Marine regions

3.3.2 Marine regions

Marine regions are somewhat more fluid and their relationships more obscure than freshwater re- gions. Although some workers have tried to place specific boundaries on marine areas, we provide only very general geographic regions because so little work has been done to determine their historical reality. Taxonomy, distribution and phylogenies are more poorly known for marine groups than for freshwater groups. The following is only a brief introduction to some of the marine areas and is in no way exhaustive or presented as

a definitive treatment. For example, we refer to McEachran and Fechhelm (1998) for the Gulf of Mexico. The emphasis here is on shorefishes which are those occurring from shore to a depth of 200 m; most pelagic groups are widely distributed, sometimes circumglobal, and deepsea families are usually poorly known both taxonomically and bio- geographically (but see Miya and Nishida 1997 for an interesting example). Nelson (1986) discussed

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56 Chapter 3

problems and prospects for historical biogeogra- phy of pelagic fishes.

The application of historical biogeographic techniques to marine fishes was slow to material- ize. Myers’ (1938) observation that primary fresh- water fishes are confined to particular drainage basins and will closely mirror geological history held the implication that marine fishes would have little to say concerning biogeography because of their capacity for dispersal. This dissuaded bio- geographers from seriously examining marine fish distributions for almost half a century. Myers’ notion of relative value of fishes for biogeography continues to have an impact on the choice of study taxa, despite the fact that taxon phylogeny and endemism are the final arbiters of biogeographical information content rather than little-known physiological tolerances and implied dispersal limitations (e.g. Dyer 1998, p. 528). With the identification of marine zoogeographic regions by Briggs (1974) and the recognition of areas with high endemism, wide dispersal could not be common to all groups. Springer’s (1982) study of the biogeogra- phy of shorefishes on the Pacific tectonic plate, the first major work on marine fishes to espouse

a modern approach, was the springboard that launched analytical marine fish biogeography. Hence, analytical marine biogeography is a very new science. Only over the last 15–20 years have the relatively restricted distributions of several groups, for example blennioids and pseudochro- mids, overcome some of the early hesitancy and lent credibility to the role of marine shorefishes in biogeographic studies.

Since Springer (1982), several authors have applied historical biogeographic interpretations to the distribution of marine shorefishes (e.g. White 1986, atherinids; Winterbottom 1986, pseudochromids; Springer 1988, Ecsenius, blen- niids; Williams 1988, Cirripectes, blenniids; Springer and Williams 1994, Istiblennius, blen- niids; Mooi 1995, plesiopids) and even to deep shelf forms (Harold 1998, sternoptychids). As in fresh- water studies, rigorous methods for comparison among cladograms has not generally been under- taken (see Lovejoy 1997 for an exception), but the search for common distribution patterns has been

useful in generating testable hypotheses for several regions (e.g. Nelson 1984 and White 1986, amphitropical/isthmian; Springer and Williams 1990, Indo-Pacific). Pattern congruence and phy- logeny will also undoubtedly help to solve the vex- ing problem of antitropical distributions, where sister species or populations are found in higher northern and southern latitudes separated by a broad equatorial distributional gap.

Some recent ecological work supports the bio- geographic evidence for restricted distributions and separate histories of some marine fishes. Swearer et al. (1999) and Jones et al. (1999) have shown, with different experimental protocols, that despite the production of planktonic eggs and larvae, offspring of some species remain close to their native shores or eventually settle back on to the reefs of their parents. The long-distance dispersal assumed for marine fishes is not univer- sal, and populations of even presently assumed widespread species are unlikely to be panmictic (Cowen et al. 2000; Pogson et al. 2001). Consistent with these studies, Gill (1999) has questioned marine fish species concepts and the reality of widespread species, suggesting that many of these are likely to comprise two or more allopatric cryptic species each with more restricted distribu- tions. These observations have clear taxonomic, systematic, biogeographic and conservation implications.

