FISH, BIODIVERSITY OF

Gene S. Helfman University of Georgia

I. Taxonomic Diversity speciose Of high diversity, having many species in a II. Geographic Diversity group or area. -taxa) A group of evolutionarily re ؍ taxon (plural lated species.

GLOSSARY A FISH IS AN AQUATIC VERTEBRATE that (usually) adipose fin A small, fleshy fin without supporting has scales, fins, and gills (Box 1 and Fig. 1). Fishes spines or rays, set far back on the dorsal surface of constitute more than 25,000 of the known 48,000 many catfishes, characins, salmons, and other species of living vertebrates and are divided taxonomi- groups. cally into three major groupings: jawless fishes (agna- ancestral The taxon from which descendant species thans), cartilaginous fishes (chondrichthyans), and are derived, often synonymized as primitive or gener- bony fishes (osteichthyans). Fishes are found almost alized. Ancestral traits or conditions are those which everywhere on Earth where water of reasonable in- appear in an ancestor. tegrity exists. Fish habitats include the deep sea to depauperate Of low diversity, lacking in species; oppo- depths of 8000 m, high mountain streams and lakes to site of speciose. 5000 m altitude, and just about every aquatic habitat derived Later-appearing taxa within a lineage, often in between. Marine fishes make up 58% of all species, synonymized as advanced or specialized. Derived freshwater species make up 41%, and 1% of fishes move traits are those which appear in a descendant species regularly between the ocean and fresh water. Tropical and are changed from the ancestral condition. areas have the highest diversity. Fish biodiversity is endemic Restricted or native to a geographically de- threatened by a wide range of human activities, but fined area. habitat modification, overharvest, and introduced spe- extant Living; opposite of extinct. cies are particularly injurious. extirpated Locally extinct. Gondwana A large supercontinent in what is now the Southern Hemisphere that separated during the Mes- ozoic, forming the modern continents of Australia, I. TAXONOMIC DIVERSITY Antarctica, South America, and Africa. species flock A large group of closely related, endemic A. Overview species in a small area; all are descended from a Approximately 25,000 known fish species inhabit the single ancestor. earth’s oceans, estuaries, rivers, lakes, and streams. This

Encyclopedia of Biodiversity, Volume 2 Copyright  2001 by Academic Press. All rights of reproduction in any form reserved. 755 756 FISH, BIODIVERSITY OF

