The Marine East Indies: Diversity and ARTICLE Speciation John C

Journal of Biogeography (J. Biogeogr.) (2005) 32, 1517–1522

ORIGINAL The marine East Indies: diversity and ARTICLE speciation John C. Briggs*

Georgia Museum of Natural History, The ABSTRACT University of Georgia, Athens, GA, USA Aim To discuss the impact of new diversity information and to utilize recent findings on modes of speciation in order to clarify the evolutionary significance of the East Indies Triangle. Location The Indo-Pacific Ocean. Methods Analysis of information on species diversity, distribution patterns and speciation for comparative purposes. Results Information from a broad-scale survey of Indo-Pacific fishes has provided strong support for the theory that the East Indies Triangle has been operating as a centre of origin. It has become apparent that more than two-thirds of the reef fishes inhabiting the Indo-Pacific are represented in the Triangle. An astounding total of 1111 species, more than are known from the entire tropical Atlantic, were reported from one locality on the small Indonesian island of Flores. New information on speciation modes indicates that the several unique characteristics of the East Indian fauna are probably due to the predominance of competitive (sympatric) speciation. Main conclusions It is proposed that, within the East Indies, the high species diversity, the production of dominant species, and the presence of newly formed species, are due to natural selection being involved in reproductive isolation, the first step in the sympatric speciation process. In contrast, speciation in the peripheral areas is predominately allopatric. Species formed by allopatry are the direct result of barriers to gene flow. In this case, reproductive isolation may be seen as a physical process that does not involve natural selection. Allopatric species formation often takes millions of years, while the sympatric process is generally much faster. Following species formation, dispersal from the East Indies appears to take place according to the centrifugal hypothesis. Keywords *Correspondence: John C. Briggs, 82-651 Sky View Lane, Indio, CA, USA. Allopatry, East Indies, Indo-Pacific, natural selection, species diversity, E-mail: clingfi[email protected] sympatry.

year. Biologists have done more work on the fishes than on INTRODUCTION any other group, yet they are still not well known. But one The East Indies Triangle, a relatively small part of the Indo- can now make a reasonably good prediction about the Pacific Ocean, is unique in terms of its species diversity and number of species that are there and can reach some general evolutionary importance. Within this area (Fig. 1), the high conclusions about their formation and geographical distri- species diversity of the major groups of marine organisms, bution. Species from the Triangle probably play a major role such as the fishes, corals, echinoderms and molluscs, is in the assemblage and maintenance of coral reef commu- almost unbelievable. Although species from the Triangle nities throughout the Indo-Pacific (Mora et al., 2003). The have been collected and described since the early part of the influence of such species is considered to be related to their nineteenth century, new ones are being discovered every mode of speciation.

