about 14 Ma. 2001. If. An important effect of the mid-Pliocene cooling of the northern oceans was the separation of boreal biotas (Briggs. if considered to terminate at Monterey Bay. has only about 2% endemism in decapods and fishes (Horn & Allen. In the North Pacific and Arctic–North Atlantic. an event subsequently called the Great Trans-Arctic Interchange. With regard to longitudinal relationships. 2000. subregions and provinces. Vermeij. Typical boreal species were no longer able to maintain amphi-Atlantic and amphi-Pacific distributions. the Arctic biota extends southwards to Cape Olyutorsky in the west and Nunivak Island to the east. near the northern tip of Vancouver Island. In the Atlantic. However.. the endemism level would probably rise to more than 10%. Stickley et al. Boschi. it was generally thought that the land bridge remained intact until c. 2002). there are also Arctic-boreal species. for the most part. The North Atlantic change is adopted here (Fig. Much of the native North Atlantic molluscan fauna originated in European waters and then spread westwards (Wares & Cunningham. 3. the Arctic/Boreal (A/B) boundary was extended northwards to Svalbard and the south end of Novaya Zemlya. and in both oceans the eastern and western faunas developed independent evolutionary trajectories. the boundary between the 19 .. Grant & Leslie. The final event apparently produced a major intensification of Northern Hemisphere glaciations between 2. it is apparent that. This region may be divided into Aleutian and Oregon provinces. including faunas as diverse as seagrass (Zostera marina. some Pliocene amphiboreal species have apparently not yet developed specific differences. 1) but the justification for the Bering Strait boundary does not appear strong. and the modern Arctic marine fauna began to develop. Their descriptions and maps indicated that. 2006). 2001) and several invertebrate groups (Wares & Cunningham. The northern boundary is concordant with the mean southern limit of the pack ice in January–February. most of the boreal species were eliminated. 2004). the east and west boreal faunas are closely related. Previously. see also Kafanov & Kudryashov. This separation is also apparent in phylogeographic studies both within and between species. (1990. Sediment cores in the polar North Atlantic detected ice-rafted debris 14– 12 Ma (Thiede et al. in each ocean. the Arctic sea surface temperature for the coldest month dropped to between +2 and )2 C.3 Ma (Gladenkov et al. The authors recognized a kingdom of temperate and cold waters that was subdivided into regions. Approximately half of the molluscan invaders have speciated and many of them are now endemic to one boreal region or the other (Vermeij. 1974). respectively). 1991a. So. Multiple biogeographic subdivisions were suggested by Golikov et al. 1998). The mid-Pliocene cooling of the northern oceans resulted in the resumed isolation of Atlantic and Pacific boreal biotas. but the northern oceans were more finely divided. with a boundary previously described at the Dixon Entrance (55 N). 2005).. (2) in the North Pacific. Addison & Hart. the cold-temperate biota of the CTZ is included.5 Ma it allowed an unrestricted mingling of biotas that had been separated for more than 30 Myr (Vermeij. 1991b). (2006) reported many California fish range terminations at the latitude of Monterey Bay (36– 37 N). respectively. 2001. including the California Transition Zone (CTZ). The Estuary–Arctic Interzonal area is probably best defined as a special ecological zone rather than as a biogeographic province. Horn et al. recent fossil studies indicate that the first opening may have occurred as early as 5. 1995). Eastern North Pacific Region Cold-temperate conditions extend from Nunivak Island in the north to about Los Angeles on the California coast.4 Ma (Mudelsee & Raymo. the cold-water Arctic Region now extends Journal of Biogeography 39. As a result. 2005).9 and 2. and cold-temperate conditions prevailed. 2005b).. the relationship is primarily due to the presence of a group of Arctic-boreal species common to both sides of the ocean. but there were some notable exceptions: (1) in the Eastern North Atlantic. (3) the collision of Australia with Southeast Asia. Olsen et al. the various regions were delineated about as they had been for the past 25 years. When the Bering Strait first opened it may have been shallow with limited passage. (2) the closure of the Tethys Sea. This coincides with a similar peak reported by Peden & Wilson (1976) based on fish distributions in British Columbia and Alaska.