Box 1 What Is a Fish and What Are Fishes? Generally defined, a fish is a cold-blooded, aquatic-living chordate with fin-like appendages, a body covered with scales, and that breathes using gills (Fig. 1). Exceptions to and variations in all traits are common. Some sharks, tunas, and billfishes are warm-blooded. Fins can be unseg- mented and spiny or segmented and soft rayed (and soft rays can be hardened). Tail types include FIGURE 1 Anatomical features of a hypothetical fish, including some (i) the heterocercal tail of sharks and sturgeons, common features that are measured during fish identification. 1, in which the notochord or vertebral axis extends premaxilla; 2, maxilla; 3, dentary; 4, barbel; 5, snout; 6, nostril; 7, considerably into the upper tail lobe; (ii) the ab- preopercle; 8, branchiostegals; 9, opercle; 10, opercular spine; 11, breviate heterocercal tail of gars and bowfin, in pectoral fin; 12, lateral line; 13, first part of (or spinous) dorsal fin; 14, second part of (or soft) dorsal fin; 15, adipose fin; 16, caudal fin; which the vertebral axis extends only slightly into 17, caudal peduncle; 18, lateral scutes; 19, anal fin; 20, pelvic or the upper lobe; (iii) the leptocercal or diphycercal ventral fin (reproduced with permission from Greenfield and Thomer- tail of lungfishes, the coelacanth, and rattails, in son, 1997). which vertebrae or tail rays extend through the middle of the tail, forming a pointed tail; and (iv) the homocercal tail of most advanced bony fishes, in which the vertebral column ends at the tail that evolved relatively recently (derived or advanced base (the urostyle) and fin rays form a symmetri- taxa) placed later in lists. cal, two-lobed tail. Scales also vary in terms of the The most primitive of the living fishes are the jawless number of layers of bony material that constitute fishes. These arguably include 22 species of marine them and the extent of spiny projections that lancelets, and less arguably about 85 species of marine cover their surface; more primitive fishes gener- hagfishes and freshwater or anadromous (migratory be- ally have heavier scales, and more advanced fishes tween freshwater and marine) lampreys. Cartilaginous have lighter scales, often with more projections. sharks, skates, rays, and chimaeras include approxi- Scale types include the placoid scales of sharks, mately 10 orders, 43 families, and 850 species of almost the ganoid scales of gars and bichirs (and stur- entirely marine, relatively large-bodied predators. geons), the cycloid scales of lower teleosts, and Skates and rays are more diverse than sharks, constitut- the ctenoid scales of higher teleosts. ing about 55% of all cartilaginous fishes. Chimaeras To ichthyologists, ‘‘fish’’ refers to one or more are cartilaginous fishes distantly related to sharks that individuals of a single species, whereas ‘‘fishes’’ consist of 3 families and 31 species. refers to more than one species, regardless of how Bony fishes make up the vast majority of living fish many individuals are involved. Hence, this article species, exceeding all other groups in species, habitat, is about fishes. reproductive, and feeding diversity. Bony fishes vary in length from the 8-mm pygmy gobies to the 12-m long oarfish, 900-kg marlin, and 1000-kg-plus ocean sunfish. There are perhaps 45 orders and 435 families among the approximately 24,000 species of bony fishes, rang- incredible diversity exceeds that of all other vertebrate ing from the relatively primitive lungfishes, coelacanth, groups combined. It forms a wealth of biological won- sturgeons, bichirs, gars, and bowfin to the more ad- der for ichthyologists (fish scientists), but such large vanced teleostean (higher bony fish) groups that in- numbers can be overwhelming to someone unfamiliar clude bonytongues, eels and tarpons, herring-like fishes with the many taxonomic groups and their names. How- and the so-called true teleostean groups of minnows, ever, the different taxonomic groups are logically ar- salmons, various deep-sea taxa, and cods. The spiny- ranged according to well-studied evolutionary relation- rayed teleosts are the most evolutionarily advanced of ships, with those more closely related to groups that the fishes and include mullets, silversides, scorpion evolved earlier in geologic time (so-called ancestral or fishes, perch-like fishes, tunas, flatfishes, and trig- primitive taxa) placed earlier in lists and those groups gerfishes. FISH, BIODIVERSITY OF 757