North Atlantic and Mediterranean

There are about 1200 species along the North American Atlantic coast including the northern Gulf of Mexico. Dominant families are the Serranidae and Gobiidae each with well over 50 species, although several families are represented by over 20 species (including Ophichthyidae, Muraenidae, Gadidae, Syngnathidae, Carangidae, Haemulidae, Sciaenidae, Scorpaenidae). It is estimated that about 600 species occur in the Mediterranean, with almost two-thirds of these considered shorefishes.

Humphries and Parenti (1999) use data from merluciids (hakes), Lophius (monkfishes) and two sponge groups to produce an area cladogram for

Historical Biogeography

(a)

Fig. 3.4 Cladistic biogeographic 5 3 analysis of real organisms from

4 3 the North Atlantic. (a) Areas of endemism for chalinid sponges that are similar to those of

northern species of the monkfish genus Lophius. 1, Arctic; 2, Boreal;

3, Mediterranean; 4, southeastern

North Atlantic; 5, western North Atlantic; 6, Caribbean. (b) Taxa and area cladogram of northern

(b)

(c)

Lophius species, with a line drawing of a typical member of the genus. Areas are numbered as in (a). (c) General area cladogram calculated from the combined data of Lophius and two chalinid sponge groups. (Source: after

4 SE North Atlantic Humphries and Parenti 1999,

4 Lophius vaillanti

6 Caribbean reproduced by permission of

6 L.gastrophysus

5 Western North Atlantic Oxford University Press. Lophius

5 L. americanus

3 Mediterranean outline from Nelson 1994;

3 L. budegassa

2 Boreal reproduced by permission of John

2 L. piscatorius

1 Arctic Wiley & Sons, Inc.)

1 L. litulon

the northern and tropical Atlantic (Fig. 3.4). They species for the tropical western Atlantic. This is found that the Mediterranean shared a more recent due in part to differences in defining the area, with faunal history with the Arctic and Boreal regions early workers considering it a single area and later than with any of the more tropical areas. The workers dividing it into the Caribbean including western North Atlantic (eastern USA), then coastal tropical Mexico and Central and South Caribbean and finally the southeastern North America, a West Indian region covering the Atlantic (western Africa, Azores, Canaries and Greater and Lesser Antilles, and a Brazilian region. Cape Verde Islands) are hypothesized as sequen- The latter has recently received attention result- tial outgroups to the Mediterranean and Arctic ing in recognition of additional endemic species regions. Too few corroborative patterns have and better understanding of its faunal history been suggested to imply plausible scenarios.

(Joyeux et al. 2001). Böhlke and Chaplin (1993) listed almost 600 species for the Bahamas. A com- prehensive recent accounting for Bermuda (Smith-

Tropical western Atlantic

Vaniz et al. 1999) lists 365 nearshore species for There are no accurate estimates of numbers of what is recognized to be a relatively depauperate

58 Chapter 3

fauna. Smith-Vaniz et al. (1999) also compares the composition of the Bermudan fauna with that of the Bahamas, Florida Keys and Carolinian Bight, comparing a total of over 800 species among these areas, which extend as far north as Cape Hatteras. However, this comparison was among selected families, so a total would be substantially higher for these areas and the Caribbean as a whole, perhaps as high as the 1500 species estimated by Lieske and Myers (1996). Dominant families with

30 or more species each include the Gobiidae, Serranidae, Labrisomidae, Apogonidae and Sciaenidae, although relatively strong representa- tion is found in the Labridae, Syngnathidae, Carangidae, Chaenopsidae and others with about