However. East–west relationships. south-western Sea of Japan. This province extends northwards along the Kurile chain of islands and the east coast of the Kamchatka Peninsula to about Cape Olyutorsky. The divisions were made on the basis of breaks in the species diversity gradient and their relationship to temperature and prevailing currents. There is about 19% regional endemism in shore fishes (Bigelow & Schroeder. 1974). Indeed. one may find a different species assemblage. W. (2000). An Oriental Province exists in three segments (Fig. The Okhotsk Province (hatched) is confined to the Sea of Okhotsk (O). In contrast. Journal of Biogeography 39. 2007). Figure 3 Three Northwest Pacific provinces. and only 5% in decapods (Boschi. The more recent literature pertaining to amphi-Pacific relationships has been reviewed by Ilves & Taylor (2007). the complex geological history with periodic isolations of the Sea of Japan and the Okhotsk Sea was probably important in generating diversity. 3). ascidians and fishes (Briggs. 1). Western North Pacific Region Cold-temperate conditions in the Northwest Pacific apply to three provinces. Although there are no recent taxonomic evaluations (known to us). phylogeographic analyses indicate that the Northwest Pacific marginal seas were isolated during glacial maxima (Liu et al. the Oriental Province extends from about Hamada to the Tsugaru Strait. See text for precise boundaries. The first extends north from the warm-temperate boundary at Wenchou and continues through the Yellow Sea. 1). together with very little endemism. Numerous species. Liparididae. 1990) (Fig. polychaetes. recognized significant differences. often called the ‘Middle Atlantic Seaboard’. 1974). the fish family Embiotocidae and the gastropod genera Nucella and Littorina may have originated on the eastern side (Ilves & Taylor. pycnogonids and fishes (Briggs. The Kurile Province (dots) extends across the Pacific side of the Kamchatka Peninsula (K) to the southern tip of Hokkaido (H) and the Sea of Japan (J). On the eastern side of the Sea of Japan. Its continuity is broken by the tip of the Korean Peninsula. On the western side. shallow basin almost completely enclosed to the north and bordered by the Alaskan Peninsula and the Aleutian islands to the south. 12–30 ª 2011 Blackwell Publishing Ltd . Previously. The fish families Cottidae. beginning with Andriashev (1939). 2007). The area is penetrated during the summer months by large numbers of tropical and warm-temperate organisms. one is confronted with a good deal of conflicting opinion. The Okhotsk Province is confined to the Sea of Okhotsk. Zoarcidae. In considering the geographic extent of this cold-temperate region. shows that it clearly belongs to the Boreal Region (Briggs. 1974). but several investigators. Bowen Aleutian and the Oregon provinces should be shifted south from the Dixon Entrance to the northern tip of Vancouver Island. 2009). C. the older literature demonstrated considerable endemism in ascidians. some of the Russian biologists preferred to recognize more provinces than those just described.. one penetrating from the north and the other from the south. 20 As noted. 2000). A faunal break exists at about the location of the Tsugaru Strait between the islands of Honshu and Hokkaido.. Japan. in the Sea of Japan the fish fauna was separated into four provinces by Kafanov et al. The Oriental Province comprises three parts: the Yellow Sea (Y). the currently used scheme indicates only two provinces in the Sea of Japan. 3). and the CTZ included within the Oregon Province. the presence of large numbers of boreal species. considered to be endemic to one side or the other. 