ª 2005 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi doi:10.1111/j.1365-2699.2005.01266.x 1517 J. C. Briggs

Figure 1 The East Indies Triangle in the Indo-West Paciﬁc Ocean, a centre of origin for dominant species.

sponges and other animals. As noted by Allen & Adrim, the DIVERSITY cryptic element generally comprises c. 3–5% of the observable A recent review of the coral reef fishes of Indonesia by Allen & species for a given locality. That such an incredible number of Adrim (2003) was instrumental in allowing a better under- species can exist in one bay of one small island, emphasizes standing of the ichthyofauna. Although the Indonesian area, as that we have much to learn about community ecology and recognized by those authors, did not reach the Philippines and evolution. the eastern half of New Guinea, most of the Triangle was The number of coral reef fishes that exists in the entire Indo- included. So, the results of the review are generally applicable Pacific has been estimated at 3764 (Allen & Adrim, 2003). This to the larger area. A total of 2057 fish species are now known to means that more than two-thirds of the reef species in the occur among the coral reefs in Indonesian waters. If the area Indo-Pacific, a region that stretches across about two-thirds of were to be expanded to include the remainder of the Triangle, the globe, also occur within the East Indies, an area that the species diversity would likely exceed 3000. There are represents a small fraction of the whole. Furthermore, 97 of the known to be c. 450 reef coral species in the latter area (Veron, species are considered to be endemic to Indonesian waters. It 1995) but there are no reliable estimates for the other also appears that these endemic species are not generally invertebrate groups. widespread within Indonesia but tend to occur in localized The enormous species diversity within the Triangle becomes populations. Another important biogeographical revelation is apparent when compared with that found for the Caribbean that the faunistic relationships of the eastern sections of the Sea, the diversity centre for the Atlantic Ocean. In the Indian Ocean (west Australian coast, Christmas Island, Cocos- Caribbean, there are only c. 700 reef fish species (Rocha, Keeling Island) belong to the Western Pacific. 2003) and c. 50 coral species (Veron, 1995). The contrast Another way to appreciate the concentration of species in became more remarkable when Allen & Adrim (2003) reported the East Indies is to consider the results from two surveys of 1111 fish species from one restricted locality on the Indonesian coral reefs that extended all the way across the Indo-Pacific island of Flores, more than occur in the entire tropical Ocean. The first (Connolly et al., 2003) analysed the ranges of Atlantic! Even that number should not be considered definitive 727 coral species and 1766 fish species. The second (Mora for there is a cryptic fauna that cannot be included in a visual et al., 2003) examined the ranges of 1970 fish species. The census. This includes species that are well camouflaged as well results were similar in that latitudinal range midpoints clus- as those that hide in tiny crevices, beneath the sand, or within tered about on the equator, thus illustrating a ‘mid-domain’