Phylum Chordata Hagfishes occur almost worldwide in temperate and Subphylum Cephalochordata; order Amphioxiformes cold-temperate oceans, usually in water deeper than 30 (lancelets, two families, 22 species, marine, tropical m. Hagfishes can reach high densities, upwards of 0.5/ and temperate) m2, on soft-bottom marine areas, which is the most Subphylum Vertebrata abundant habitat type in the world. Hence, hagfishes Superclass Agnatha could be ecologically important as predators and scav- Class Myxini; order Myxiniformes (living hagfishes; one engers. They are also common in the diets of seals family, 43 species, temperate marine) and sea lions. Hagfishes are commercially important Class Cephalaspidomorphi; order Petromyzontiformes because their hides are the popular ‘‘eelskin’’ of wallets, (living lampreys; one family, 41 species, temperate purses, and briefcases. Overfishing has depleted hagfish fresh water and anadromous) stocks in Korea and Japan, and new fisheries are being exploited, and probably overexploited, in the eastern B. Cephalochordates Pacific and western Atlantic. This unfortunate chain of events has characterized marine fisheries worldwide. Lancelets may not be fishes because they lack scales, Drastic reductions in hagfish populations brought on fins, and gills. However, their evolutionary and anatom- by overfishing could potentially disrupt a widespread ical affinities are similar to what most workers believe ecosystem; not enough is known about hagfish ecology characterized the ancestors of fishes, and lancelets are to predict these impacts. studied primarily by ichthyologists. Lancelets possess a dorsal nerve tube, in common with all vertebrates. 2. Lampreys They also have a notochord, which is a cartilaginous Lampreys also have a notochord rather than vertebrae. rod that runs the length of the body and is shared with Lampreys go through a long-lived larval phase; the free- all embryonic vertebrates as well as with the adults of living, blind, toothless larva lives in silty stream-beds many primitive living fishes. Lancelets are small (to 3 in which it filters microscopic organisms from the water cm), slender organisms that as adults occupy sandy, for up to 7 years before transforming into an adult. usually shallow bottoms in all major tropical and tem- In brook lampreys, larvae transform into non-feeding perate oceans. A commercial fishery for lancelets involv- adults, live for 6 months, spawn, and die. Other species ing bottom dredging exists in southern China. The transform into feeding adults and live for 1–3 years as fishery is in apparent decline. parasites on other fishes. They rasp holes in their host’s skin and live off its body fluids. Accidental introduction C. Agnathans of the parasitic marine lamprey into the Great Lakes of North America has contributed to the decline of lake The first fishes lacked jaws. Modern jawless fishes— trout, whitefishes, and blue pike. hagfishes and lampreys—look approximately similar, Lampreys are cool-water species (30Њ north and with slippery, eel-like bodies and jawless heads. The south latitude or higher). Most lampreys live in fresh fossil record, however, indicates that they have been water, but some parasitic species are anadromous. separated evolutionarily for hundreds of millions of Brook lampreys typically live in headwater streams, an years. Hence, most similarities are due to convergent ecosystem type frequently disrupted by human activi- or parallel evolution. ties. Hence, several U.S. brook lampreys are imperiled. North America’s smallest lamprey, the Miller Lake lam- 1. Hagfishes prey, was poisoned into extinction because it parasit- Hagfishes, known also as slime eels or slime hags, pro- ized introduced trout in its only habitat—Miller Lake, duce copious mucus from many pairs of slime glands. Oregon (Fig. 2). A disturbed 2-ft-long hagfish can fill a 5-gallon bucket with slime. However, a hagfish covered in its own slime will suffocate. To rid itself of slime, a hagfish ties a D. Cartilaginous Fishes knot in its tail and passes the knot forward until the Elasmobranchs (sharks, skates, and rays) include about slime is pushed off. Hagfishes are nocturnal predators 800 species that live in all the world’s oceans; a few on small invertebrates but are better known for their live in fresh water. Elasmobranchs are characterized by scavenging behavior, which involves burrowing into a cartilaginous skeleton hardened by calcium deposits a dead or dying fish and consuming the prey from and usually five (sometimes six or seven) gill slits. They the inside. lack lungs or gas bladders but instead have large, oil- 758 FISH, BIODIVERSITY OF

FIGURE 2 The extinct Miller’s Lake lamprey (reproduced with permission from Miller et al., 1989).