53% in all molluscs (Vermeij. To the north of this point. The Bering Sea is essentially a broad. it continues southwards on the outer coast of Honshu Island to Cape Inubo. Most of the disagreement is concerned with the relationship of the fauna that occupies the area between Cape Hatteras and Cape Cod. as well as the genera Oncorhynchus and Sebastes (Hyde & Vetter. 21% in opisthobranch gastropods (Garcı´a & Bertsch. From that point. 1953). Western Atlantic Boreal Region This region extends from the Strait of Belle Isle to Cape Hatteras (Fig. 1999). The provinces that were identified reflected interesting ecological differences but did not exhibit sufficient endemism to qualify as provinces according to the 10% rule. The absence of obvious barriers might lead one to expect a homogeneous marine fauna. and northern Honshu Island (Ho). Briggs and B. Eastern Atlantic Boreal Region This region is now considered to extend from Svalbard and the southern end of Novaya Zemlya to the southern entrance of the English Channel (Golikov et al. defined in part by the complex geological history of the Sea of Japan and adjacent regions (Wang. but it then continues up the north side of the peninsula to about Chongjin. probably underwent major radiations in that area. 2005c). This has often resulted in its being allied with the Carolina Province to the south. In contrast. both along the outer coast and within the Sea of Japan. of the Kurile Province (Fig. are documented among the anomuran crabs. it was probably isolated during the glacial stages and perhaps earlier. Although this sea is now confluent with the North Pacific through the Kurile Islands and with the Sea of Japan around Sakhalin Island.J. 2007). For example.

An exception is the narwhal. More than 50% of those species had their ultimate origins in the North Pacific. From this point. 2004).. and Garcı´a & Bertsch (2009) estimate 25% for the opisthobranch gastropods alone. The Southwestern (Flindersian) and Southeastern (Peronian) provinces of Australia (Briggs. 1). 2010). Endemism in Arctic fishes Journal of Biogeography 39. 1995). although the provincial 21 . The salinity is relatively stable and decreases gradually towards the inner end of the long. 1974). Furthermore. an early estimate was 20–25% endemism for both fishes and invertebrates (Briggs.000–20. when considered with the strong European relationship of Iceland. with no endemics or any indication of relationship to the Western Atlantic (Magnussen. Monodon monoceros. 124 species had transatlantic ranges. The demersal fish fauna is closely related to that of the North Sea. 1974) suggested a purely boreal component. or at least the south and east shores. and of endemics with nearby tropical ancestors. as demonstrated by the superior species diversity in fishes (Wheeler. the Falkland Islands. Although Golikov et al. there are very few endemics at the higher taxonomic levels.4 Ma (Mudelsee & Raymo. Waters. there is considerable longitudinal relationship due to the influence of the West Wind Drift (WWD).. The polar cod (Boreogadus saida) may be considered an indicator species because it extends to all parts of the region but no farther (Cohen et al. although there are significant numbers of endemic species. The spiny lobster genus Jasus provides another example (Pollock. amphipods and echinoderms (Briggs. It was apparently the ancestral habitat for the seastar Patiriella exigua.000 years ago). the present glacial regime probably began about 2.9–2. a group of 21 volcanic islands located between Iceland and the Shetlands. 2004). host species that are either boreal or Arctic-boreal. For the entire region. 1999). 1960).5 Myr. 1990). The Arctic Region occupies all of the area north of the cold-temperate boundaries previously identified. 2005). The Agulhas Province extends from the Cape of Good Hope north-eastwards to about the mouth of the Kei River. 