1518 Journal of Biogeography 32, 1517–1522, ª 2005 Blackwell Publishing Ltd The marine East Indies effect (Colwell & Lees, 2000). And, in both surveys, the (Briggs, 1999b). When many disjunct patterns, representing longitudinal range midpoints clustered directly over the East various reef species are examined, it may be noted that in Indies, a location far to the west of the midpoint of the Indo- almost all cases a distributional gap appears in the central East Pacific Ocean. The anomalous position of the longitudinal Indies. Sometimes, the vacated areas may be very large with the peak was attributed by Connolly et al. to the effect of westward affected species occupying only the outermost fringes of the flowing currents driving the biodiversity pattern. On the other Indo-Pacific. In other instances, the gap may be small but will hand, Mora et al. concluded that speciation in and dispersal include the East Indies. Most of the disjunctions occur in a from the East Indies was responsible for the longitudinal peak, north–south direction and are termed ‘antiequatorial’ but and that the East Indies had played the major role in others may be found along an east–west axis. assembling communities throughout the Indian and Pacific Some of the disjunct species occur in patterns with related oceans. species so that a competitive replacement is suggested. For The conclusion of Mora et al. (2003) supported an earlier example, Fricke (1988) has shown that the fish species suggestion (Briggs, 1966) that the East Indies Triangle Diplogrammus xenicus (Jordan & Thompson) (Callionymidae) represented a centre of evolutionary origin. In its modern is antiequatorial and that the intervening area is occupied by version, the East Indies centre of origin hypothesis rests upon the closely related D. goramensis (Bleeker). Springer & evidence provided by six biogeographical patterns: (1) a Williams (1994) demonstrated that the genus Istiblennius distance-correlated decrease in species diversity (richness) that (Blennidae) included a monophyletic triad consisting of extends outward from the East Indies, (2) an average outward I. bellus (Gunther), I. zebra (Vaillant & Sauvage), and increase in geological age demonstrated by several genera and I. muelleri (Klunzinger). Istiblennius bellus exists in two widely species, (3) a pattern of historical dispersal tracks that radiate separated populations, one in the Indian Ocean and the other outward from the East Indies, (4) phylogenetic regression in the Western Pacific. Istiblennius muelleri occupies the East patterns indicating that the East Indies are occupied by the Indies while I. zebra is endemic to Hawaii. It these cases, it most apomorphic species, (5) evidence that some extinction appears that the dominant species occur in the East Indies and patterns begin in the East Indies and gradually move outward, may have displaced their siblings. These and other apparent and (6) indications from the mtDNA of some species that a fish replacement examples are illustrated in Briggs (1999b, Figs decreasing gradient of genetic diversity extends outward. 1–3 and 5). Literature references that substantiate the foregoing patterns In regard to invertebrates, Stanley & Newman (1980) gave and accounts of alternative theories may be found in Briggs several examples of relict distributions in the barnacle super- (1999a, 2004a). More recently, information on the phylogeny family Chthamaloides. This 70-Myr-old group is evidently of several fish families was interpreted (Briggs, 2004b) to being out-competed by the younger Balanoidea. Once con- substantiate evidence of an evolutionary flow that extended tinuous distributions in the former have become disjunct in outward from the diversity centre. This flow was considered to fringe areas where the balanoids are not yet dominant. be the marine equivalent to Wilson’s (1961) taxon cycle and A molluscan example provided by Cernohorsky (1976) is the Erwin’s (1979) taxon pulses. For the East Indies, this means gastropod Mitra chinensis (Gray) (Mitridae). Its antiequatorial that dominant species from the centre should displace older populations are separated by the closely related M. triplicata species that become extinct or are forced to retreat into deeper (Martens) that exists in the East Indies. water or farther out into the horizontal periphery. While disjunct patterns demonstrating an absence in the Critics of biogeographical theories that involve historical East Indies are common, it is generally not possible to identify dispersals often complain about the proposal of untestable a specific competitor that may have been responsible. But that hypotheses. However, if one wished to disprove the hypothesis location has a high diversity of predators, parasites and disease of an East Indies centre of origin, it may be done by finding organisms. Any of these factors could play a role in the evidence against the six patterns that provide its support; that extinction process. In regard to antitropical disjunctions, is, if one could demonstrate diversity gradients that increase eurybathic organisms may avoid the tropics by undergoing with distance from the centre, geological age gradients that isothermic submergence and therefore appearing to be decrease with distance, dispersal tracks that extend inward, disjunct. Some species have been able to transgress the tropics phylogenetic patterns with plesiomorphic species in the centre, during periods of cooler surface temperature, but most widespread extinction patterns that do not originate in the tropical disjuncts appear to have been caused by extinctions East Indies, or a gradient of increasing genetic diversity that (Briggs, 1995). If over time, older species in the East Indies extends outward, then the hypothesis may be called into begin to lose territory, become disjunct, and retreat to the question. edges of their ranges, some of them should still be found in distant places. In the tropical Indo-Pacific, short-range endemic species EXTINCTION AND REPLACEMENT tend to be concentrated around the periphery in places such as The competitive displacement of relatively old species appears the Red Sea, Western Indian Ocean, east and west Australia, to become evident when one considers geographical patterns isolated oceanic islands, and the Eastern Pacific. It has been that are probably caused by extinction and replacement pointed out (Briggs, 2002; Hughes et al., 2002) that these