filled livers which may aid in buoyancy. Their teeth are primarily offshore, pelagic inhabitants. The squali- and pedestal-like placoid scales develop from the same form dogfishes, the second largest shark order, are most embryonic structures. Teeth are continually lost and successful and abundant in the North Atlantic, North replaced, and a shark may produce as many as 30,000 Pacific, and deep-sea regions. teeth during its lifetime. All elasmobranchs have inter- Sharks range in size from the 15-g, 16- to 20-cm nal fertilization, and many bear live young that are dwarf dogshark to the 12,000ϩ -kg, 12ϩ -m long whale nourished by the mother via a complex umbilicus and shark, the largest fish in the world. White sharks as placental structure analogous to that found in mam- large as 6 m and 3324 kg are known; larger individuals mals. Slow growth, late maturation, and low reproduc- are suspected. Other large sharks include basking tive output make many elasmobranchs exceptionally sharks (9.8 m), great hammerheads (5.5 m), Greenland vulnerable to human exploitation. sharks (6.4 m), tiger sharks (5.9 m), and megamouth (5.4 m). Class Chondrichthyes: cartilaginous fishes Sharks inhabit all oceans except the Antarctic. Depth Subclass Elasmobranchii: shark-like fishes records for sharks are held by the Portuguese shark at Superorder Euselachii: modern sharks and rays 3690 m and an unidentified dogfish at 4050 m. A few Order Heterodontiformes: 1 family, 8 species, bullhead carcharhinid sharks enter fresh water; bull sharks have and horn sharks been captured 4200 km up the Amazon River and 1200 Order Orectolobiformes: 7 families, 31 species, includ- km up the Mississippi River. Large pelagic sharks may ing wobbegongs, nurse, and whale sharks cross entire ocean basins; blue sharks have been tracked Order Carcharhiniformes: 8 families, 210 species, in- across the North Atlantic Ocean and back, a distance cluding catsharks, requiem, and hammerhead sharks of 16,000 km. Order Lamniformes: 7 families, 16 species, including Most sharks are predatory on large prey, but three sand tiger, megamouth, thresher, basking, and mack- of the largest sharks—the basking, megamouth, and erel sharks whale sharks—feed on zooplankton. A small, 40-cm- Order Hexanchiformes: 2 families, 5 species, frill and long, midwater species, the cookie-cutter shark, is an cow sharks ectoparasite on the sides of tunas, dolphins, whales, an Order Squaliformes: 4 families, 74 species, including occasional megamouth shark, and even rubber sonar sleeper and dogfish sharks domes of nuclear submarines. Some sharks use struc- Order Squatiniformes: 1 family, 12 species, angel sharks tures other than jaw teeth to capture prey. Thresher Order Pristiophoriformes: 1 family, 5 species, saw- sharks use the long upper lobe of their tails to stun sharks schooling prey. Sawsharks (and rajiform sawfishes) Order Rajiformes: 13 families, 456 species, including have elongate, blade-like snouts studded with lateral sawfishes, electric rays, guitarfishes, skates, sting- teeth which they slash laterally to disable prey. Ham- rays, eagle rays, and manta rays merheads use their broadened hammer-shaped head to pin stingrays against the bottom before biting chunks out of the rays’ wings. 1. Sharks Sharks are sensitive to chemicals, able to detect 1 Approximately 350 shark-like species are alive today part fish extract per 10 billion parts seawater. Sharks (Fig. 3A). Sharks are generally large (Ͼ1 m), predatory have good vision, although they tend to be slightly fishes. The diverse requiem or ground sharks (carchar- myopic (farsighted). Sharks are also highly sensitive to hiniforms) include the tiger, gray reef, bull, blue, lemon, sounds, including infrasonic sound below 10 Hz, and and hammerhead sharks. Lamniform mackerel sharks can localize the direction from which underwater FISH, BIODIVERSITY OF 759