1990). but recent works indicate an essentially homogeneous biota so that a single region is now recognized. The Faroes. Iceland possesses an interesting biotic mixture. based on the work of Bennett & Pope (1953. The Agulhas Province contains large numbers of eurythermic species shared with the tropical WIO Province. 12–30 ª 2011 Blackwell Publishing Ltd has been estimated at about 25% (Eastman. Recently. 2001). A number of taxa originating in Australia–New Zealand were able to achieve circum-global ranges via the WWD. In the Southern Hemisphere (Fig.. Cold Arctic Region Although sea ice has been detected in the Arctic as far back as 47 Ma (Stickley et al. 2009). The richest boreal fauna occurs on the eastern side of the North Atlantic. The relationships are almost entirely with the Eastern Atlantic. and the earlier literature indicates high endemism in sponges. The Arctic seas have traditionally been divided into a number of separate zones and provinces.Marine biogeographic provinces the northern boundary was located at the base of the Murmansk Peninsula. are still recognized in the recent literature (Waters et al. Southern Hemisphere Warm-temperate provinces Owing to the presence of many tropical eurythermic species. 1999.. pure Arctic. This. Pacific and Atlantic oceans (Waters & Roy. this suggestion has been reinforced by genetic evidence (Burridge & Smolenski. Tristan da Cunha and the tip of southern Africa (RochaOlivares et al.. 1997). 1974). Phylogeographic analyses of the Arctic charr (Salvelinus alpinus) demonstrate that most of this region was recolonized after the last glacial retreat (10. Vermeij (2005c) determined that all of the transatlantic species had apparently dispersed from Europe to America within the past 3. Genetic and morphological evidence has indicated that two species of chironemid fishes (Chironemidae) found on Juan Ferna´ndez Island represent colonizations from Australia–New Zealand (Burridge et al. The Baltic Sea is the world’s largest estuarine area. they reached. with a relict distribution (Jefferson et al. This suggestion has been verified by Vermeij (2005c). who examined mollusc distribution in the North Atlantic. this designation does not appear to be justified on the basis of endemism. Three species of the fish genus Sebastes (Scorpaenidae) dispersed from the North Pacific southwards through the Eastern Pacific to Tierra del Fuego. the faunal relationship of most northern warm-temperate provinces is closest to their adjoining tropical province. (1990) recognized a Baltic Province.. narrow basin. so the polar biota of the Arctic Region is much younger than that of the Antarctic.. 1993). and some eurythermic temperate forms. which was subsequently transported by the WWD across the Indian. New data from Vermeij (2005c) indicate that about 69% of the molluscs are endemic. indicating a lack of substantial biogeographic barriers (Brunner et al. 1974). should be included in the Eastern Atlantic Boreal Region. Other recent phylogeographic studies have produced strong evidence for WWD dispersal in the Southern Hemisphere (Waters & Burridge. The older literature (Briggs. Arctic-boreal. apparently via the WWD. He found 402 extant species on the eastern side (69% endemic) and 262 on the west (54% endemic). 1974). This province has been reported to possess high levels of invertebrate endemism (Griffiths et al. indicated that the principal evolutionary centre for the recent Atlantic boreal fauna was on the eastern side of that ocean (Briggs. Consequently. The absence of any special American relationship and the almost complete absence of endemics (except for a few subspecies) indicate that Iceland. 2006). 2009) but it is not known how many of the suspected endemics actually extend northwards into the tropics of the south-eastern African coast.. 2008). however. Such a dispersal history had been proposed for the fish families Cheilodactylidae and Latridae (Briggs. 1969). 2002). East–west relationships.