20 species each. Endemic families of the western Atlantic include the Grammatidae and Inermi- idae, but there is considerable endemism within most represented shorefish families. The Caribbean has received substantial attention by historical biogeographers. Rosen (1976, 1985) pro- vides plate tectonic models to explain biotic distri- butions. Rauchenberger (1989) and Lydeard et al. (1995) used freshwater fishes to hypothesize area relationships, but marine groups have played very little part in examining Caribbean biogeography. Page and Lydeard (1994) summarize the require- ments for advancing Caribbean biogeography. That the Caribbean and tropical eastern Pacific have a close biological connection with many amphi-American taxa has been known for well over a century. Briggs (1974, 1995) and Rosen (1976) provide summaries from differing perspec- tives on this connection. Fishes have provided numerous examples of amphi-American relationships including Atherinopsidae, Dactylo- scopidae, Labrisomidae, Chaenopsidae and Cen- tropomidae (sensu Mooi and Gill 1995), plus many others. Sister species and sister taxa at various supraspecific levels are known to be separated by the Isthmus of Panama. Kocher and Stepien (1997) provide some molecular examples, whereas Nelson (1984), White (1986) and Hastings and Springer (1994) provide morphological studies showing relationships among amphi-American and pan-Isthmian taxa.

Eastern North Pacific

The temperate to boreal areas of the North Pacific have a surprisingly diverse shorefish fauna. Along the North American coast alone, from California to Alaska, there are over 600 species in the top 200 m. The dominant families are in the Scorpaenoidei (sensu Mooi and Gill 1995), with scorpaenids (70 species), cottids (85 species) and agonids (over 20 species) making up most of these. The liparidids comprise about 15 species in water less than 200 m deep, the remaining 35 species being found at depths between 200 and 7600 m. Other dominant families of the eastern North Pacific include the zoarcids (40–50 species) although most are found below 50 m, stichaeids (25 species), pleuronectids (22 species) and em- biotocids (20 species). The latter family is endemic to the North Pacific and is unusual in having inter- nal fertilization and true viviparity. Various other taxa suggest a close relationship for North Pacific taxa. For example, additional North Pacific endemic families include the gasterosteiform aulorhynchids, the scorpaenoid Anoplopomatidae (two species) and Hexagrammidae (over a dozen species). Other North Pacific taxa of note are, of course, the Salmonidae with about 11 species, the most famous of which are anadromous and die after impressive migrations upriver to spawn, and the Pacific halibut (Hippoglossus stenolepis) grow- ing to over 2.5 m and 360 kg.

Tropical eastern Pacific

There are about 750 species in the tropical eastern Pacific, which includes the Gulf of California to Ecuador and offshore islands, and over 80% of these are endemic (Allen and Robertson 1994). The dominant families are the Sciaenidae (80 species) and Gobiidae (80 species), with blennioids also forming a substantial part of the shorefish fauna with Chaenopsidae (30 species), Dactyloscopidae (24 species) and Labrisomidae (25 species). Other well-represented families include the Serranidae, Haemulidae and Labridae, all of which have about

30 species, and Ariidae, Muraenidae and Pomacen- tridae that each have about 20 species. All recorded 30 species, and Ariidae, Muraenidae and Pomacen- tridae that each have about 20 species. All recorded

Tropical Indo-Pacific

Dividing this huge area, which includes the Red Sea and east coast of Africa to Hawaii and Easter Island, into identifiable subregions is difficult. Essentially all parts of this region require substan- tial sampling to produce a clear picture of the fauna. In addition relationships among the fishes remain obscure, and species limits and distribu- tions are frequently poorly delimited. Springer (1982) made an estimate of 4000 species for this area, but this is likely to be too low. The following discussion of species numbers and endemism fol- lows Randall (1998) and references therein.

The highest number of species is found in the East Indian region, which includes Indonesia, New Guinea and the Philippines, This region is esti- mated to be home to 2800 species of shorefishes. Because the region is so large, it is misleading to provide levels of endemism because many fish are restricted to smaller subregions. The diversity in this area is probably a result of its complex geologi- cal history. Several islands, including Sulawesi, New Guinea and others, are known to be compo- sites with smaller pieces having accreted during a basically counterclockwise rotation of the Sahul Shelf into the Sunda Shelf over the last 30 million years, with significant tectonic change even over the last 3–5 million years (Burrett et al. 1991; Hall 1996, 1998; Moss and Wilson 1998). Significant fluctuations in sea level during glacial cycles are also hypothesized to have resulted in speciation. Shorefishes that appear to provide examples of patterns consistent with these geological models are discussed by Winterbottom (1986), Woodland (1986), Springer (1988), Springer and Williams (1990, 1994) and Springer and Larson (1996). The region also exhibits an incredible variety of habi- tats, including silicate and volcanic sands, muds, mangroves, estuaries, lagoons and protected

and exposed reefs. The fauna is dominated by the perciforms, especially the Gobiidae (200+),