FIGURE 3 Taxonomic distribution and representative orders of (A) sharks and (B) skates and rays (modified with permission from Compagno, 1990). sounds originate. Sharks can also locate prey by de- 70–100 years and lemon sharks 50–60 years. Sharks tecting the weak electric fields prey emit. This electro- also reproduce slowly. Lemon sharks may not mature sensitivity may also allow them to navigate using the until they are 24 years old, and spiny dogfish may not earth’s geomagnetic fields. Many sharks have relatively mature until they are 35 years old. Sharks produce large brains, with brain to body weight ratios compara- relatively few, large young with a long gestation period. ble to those of some birds and mammals. Clutch size varies from 1 or 2 live young (sand tigers, Sharks grow slowly and live long: spiny dogfish live threshers, and makos) to 300 in the whale shark. Gesta- 760 FISH, BIODIVERSITY OF tion periods average 9–12 months but may be 2 years 3. Chimaeras in spiny dogfish and 3.5 years in basking sharks. Bull- head and nurse sharks lay eggs, but most sharks bear Class Chondrichthyes live young. The requiem sharks have the most advanced Subclass Holocephali developmental pattern, in which an umbilical cord con- Order Chimaeriformes: 3 families, 31 species, chi- nects mother and embryo, transporting nutrients and maeras oxygen to the embryo and carrying metabolic wastes to the mother. Chimaeras, also known as rat- or rabbitfishes, share a Because many sharks mature late, reproduce at long intervals, and have low reproductive output, shark pop- cartilaginous skeleton and other features with elasmo- ulations are easily and frequently overfished. Thresher branchs. They differ by having (i) the upper jaw perma- sharks, school sharks, spiny dogfish, porbeagles, bask- nently attached to the braincase, (ii) continually grow- ing sharks, bull sharks, and soupfin sharks are all ex- ing tooth plates in the jaws instead of replaceable teeth, amples of shark stocks that have been overexploited. (iii) a single gill flap instead of five or more gill slits, White sharks are protected in Australia and South Af- and (iv) no scales. Chimaeras swim by flapping their rica; white sharks and whale sharks are listed as endan- pectoral fins and by undulating their bodies. All chimae- gered by the International Union for the Conservation ras are egg-layers, the egg being protected by a horny of Nature. shell. Adult chimaeras range in size from 60 to 200 cm. Chimaeras are cool-water, marine fishes that live in shallow to moderate depths between 80 and 2600 m, 2. Skates and Rays where they usually swim just above the bottom. Chi- maeras eat predominantly hard-bodied benthic inverte- The rajiform skates and rays are 450 species of mostly brates, which they crush with their tooth plates. Sur- benthic (bottom-living), mostly marine forms (Fig. 3B). prisingly little is known about their general biology and In skates and rays, the pectoral fins are fused to the natural history. sides of the head and the five gill slits are under the head. Skates are most diverse in deep water and at high latitudes, whereas stingrays are most diverse in tropical, E. Bony Fishes inshore waters. Some rajiforms live much or all of their lives in fresh water. Largetooth sawfish frequently swim Modern bony fishes, often referred to as Osteichthyes up rivers in Central and South America. Two stingray (literally ‘‘bony fishes’’), consist of seven major taxo- families contain entirely freshwater species—the river nomic groups. The first six, primitive taxa belong to stingrays of South America and several species in the groups that were much more diverse during Paleozoic large stingray family Dasyatidae. The latter inhabit Afri- and Mesozoic eras. Several of these primitive but mod- can, Southeast Asian, and New Guinea rivers. ern fishes are classified as ‘‘bony’’ even though they Skates and rays feed mostly on benthic invertebrates, have cartilaginous skeletons. Their skeletal condition is except for the huge (up to 6-m wide) manta rays, which actually an advanced, specialized trait; their immediate capture small crustaceans and fishes in the water ancestors were bony. column. Torpedo rays stun prey with powerful electri- cal discharges (50 V and 50 A ϭ 1 kW output). Raji- Grade Teleostomi (more commonly Osteichthyes) forms reproduce by either laying eggs (skates) or Class Sarcopterygii bearing live young (rays). Embryonic skates develop Sub- (or infra-) class Dipnoi inside the ‘‘mermaid purse’’ egg cases for as much as Order Ceratodontiformes: 1 family and 1 species, the 15 weeks. Australian lungfish Skates in some locales are actually increasing in num- Order Lepidosireniformes: 2 families, 5 species, South ber because of overexploitation of competing bony American and African lungfishes fishes, such as cod in the North Atlantic. However, the Subclass Coelacanthimorpha giant barndoor skate of the northwest Atlantic and its Order Coelacanthiformes: 1 family and 2 species, the relative, the common skate of the northeast Atlantic, coelacanths are caught incidental to bottom trawling for bony fishes; Class Actinopterygii they have been seriously depleted and may face extinc- Subclass Chondrostei tion. Largetooth sawfish in Nicaraguan lakes have been Order Acipenseriformes: 2 families, 26 species, stur- drastically overfished. geons and paddlefishes FISH, BIODIVERSITY OF 761