Cyclostomata and Cheilostomata) was determined by Griffiths et al. southern New Zealand and nearby islands. (2009) illustrates a sharp change from a warm-temperate to coldtemperate fauna near the Valdes Peninsula (42 S). South American Region. Clinidae. with two provinces in the New Zealand Region and five provinces in the Sub-Antarctic. 1974). The external relationship proved to be stronger to the south-west Pacific than to the Chilean coast. This procedure resulted in the designation of 12 provinces and four regions within the Southern Ocean area. but was designated as a province by Briggs (1974). 296 gastropods. Garcı´a & Bertsch (2009) reported 24% endemism in opisthobranch gastropods in the same area. respectively (Griffiths et al. concordant with the faunal barrier for sea anemones (Haussermann & Forsterra. However. Endemism in four classes of benthic invertebrates (Bivalvia. the southern boundary was extended to the Valdes Peninsula. Later. The Argentinian Province lies between Santa Catarina. 2009). Briggs and B. all located in warm-temperate provinces. W. Argentina (Fig. The Auckland Province shares many species and has a historical relationship with southern Australia (Waters et al. In the south-eastern Atlantic. 1995). endemic rates of 60% for bivalves. 12–30 ª 2011 Blackwell Publishing Ltd . These areas had been apportioned into four regions and seven provinces (Briggs. The early literature on fishes. the invertebrates so far as known appear to be unrelated. Although the endemism percentages varied among the classes. 1974). The inclusion of Macquarie brings the total number of cold-temperate provinces to 13. especially by Linse et al.5 S) as the southern limit of the province. 69% and 69%. 2009) and 31% for ascidians (Primo & Va´zquez. Nearby Gough Island probably should be included so we can recognize a Tristan–Gough Province. 2000). However. invertebrates and algae indicated considerable endemism. Tasmania and Victoria in Australia.. Briggs. 2008). (2006). Boschi (2000). Brazil (Floeter et al. it extended from Cape Frio. and the beginning of the cold-temperate waters at 43º S.J. Tasmanian. corroborated by a recent survey of the littoral fish fauna that revealed 25. 2008). and SubAntarctic regions. 1995): South America (Magellan). (2009). originally considered to be a distinct province (Briggs. 2007). An endemism rate of 13% was noted for the decapod fauna (Boschi. The endemic rate in the Benguela Province is about 12% for fishes and 53% for opisthobranch gastropods (Garcı´a & Bertsch. The Kermadec Province invertebrates revealed a large number of species. Amsterdam–St Paul. The Juan Ferna´ndez Province consists of three islands 650 km west of Valparaiso. Macquarie Island was not surveyed by Griffiths et al. For New Zealand.. 31% of the ascidians are endemic (Primo & Va´zquez. Lee et al. Nototheniidae. Five cold-temperate fish families (Bovichtidae. Previously labelled the Eastern South America Region (Briggs. Gastropoda. there may be a considerable endemic component although there are no published estimates. The Cape is the dividing line between the Benguela and Agulhas provinces.. by itself.5% endemism (Pequen˜o & Sa´ez. an intermediate zone from 40 to 43 S. 100% for gastropods (Griffiths et al. extending from the Gulf of Guayaquil to about the Taitao Peninsula. Briggs (1995) designated Chiloe Island (41. 2007) indicate a highly distinct invertebrate fauna. Brazil and the Valdes Peninsula. 1991. 1964). Tristan da Cunha and the Kermadec Islands. the warm-temperate Benguela Province is located between Mossamedes and the Cape of Good Hope (Floeter et al. 1). Two sets of widely separated offshore islands. illustrating the dramatic faunal transition in this boundary area. Clarke et al. we recognize the following geographic areas as provinces if at least two classes have endemism rates of more than 10%.. Congiopodidae. the southern cold-temperate and cold regions and provinces can now be more confidently defined. (2008) referred to a Peruvian/Chilean Province extending to around 40 S. for both the fishes and the decapod fauna (Boschi. Marine fishes of southern Chile also show three latitudinal fish zones but they are located farther south by Sielfeld & Vargas (1999).. The provincial status was based on a 30–40% endemism rate for the shore fishes. (2009). the former exhibiting an Atlantic relationship and the latter linked to the 22 Indo-Pacific. Brazil to the Rı´o de la Plata. Moridae) do not extend north of this region. 2005). 1995) and the northern boundary was set at Santa Catarina. there are few endemic fishes. with endemism rates of 71%. New Zealand. formerly comprised the warm-temperate West Wind Drift Province (Collette & Parin. because of much recent work on the cold-temperate invertebrate fauna. Tristan–Gough and Amsterdam–St Paul. reported 13% endemism in the decapod crustaceans. However for Tristan da Cunha. The warm-temperate Peru–Chilean Province (Fig. the Auckland and Kermadec provinces are recognized. Previously the cold-temperate waters of South America were united in a Magellan Province spanning from Chiloe Island on the Pacific side to Rı´o de la Plata Journal of Biogeography 39. Normanichthyidae) extended as far as 45–46 S. These authors indicated that species belonging to typical warm-temperate families (Blenniidae. 2009). but the Taitao Peninsula now seems to be more appropriate. 2000). 2007). Cold-temperate regions Cold-temperate waters are found around the tip of South America and the Falkland Islands. 1). should retain that status. All three showed exceedingly high endemism. 77 bivalves and 203 cheilostome bryozoans in this small area. who accepted the Cape Frio boundary. 2000). Although the fish fauna indicates a relationship between the two island groups. Eleginopidae. (2009) based on more than 7000 specimens collected from all parts of the Antarctic and SubAntarctic. Argentina (Briggs. and the Sub-Antarctic (Fig. Bowen names have varied. A new assessment of the reef fish fauna by Galvan et al. Chile. so they were not included in the following cold-temperate arrangement. (2007) and Griffiths et al. Three of the collection localities were in South Africa. based on a 66% molluscan endemism (Dell. 1) includes the major part of the western coast of South America. C.