Journal of Biogeography 32, 1517–1522, ª 2005 Blackwell Publishing Ltd 1519 J. C. Briggs populations of endemics are not the best conservation targets newest development in speciation research is the emphasis on because many are phylogenetic relicts that have accumulated ecological speciation or ecological genetics (Via, 2002). The over vast periods of time. Their small populations have low term ‘ecological speciation’, as used by most authors, is genetic variability and probably represent advanced stages in generally equivalent to the traditional concept of sympatric the extinction process. On the other hand, their very presence speciation. appears to corroborate an extinction pattern that began in the Recent studies, combining research on phylogeny and East Indies and progressed outward. As noted, there is also a ecology of natural populations, have cast new light on patterns, relatively small number of endemic species in Indonesian time-scales and mechanisms, and have emphasized the role of waters, but these are apparently newly formed, neo-endemics ecological factors in speciation. Studies of parallel speciation that do have a biogeographical potential. have provided a strong case for sympatric speciation and Although the East Indies Triangle has been recognized as the for natural selection in generating reproductive barriers predominant centre of origin for the marine tropics (Briggs, (Johannesson, 2001). Numerous examples exist for the marine 2003), this does not mean that it is the only place where environment (Hellberg & Vaquier, 1999; Hendry et al., 2000; successful evolution is taking place. An important secondary Ruber et al., 2003). They not only illustrate ecological (symp- centre is located in the Caribbean Sea, and parts of the Indo- atric) speciation but also show that this process can occur very Pacific that are separated by biogeographical barriers demon- rapidly. In the case of a salmon population that colonized strate independent evolutionary activity. For example, certain divergent reproductive environments, Hendry et al. showed fish families in the Eastern Pacific, such as the Sciaenidae, evidence of reproductive isolation after fewer than 13 genera- Gobiesocidae and Chaenopsidae, have undergone remarkable tions. In a pair of cryptic species of sea urchin inhabiting the radiations. The same is true for certain other families in the Indo-West Pacific, Landry et al. (2003) found that divergence in Western Indian Ocean. And warm-temperate provinces that sperm morphology and gamete recognition proteins had taken border the tropics to the north and south also demonstrate place within the past 250,000 years. In sympatric Drosophila considerable evolutionary independence. populations, speciation is completed in c. 200,000 years, but allopatric populations require c. 2.7 Myr (Coyne & Orr, 2004). It is interesting to note that the new emphasis on ecological SPECIATION speciation has resulted in a return to the Darwinian view that In the Indo-Pacific, successful evolution (the formation of speciation is the result of adaptation to different natural or dominant, widespread species) is apparently concentrated in sexual environments. In regard to the East Indies Triangle, the the East Indies. Why should this be so? Although most production of so many dominant species that have become published examples of centrifugal speciation had been con- widespread suggests an unusually effective speciation process. In cerned with evolutionary changes taking place among species addition, the high level of species diversity together with the within genera that occupied limited geographical areas, it was presence of local neo-endemics may indicate a relatively rapid proposed that this theory provided a much better model for the speciation rate. Both the evolution of dominant species and an operation of a centre of origin than does peripatric speciation increase in the rate of species production could be due to natural (Briggs, 2000). Both models are not so much concerned with selection operating in a competitive (sympatric) mode. There the process of speciation itself, but with the fate of the newly seems to be no other logical cause. Assuming that this is true, formed species. A review of the modes of speciation within the then the contrast between such dominant species and those Indo-West Pacific (Briggs, 1999c) indicated that all three basic generally produced elsewhere becomes apparent. Furthermore, types (allopatric, parapatric, sympatric) had been active. The the Darwinian view suggests that allopatrically formed species centrifugal mechanism emphasizes the evolutionary import- are accidental by-products of genetic divergence, while natural ance of relatively large, central populations. We know that selection directly favours the multiplication of species during genetic variation is driven mainly by population size and, the ecologically based sympatric speciation (Coyne & Orr, 2004). greater the genetic diversity, the greater the adaptive potential. As many systematic works have demonstrated, widespread There is experimental evidence for greater responses to geographical patterns, consisting of a mosaic of closely related selection in larger populations (Weber, 1990). And there is a species or subspecies, are often traceable to the previous highly significant correlation between heterozygosity and existence of a broadly distributed ancestral species. The population fitness (Reed & Frankham, 2003). presence of such mosaics, with their evidence of many Does the mode of speciation have anything to do with the allopatric (often vicariant) speciation events, has led many eventual success of the newly formed species? Until recent investigators to conclude that this mode has been predominant years, the allopatric process, involving either dispersal or or even exclusive in the marine environment. Granted that vicariance, had been considered the almost exclusive driving allopatry has probably been the most common, we can now force for evolution in the sea. Then, sympatric speciation was consider that it usually represents accidental speciation. In revealed to be quite common (Knowlton, 1993) and was noted allopatric speciation, the initial step of reproductive isolation to evidently occur in the East Indies (Briggs, 1999c). Parapatric does not involve natural selection and the subsequent speci- speciation may also be common but it has been difficult to ation process proceeds very slowly. In sympatric speciation, demonstrate in nature, except in pelagic species. Now, the reproductive isolation does involve selection and the speci-