Labridae (100+), Pomacentridae (100+), Serrani- dae (about 100) and Blenniidae (about 100). Other common and conspicuous families are the Apogonidae (about 80), Pseudochromidae (about 60), Chaetodontidae (about 50) and Acanthuri- dae (about 45). The only non-perciform families with 50 or more species are the Muraenidae and Syngnathidae.

Numbers of species in the Pacific decrease away from the East Indian region. To the north, Taiwan is estimated to have about 20 species, with al- most 14% of these not found to the south in the East Indian region. The Ryukyu Islands probably have about 2000 species. Almost 1600 species are expected to be found in Micronesia, but numbers decrease eastward through the archipelago (Palau 1400, Carolines 1100, Marianas 920, Marshalls and Gilberts 850). Myers (1989) provided an excel- lent review of the geography, geology and ecology of Micronesia and its shorefishes. Springer (1982) referred to Micronesia as the Caroline Islands con- duit, forming a potential set of stepping stones on to the Pacific plate and discussed the biogeography of the Pacific in general. The oceanic island groups, of course, offer fewer habitats and have a less diverse fauna. However, endemism is often high. Hawaii has about 560 species, with over 130 not found elsewhere. The Great Barrier Reef of eastern Australia and New Caledonia have the greatest diversity of the South Pacific, both with about 1600 species. Randall et al. (1998) provide the only comprehensive summary of Great Barrier Reef fishes. To the east of New Caledonia, numbers of shore species decrease to about 900 in Samoa, 630 in the Society Islands, 260 in Rapa and 126 in Easter Island (Randall 1998). Highest endemism in the South Pacific is found at Easter Island, with 22% of its species found nowhere else. The Marquesas Islands have 10% endemism, but most island groups exhibit about 5%. Briggs (1995, 1999a) has attributed these diversity differences to centre of origin hypotheses and competitive exclusion. However, such explanations have been discredited and replaced by allopatric speciation models incorporating plate tectonics, sea-level

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fluctuations and other geological mechanisms and parameters such as island age (e.g. Springer and Williams 1990, 1994). Woodland (1986) suggested that Wallace’s Line, usually reserved as a boundary for terrestrial taxa (see section on Southern Asia), might be descriptive for some marine fish groups, and Barber et al. (2000) have found sharp genetic breaks for stomatopods in this same area.

West of the East Indian region, numbers of species are surprisingly similar among localities. The Chagos Archipelago has 784 reported species but is expected to approach the total of the Maldives at about 950 species (Winterbottom and Anderson 1999). Almost 900 species are reported from the Seychelles, a substantial underestimate of the Mauritius shorefish fauna sets its total at 670 species, and over 900 species are recorded for Oman (Randall 1998). At least 1160 species occur off the coast of southeast Africa and the Red Sea has a similar number of species but exhibits a much higher level of endemism at about 14%, although some families have much higher levels, for example almost 90% in pseudochromids. Unfortunately, there are no recent summaries of the fauna of India, although Anderson (1996) estimated over 1000 for Sri Lanka alone. Several authors have proposed that disjunct distributions among sister taxa across the Indian Ocean is a result of vicariance imposed by the movement of India northwards towards Asia (Winterbottom 1986; Hocutt 1987; Springer 1988; Mooi 1995). Briggs (1990) criticized these hypotheses based mostly on the proposed age of taxa, but Patterson and Owen (1991) effectively countered these argu- ments. Within the Indo-Pacific region, the western Indian Ocean exhibits considerable endemism. Several taxa, such as butterflyfish species pairs and pseudochromid genera, have sister-group clades with one member in the western Indian Ocean and the other in the eastern Indian Ocean/western Pacific Ocean (Winterbottom 1986; Gill and Edwards 1999).