23% for the Bounties. In contrast. Prince Edward Island is a separate province and probably should include the Marion Islands based on proximity. Southern Argentina (SA) and the Falkland Islands (F). their data indicate about 35% endemism for the Pacific side and about 18% for the Atlantic. 5).. 2009. 2009) indicate that all four areas should be designated as provinces within a South American Region (Fig. Journal of Biogeography 39. 23 . 1999). Antipodes (AN) and Bounty Islands (B). but now must be assigned to a separate province based on two of the four invertebrate classes (Griffiths et al. Although Boschi & Gavio (2005) recognized a single Magellan Province for the decapod crustaceans. but now we recognize six (Griffiths et al. 2009). 2003). New provinces are Southern Chile (SC).. and about 34% of the shore fishes may be endemic (Briggs. demonstrates elevated endemism for the two molluscan classes (Griffiths et al. (2009). Tierra del Fuego. the shore fish fauna of the region shows no indications of provincial endemism (Sielfeld & Vargas. southern Argentina and the Falkland Islands (Griffiths et al. Antipodes. However. and asked if all these island groups should be provinces. Notably. The shelf waters of the Antarctic and Sub-Antarctic are occupied by a highly distinctive fauna that owes its origin to four historical factors: (1) the persistence of a small ancestral group of Mesozoic and early Cenozoic taxa. Five provinces in the Sub-Antarctic Region were previously recognized (Briggs. Haussermann & Forsterra (2005) noted that the polychaetes and anemones indicated a barrier between the Pacific and Atlantic sections because those faunas showed very little overlap.. 1974).5%) and the gastropods (86. the molluscs of the Chilean coast do not demonstrate a reduction in species diversity at higher latitudes. The coastal fish fauna of 270 species has 25% endemism (Walrond. an area not reported on by Griffiths et al. Sub-Antarctic Region. Likewise. 1974). The fauna of South Georgia is highly endemic for three out of the four invertebrate classes. The Crozet Islands were previously considered part of the Kerguelen Province.Marine biogeographic provinces (Argentina/Uruguay border) on the Atlantic side (Briggs. this province may extend to the Victoria coast of Australia (Briggs. 6). Tierra del Fuego (TF). Is there sufficient endemism to recognize a separate province? The Antipodes Province (Fig. 2009).. (2009). Kerguelen Island has a distinctive molluscan fauna. consisting of the Aucklands (A). New Zealand has very high endemism for all four invertebrate classes. 12–30 ª 2011 Blackwell Publishing Ltd 150° 160° C 180° Figure 5 Cold-temperate provinces of the Southwest Pacific. but rather a sharp increase in diversity above 42 S (Valdovinos et al. However. the very high endemism rates for invertebrates in southern Chile. Even so. Fig. 2009). and the endemism figures provided by MacDiarmid & Patuawa (2010) for the bivalves (85. the answer still eludes us. consisting of the Auckland.. (2) the extinction of many early Tertiary warm-temperate species. New Zealand–Australia Region. Almost 50 years later. High endemism for other New Zealand marine groups are indicated in the online summary edited by MacDiarmid (2010). In their review of invertebrate zoogeographic patterns in the Magellan Province.. 2009). (3) geographical isolation produced by the opening of Drake Passage. and 23% for the Snares. It should be noted that the north coast between Auckland and East Cape is in the warm-temperate zone.6%) are considerably higher than those listed by Griffiths et al. Campbell (C). 4). The Tasmania (Maugean) Province (Fig. This represents a significant change from the previous scheme that assumed an undivided fauna for the entire region. Dell (1962) had found molluscan endemism rates of 16% for the Aucklands. 