1520 Journal of Biogeography 32, 1517–1522, ª 2005 Blackwell Publishing Ltd The marine East Indies ation process is apparently completed very rapidly. That is, fishes and corals of the East Indies. The enormous diversity in allopatric isolation involves a physical splitting of populations each group, and presumably for many other marine groups through the action of a barrier to gene flow, while sympatric that are less well known, far exceeds that of any other part of isolation requires the selection of an alternative habitat. the tropical seas. In fact, the number of reef fish species The proposition that successful speciation, i.e. the produc- observed in one locality, a bay on the Indonesian island of tion of dominant species that have the ability to spread out Flores, exceeds the number known from the entire tropical from their places of origin, usually results from natural Atlantic. Investigations of Indo-West Pacific palaeontological, selection, does not mean that allopatric or accidental speciation phylogenetic, genetic, and contemporary biogeographical pat- is invariably unsuccessful. In the event that two large popula- terns, along with a broad-scale survey of Indo-Pacific reefs, tions in a peripheral location have been separated by a barrier, showed that the East Indies was functioning as a centre of evolutionary adaptation would continue in each population but origin. It apparently has played the major role in assembling the initial impetus provided by natural selection would be reef communities throughout the Indian and Pacific oceans. missing. Eventually, over geological periods of time, peripheral The function of the East Indies as a centre of origin may be species formed by allopatry could evolve to become successful related to the mode of speciation taking place in that area. but, in the meantime they would be vulnerable to invasion by Recent studies on the phylogeny and ecology of natural better adapted species from an evolutionary centre. If the populations have emphasized the role of ecological factors and imposition of a barrier, or a dispersal across a barrier, results in provided a strong case for sympatric speciation. This mode the isolation of a small population, its prospects for long-term evidently occurs in the East Indies along with the allopatric and survival, despite speciation, are very poor. Small populations parapatric types. But certain characteristics of the East Indian may evolve rapidly but they possess relatively little genetic fauna such as the production of dominant, apomorphic variation and are subject to the debilitating effects of inbreeding species; the high species diversity; and the presence of and genetic drift. apparently newly formed species; suggest an unusually efficient speciation process. There is good evidence for the Darwinian view that natural selection involving sympatric speciation CONSERVATION rapidly produces better adapted species. On the other hand, it The world’s greatest diversity of marine animals may be found appears that allopatric speciation, usually the result of an among the coral reefs of the East Indies. Much of Malaysia and accidental or fortuitous interruption of gene flow, does not the Philippines support dense human populations and all of rapidly produce better adapted species. In the latter mode, their coral reefs have been degraded, many severely so. Aside natural selection is not involved in the initial step of from the reef structures, overfishing is evident in many areas so reproductive isolation. that artisanal and commercial fisheries are no longer produc- It is proposed that speciation by natural selection, probably tive. These reasons alone have been the causes of considerable involving the sympatric mode, is predominant in the East conservation concern. But, more important for the future is Indies, thus accounting for its evolutionary efficiency. In the the fact that the East Indies is an area of evolutionary origin vast peripheral areas of the Indo-Pacific, the allopatric process where dominant species are being produced. That is, the is probably predominant. This does not mean that the evolutionary process concentrated in the East Indies has allopatric process could not eventually give rise to a the function of supplying species that help to maintain the continuing lineage. In the long run, evolutionary change will diversity patterns in the rest of the Indo-Pacific. occur in any reproductively isolated population, but whether About 97 fish species have been identified that are confined to or not it is advantageous in a competitive sense will depend Indonesia (the inner part of the East Indies Triangle). These on factors such as natural selection, rapidity of speciation, species have been considered to be neo-endemics that were community diversity, and geographical location. The enor- recently derived and had yet to expand their ranges (Mora et al., mous diversity and evolutionary importance of the East 2003). Such species, and the other short-ranged neo-endemics Indies call for priority consideration in marine conservation belonging to the many invertebrate groups, represent the planning. immediate evolutionary and ecological future of the Indo- Pacific. In other words, they represent the next phase of a ACKNOWLEDGEMENT dynamic production system that has its origin in the East Indies. As they spread out, they will replace many of today’s common, I wish to thank Eila Hanni for her assistance with the widely distributed species. Future human generations will be manuscript. able to appreciate these new species and their descendants provided we protect significant portions of the East Indies. REFERENCES

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Briggs was named Professor Emeritus upon his fertilization selectivity and lysine cDNA sequences in tegu- retirement from the University of South Florida in 1990. His line gastropods. Molecular Biology and Evolution, 16, 839– research deals primarily with the origin and distribution of 848. contemporary groups of organisms. His biogeographical Hendry, A.P., Wenberg, J.K., Bentzen, P., Volk, E.C. & Quinn, books include Marine zoogeography, McGraw-Hill, 1974; T.P. (2000) Rapid evolution of reproductive isolation in the Biogeography and plate tectonics, Elsevier, 1987; and Global wild. Science, 290, 516–518. biogeography, Elsevier, 1995. In 2005, Professor Briggs received Hughes, T.P., Bellwood, D.R. & Connolly, S.R. (2002) Biodi- the Alfred Russel Wallace Award from the International versity hotspots, centres of endemicity, and the conservation Biogeography Society for his lifetime contributions to of coral reefs. Ecology Letters, 5, 775–784. biogeography. Johannesson, K. 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