Antitropical distributions

Most distributions do not fit classical regions because the regions themselves do not have a his-

torical basis. Particularly striking examples are antitropical (antiequatorial) distributions, which are distributions of the same or closely related taxa that occur in southern and northern subtropical or temperate regions with an intervening equatorial disjunction. Briggs (1999a,b) continued to argue that such distributions are a result of competitive exclusion of newer taxa over older taxa. White (1989) succinctly discussed the numerous prob- lems with Briggs’ approach and noted several logical alternative hypotheses. As with many biogeographic questions, the solution lies in the recognition of pattern. If the antitropical distribu- tions are truly congruent in time and space, then a common explanation can logically be sought and tested. However, if each antitropical distribution is unique, then any explanation of each pattern will also be unique and will remain speculative. White (1986, 1989) hypothesized a common process of global Eocene/Oligocene cooling followed by a mid-Miocene equatorial warming event as the main factor in shaping the repeating patterns of antitropical distributions of taxa, us- ing atherinopsines (silversides) as a model. Humphries and Parenti (1986, pp. 84–6) described a novel explanation for antitropical patterns involv- ing a north/south division of a pre-Pangaean conti- nent called Pacifica. Nelson (1986, p. 216) provided an example from the Engraulidae that, if not sup- porting the Pacifica model, suggested taxonomic differentiation in a growing rather than shrinking Pacific Basin. Geological explanations that are not mainstream have been dismissed by some (Hallam 1994; Holloway and Hall 1998), but mainstream geologists do not have a stranglehold on truth; plate tectonics itself was dismissed as radical until recently.

Temperate Indo-Pacific (South Africa, southern Australia, New Zealand)

Although not recognized as a traditional region, this area deserves special mention because of the high diversity and endemism exhibited and its faunal connection. In total, this area is home to at least 2100 shorefishes, with about 900 of these found nowhere else. South Africa has over 20 Although not recognized as a traditional region, this area deserves special mention because of the high diversity and endemism exhibited and its faunal connection. In total, this area is home to at least 2100 shorefishes, with about 900 of these found nowhere else. South Africa has over 20

Wilson and Allen (1987) estimated that of 600 inshore species for southern Australian temperate seas an incredible 85% are endemic, far outstrip- ping levels of endemism in any tropical region (peak of 23% in Hawaii). Families with 20 or more endemics are Clinidae, Gobiesocidae, Gobiidae, Labridae, Monacanthidae and Syngnathidae, which includes the famous seadragons. These are also the most speciose families, making up over one-quarter of inshore species. Other inshore families with five or more endemic species include Apogonidae, Antennariidae, Atherinidae, Callionymidae, Ostraciidae, Platycephalidae, Plesiopidae, Pleuronectidae, Rajidae, Serranidae, Scorpaenidae, Soleidae, Tetraodontidae, Urano- scopidae and Urolophidae. Gomon et al. (1994) listed 730 species for the southern coast of Australia, although the inclusion of species from the pelagic and deepsea realms and geographic coverage were arbitrary. Even from this source, southern Australia boasts well over 400 endemic species representing 100 families and over 60% endemism. Neira et al. (1998) estimated 1500 temperate Australian species including deepsea and oceanic taxa.

Paulin et al. (1989) listed just over 1000 species for New Zealand and adjacent islands, with 110 endemic species (11%). Paulin and Roberts (1993) split the New Zealand fauna into ‘rockpool’ and ‘non-rockpool’ fishes and determined that the 58 of the 94 species in the former group were endemic whereas 52 of the remaining 900+ fishes were endemic. Dominant rockpool endemic groups are Tripterygiidae (20 endemics), Gobiesocidae (9 endemics), Syngnathidae and Plesiopidae (four endemics each). Of non-rockpool species, domi- nant endemic families are the Galaxiidae (12 endemics; spawns in fresh water but at least part of the life cycle is marine), Macrouridae (about 12 endemics) and Pleuronectidae (eight endemics).