5) has an exceedingly high endemism for both molluscan classes (Griffiths et al. Campbell and Bounty Islands. These references provide additional justification for a separation between Southern Chile and Southern Argentina provinces. including the Tasmanian Province (T). However. the province is very distinct. T 40° N B AN 50° A 140° Figure 4 The cold-temperate South American Region (northern boundaries are Valdez Peninsula on the east coast and Taitao Peninsula on the west coast). so the endemism figures probably will be adjusted downwards in the future. and (4) invasions by cold-temperate species from the North Pacific. the New Zealand Province (N) and the Antipodes Province. The Bouvet Province is now known to have a very distinct fauna with 50% endemism in gastropods. 1995). 1995).

J. (2009). c. 2007) and the anemones (Rodrı´guez et al. the Arctic Ocean did not decline to similar temperatures until about 2. 2005). 1974) indicates that about 66% of the Kerguelen shore fishes may be endemic species. endemism and new biogeographic provinces. and four notothenioids (ice fishes.9–2. Previously. Kerguelen (K). 2007). The fauna of the Antarctic Region is relatively old compared with that of the Arctic Region. Africa (AF). 2005. (2009). Considering the 20+ Myr of isolation. but see Stickley et al. Eastman. Together. As noted. 2007) and older references indicate a 64% molluscan endemism (Dell. 2005). W. C. 12–30 ª 2011 Blackwell Publishing Ltd . 1981). but four of nine ascidian species are endemic (Primo & Va´zquez. and the unique notothenioids probably arose in the Antarctic. with the notable exception of the ephemeral Indo-Pacific barrier. and Macquarie Island (M). NZI). underexplored regions of the planet have revealed high diversity. South Sandwich and South Shetland islands are all below the February 1 C isotherm (Dietrick. Many temperate marine biota originated in the tropics. 2007). The Antarctic Continent and the South Orkney. 2010). the phylogenetic and taxonomic affinities of warmtemperate and adjacent tropical provinces (relative to coldtemperate provinces) indicate that they should be united in a single warm region.. 6). the ascidians (Primo & Va´zquez. New Zealand (NZ) and New Zealand Islands (Antipodes. Early literature (Briggs. Macquarie Island was not reported on by Griffiths et al. 67 liparidids (Liparididae) and 23 zoarcids (Zoarcidae). (2009). The fishes have restricted taxonomic diversity but an endemism rate of 88% on the continental shelf and upper slope (including depths to 1200 m. They also 24 determined a general endemism level between 42% and 56% for the four invertebrate classes surveyed in Griffiths et al. First. but there are also exceedingly large numbers of endemic genera (76%. Second. Australia (AU). Janko et al. 2008)... This does not affect the status of individual provinces in the tropics and warm-temperate zones. CONCLUSIONS AND CONSERVATION IMPLICATIONS The vast reservoir of new biogeographical information emerging since Briggs (1995) has revealed several trends. Bouvet Island (B).. the Crozet Islands (C). sufficient phylogeographic information now exists to conclude that genetic architecture (primarily within species) and the biogeographic structures defined by endemism are largely concordant. which for the most part is not related to cold-temperate faunas in adjacent regions. The Sub-Antarctic provinces include South Georgia (SG). Eastman. We anticipate that the combination of warm (temperate and tropical) regions should more closely align marine biogeography with the evolutionary relationships discovered in the oceans. Also indicated are the southern tips of South America (SA). and the alignment of tropical and warm-temperate regions is intended to accommodate this relationship. this degree of species endemism is not unexpected. The fast-moving Antarctic Circumpolar Current must facilitate the high dispersal observed in this region. Bowen Figure 6 Antarctic and Sub-Antarctic regions. Prince Edward Islands (PE). Knox (1994) had observed that more than half the invertebrate species were endemics. 