Order (or subclass) Polypteriformes: 1 family, 11 spe- Coelacanths are large (to 180 cm, 95 kg), old (proba- cies, bichirs and reedfish bly 40–50 years), and produce relatively few, live young Subclass Neopterygii (5–26 young per clutch). The gestation period is about Order Lepisosteiformes: 1 family, 7 species, gars 13 months. Replacement rate in the population is there- Order Amiiformes: 1 family and species, the bowfin fore slow. Coelacanths are captured primarily as by- catch in the hook-and-line fishery for oilfish, and it is unlikely they can sustain even the current by-catch rate 1. Lungfishes of 5–10 animals per year. Counts from small subma- Lungfishes today are represented by three families and rines indicate the Comoran species is declining. The six species, all located on the former Gondwanan conti- Comoran government has outlawed its capture, and nents of Australia, South America, and Africa. Lung- trade in coelacanths is outlawed by the Convention on fishes lack jaw teeth but have unusual toothplates on the International Trade in Endangered Species (CITES). the mouth roof and floor. The Australian species, Neoce- ratodus forsteri, is limited to four river systems of north- 3. Sturgeons and Paddlefishes eastern Australia (Fig. 4A). It is large (1 m), with large The most primitive actinopterygian (ray-finned) fishes scales, flipper-like fins, a broad tail, and a single lung. It are the chondrostean sturgeons and paddlefishes. All crushes benthic crustaceans, mollusks, and small fishes 24 species of sturgeons live in the Northern Hemi- with its tooth plates. It can, but does not have to, breathe sphere. All spawn in fresh water, although some species atmospheric oxygen. Its young lack external gills. Neo- move seasonally between marine and fresh water. North ceratodus populations have declined dramatically and American freshwater species include the lake sturgeon the fish is protected; species recovery efforts include and three river sturgeons. Anadromous species include transplantation into several Queensland reservoirs the Atlantic and white sturgeons, the latter being an and rivers. occupant of west coast bays and rivers. White sturgeons The one South American and four African lungfishes attain the largest size of any North American freshwater have eel-like bodies; slender, almost-filamentous paired fish (3.8 m, 630 kg). The world’s largest freshwater fish fins; lack scales; have paired lungs; have larvae with is the beluga sturgeon of eastern Europe and Asia, Huso external gills; and must breathe air to survive. The four huso, at 8.6 m and 1300 kg. African species occur across central and south Africa, Sturgeons have four barbels ahead of a ventrally lo- often in swampy areas that frequently experience cated mouth, five rows of large bony shields on an drought. When a swamp dries up, African lungfishes otherwise scaleless body, and a heterocercal tail (Fig. dig a burrow and can wait 4 years for rains to return. 4C). They are exceptionally long-lived (118 years for The South American species occurs in swampy regions beluga and 70–80 years for white sturgeon), mature of the Amazon and Parana river basins. Comparatively slowly (as late as 30 years old), and spawn infrequently little is known about its biology. (every 3–5 years). They migrate up rivers to spawn in clean sand and gravel areas; hence, dam building and 2. The Coelacanths siltation of rivers both impede their reproduction. A Coelacanths were thought to have gone extinct at the spawning female can be worth thousands of dollars for end of the Cretaceous, 65 million years ago, until a live her caviar alone, and many sturgeon stocks have been one was trawled up in 1938 off South Africa (Fig. 4B). reduced 99% from historical levels. The shortnose stur- Today, a small, endangered population of 200–600 coe- geon of Atlantic coastal rivers is listed as endangered, lacanths lives at 100- to 500-m depths off two small and lake sturgeon have been extirpated from a large volcanic islands in the Comore Archipelago, between part of their native range. Three species endemic to the Madagascar and Mozambique. In 1998, another species Aral Sea may be extinct due to extensive drying of that was discovered at similar depths in northern Indonesia. once huge body of water, and several species in the The living coelacanths have fleshy pectoral, pelvic, anal, former Soviet Union are fished mercilessly. and second dorsal fins (= the lobed fins that define the Two species of paddlefishes occur in large rivers of class Sarcopterygii); a symmetrical, three-lobed tail with North America and China. Paddlefishes also have a a central extension; hollow neural spines (hence ‘‘coel- heterocercal tail, unconstricted notochord, largely carti- acanth’’ or ‘‘hollow spines’’); a unique unconstricted laginous skeleton, and scaleless body. The most distinc- notochord; a joint in the dorsal braincase that aids tive feature of the paddlefishes is their paddle or spoon- jaw opening; relatively large, thick, bony scales; and bill, which is flat and rounded in the North American live young. paddlefish and elongate and almost spear-like in the 762 FISH, BIODIVERSITY OF

FIGURE 4 Living representatives of primitive groups. (A) Australian lungfish (reproduced with permission from Norman, 1905); (B) the living coelacanth, showing anatomic features (reproduced with permission from Musick et al. 1991); (C) Atlantic sturgeon (reproduced with permission from Vladykov and Greeley, 1963); and (D) alligator gar (reproduced with permission from Suttkus, 1963).