These localities have a surprising number of

taxa in common. Cool temperate New Zealand and Australia share over 80 species (in over 40 families) that are found nowhere else; an addi- tional 70 or more species are shared by warm tem- perate environments in these areas (Paulin et al. 1989). Families endemic to this region include Ar- ripidae (four species) and Chironemidae (four species). An additional 50 or more species are shared with Australia, South Africa and southern South America to the exclusion of other regions. Some of these latter species are from deep water and their apparent restricted distributions might

be due to poor sampling. However, families such as the Aplodactylidae are found only in Australia, New Zealand, Peru and Chile. Detailed phyloge- netic studies are necessary to test whether some marine fish distributions might support patterns shown by freshwater fishes that indicate a shared history among Australian, South American and African taxa and the implied Gondwanan origin.

Antarctic

The Antarctic Ocean is a well-defined body of water that is bounded by the continent of Antarctica to the south and by the Antarctic convergence, or polar front, which encircles the continent at 50–55° S and is where cold Antarctic surface water (2–6 °C) meets warmer Southern Ocean water (8–10 °C). Miller (1993) listed almost 300 species for the region, although over one-third of these are not shorefishes (they are pelagic or deepwater species). As would be expected, the fish fauna is dominated by those that can withstand ex- ceedingly low temperatures. The Notothenioidei make up well over one-third of the total fauna and over half of the shorefishes, with Nototheniidae (over 40 species), Artedidraconidae (over 20 species), Channichthyidae (about 20 species), Bathydraconidae (16 species), Harpagiferidae (eight species) and Bovichthyidae (one species). Most of these species are endemic to the Antarctic and, given that some species live in water with an average temperature of about -2 °C, exhibit some incredible adaptations: they have special glycopro- teins in the blood that lower its freezing point or lack haemoglobin altogether in the cold oxygen-

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rich waters. This suborder is mostly benthic and more on geological hypotheses and environmental without air bladders, but some species have en- and dispersal assumptions (Holloway and Hall tered the water column and have evolved lipid- 1998; Avise 2000); some welcome this shift to a filled sacs that help to provide neutral bouyancy. process/modelling approach (Heaney 1999) where- Bargelloni et al. (2000) hypothesized a phylogeny as others do not (Humphries 2000). Although an based on mitochondrial DNA; a candidate for sis- integration of the two paradigms (process and ter taxon has yet to be established. The Liparidae pattern) will advance the field, process narratives (about 40 species) and Zoarcidae (about 25 species) should be incorporated cautiously. Repeated his- are also common faunal elements but are mostly torical patterns that can form the basis of robust found deeper than 100 m, with some deeper than mechanistic explanations are only very slowly 7000 m.

coming to light but we should be patient in waiting for them to avoid the shortcomings of previous eras in biogeography.

Historical biogeography in ichthyology is a rela- tively new science and, with few exceptions (e.g. Although biogeographers have been assembling North America), quality data for pattern recogni- taxon and area cladograms for about 25 years, there tion remain scarce; opportunities to make con- have been relatively few successful attempts to tributions to the field abound. Future work in compare and evaluate potential congruence. New biogeography will continue to rely on excellence methods in biogeography are continually devel- in taxonomy and phylogenetics, basic field data for oped, and choosing which advance the field is complete distributional records, and a method difficult. Morrone and Crisci (1995, pp. 392–4) to identify congruence among area cladograms. advocated an integrated approach that can take There are areas of the world for which a synthesis advantage of the merits of each method; such an of phylogenies using rigorous techniques would approach is perhaps as likely to incorporate their help direct future ichthyological work, particu- various disadvantages. Despite the number of larly Neotropical and Australian fresh waters and methods, there is consensus for reliance on the Indo-Pacific. phylogeny reconstruction, use of some form of area cladogram, and increased use of geology and palaeontology. A continuing serious challenge to

3.4 CONCLUSIONS

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Part 2

Production and Population Structure