2009). most notably in the Southern Ocean. The latter two families represent invasions from the North Pacific. There is evidence of a major ice sheet by late Oligocene times. 1964). Briggs and B. in contrast to many earlier workers who divided the continent into various segments. The cold Antarctic Region is indicated by shading. The shelf and upper slope support about 222 fish species. and about 73% of its ascidian species are endemic (Primo & Va´zquez. In contrast. 2007). Estimates for other invertebrate groups are also high: 51% for sponges. Third. indicate similar levels of endemism.. The Region includes all of the waters surrounding the continent and the noted island groups (Fig. this province probably should include the nearby Heard and McDonald islands. Cold Antarctic Region. This agrees with the conclusion of Griffiths et al. showing little (or no) population structure across this vast region for two decapods (Raupach et al. 57% for polychaetes and 75% for molluscs (Arntz et al. Phylogeographic studies are consistent with a single Antarctic Province. but more accurately reflects the alignment of provinces into warm (temperate and tropical) and cool (temperate and cold) regions. 24 Ma (Ivany et al. who recognized a single Antarctic ‘Province’. This Journal of Biogeography 39. Other invertebrate classes recently investigated. 1997).4 Ma (Mudelsee & Raymo. one nemertean (Thornhill et al. 2006). including 96 notothenioids (five families).. indicating close to modern conditions. these three groups account for more than 85% of the fish fauna..

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& Anderson.A. C.J. K. Brian W. & Gurgel. Zuccarello. Connell. (2010) Australia’s marine biogeography revisited: back to the future? Austral Ecology.C.. T. Wernberg. C. Thomas... J.. 34. 686–693. and ecosystem connectivity in the Papahanaumokuakea Marine National Monument in the Northwest Hawaiian Islands. Briggs is a professor emeritus from the University of South Florida and is now affiliated with Oregon State University.. His research deals primarily with the origin and distribution of contemporary groups of organisms.. Kraft. Smith Institute of Ichthyology Ichthyological Bulletin. (2001) Trends.M. London. A. He studies the phylogeography of marine vertebrates. & Billups. 1987) and Global Biogeography (Elsevier. In 2005. with a focus on the origin and maintenance of marine biodiversity. M. 35. Central Indian Ocean. J. Thomsen. J.. Waters. Pagani.D. 678–687. Editor: Alistair Crame 30 Journal of Biogeography 39. 66. J. L. M. R. J. Macmillan.C. Science. (1969) The fishes of the British Isles and North-West Europe. 1– E. Bowen is a research professor for the University of Hawaii at the Hawaii Institute of Marine Biology (http:// www. R.. Briggs and B. 1995). 988–992. G. 1974).B. 292. Biogeography and Plate Tectonics (Elsevier. Wheeler.F.C.T. (1997) A revised checklist of the epipelagic and shore fishes of the Chagos Archipelago. Journal of Biogeography. Zachos.S. Sloan. Winterbottom. West.L.D. G. Sanderson. S.hawaii. BIOSKETCHES John C... 12–30 ª 2011 Blackwell Publishing Ltd . rhythms and aberrations in global climate 65 Ma to present. W. Professor Briggs received the Alfred Russel Wallace Award from the International Biogeography Society for his lifetime contributions to biogeography. Bowen gastropods: implications for trans-Tasman dispersal. His biogeographical books include Marine Zoogeography (McGraw-Hill.