Chinese paddlefish. Paddlefishes are water column on the paddle suggest an additional, unknown function. swimmers that feed on zooplankton and fishes. The North American paddlefish may live 30 years and grow paddle of the North American species may help direct to be 2.2 m in length and 83 kg in weight. Late matura- food into its mouth, but abundant electroreceptor cells tion (at 10 years of age), infrequent spawning (every FISH, BIODIVERSITY OF 763

2–5 years), lack of clean gravel spawning habitat, and shaped, interlocking ganoid (heavy and bony) scales. overfishing have all contributed to population reduc- Gars are the only freshwater fish in North America with tions and range contraction. The exceedingly rare and poisonous eggs. Alligator gars can be 3 m long and poorly studied Chinese paddlefish is larger, reaching weigh 140 kg. In recent years, alligator gars have come more than3minlength. This paddlefish may be the under intense commercial fishing, and concern for their most endangered fish in China because of overfishing, well-being is increasing. habitat destruction, pollution, and dam construction that blocks spawning migrations in the Yangtze River basin where it lives. 6. The Bowfin The bowfin, Amia calva, is the only living member of its genus, family, and order. The bowfin has the abbreviate 4. Bichirs heterocercal tail and spiral valve intestine of the gars The polypteriform bichirs and ropefish of west and but also has teleost-like biconcave vertebrae as well as central tropical Africa are difficult to assign to any par- cycloid scales, a relatively light scale type also possessed ticular taxonomic group. Their gilled larvae, lobe-like by many teleosts. The bowfin’s head is exceptionally fins, thick ganoid scales, and modified heterocercal tail bony and the throat is covered by a distinctive large suggest affinities with several living groups, particularly bone, the gular plate. Bowfin swim slowly forwards or the chondrostean sturgeons and paddlefishes. However, backwards by passing undulations back and forth along their unique dorsal, caudal, and paired fins and unusual their long dorsal fin. Bowfin occur throughout much of chromosomes place them apart from all extant groups. the eastern North America in backwater, often swampy Taxonomists usually place them either in the Chon- areas; they also have a highly vascularized gas bladder drostei or in their own subclass, the Brachiopterygii. which functions as a lung. They are relatively large and Ten species of polypteriforms are called bichirs robust (to 1 m and 9 kg) and predatory on anything (Polypterus) and an 11th, elongate species is the reedfish that moves. Bowfin males guard the young vigorously or ropefish, Erpetoichthyes. Bichirs reach 120 cm in until they are relatively large (10 cm). length and ropefish 90 cm. All are predatory and inhabit shallow, swampy regions, in which they breathe atmo- 7. Teleosts spheric oxygen with their paired lungs. Bichirs, also called ‘‘flagfins,’’ have unique dorsal and pectoral fins. The division Teleostei (‘‘perfect bone’’) contains most Each dorsal finlet consists of a vertical spine with living fishes. Teleosts are not only taxonomically di- attached horizontal rays, looking like a flagpole with verse but also ecologically diverse, occupying every streaming banners. In more usual ray-finned fishes, dor- aquatic habitat type and niche imaginable. The 23,600 sal fin rays emerge vertically from the body of the fish. living teleostean species are placed in 4064 genera, 426 The pectoral fin has an internal arrangement that in- families, and 38 orders. This incredible diversity is gen- volves a wishbone-shaped, flattened plate, again unlike erally organized into four taxonomic subdivisions that that in any other fish. Bichirs are not particularly well reflect patterns of evolution that date back to the Meso- studied; no evidence exists to suggest they are im- zoic. These four main subdivisions are the osteoglosso- periled. morphs (bony tongues), elopomorphs (eels and tar- pon), clupeomorphs (herrings), and the euteleosts, with the latter group containing the vast majority of modern 5. Gars bony fishes. The lepisosteiform gars are seven species of elongate, predaceous fishes that occur in eastern North America a. Class Actinopterygii and Central America (Fig. 4D). They typically inhabit backwater areas of lakes and rivers, such as oxbows and bayous, and breathe atmospheric oxygen using a Subclass Neopterygii highly vascularized gas bladder. Gars have bony skele- Division Teleostei tons, but their vertebral centra are unique, being convex Subdivision Osteoglossomorpha anteriorly and concave posteriorly. In most fishes, the Order Osteoglossiformes: 6 families, 217 species, in- vertebrae are concave on both surfaces. Gars have an cluding bonytongues, mooneyes, African knifefishes, abbreviate heterocercal tail and hinged, diamond- and elephant fishes 764 FISH, BIODIVERSITY OF

Osteoglossiforms derive their name ‘‘bonytongue’’ waters. The 15 families of anguilliform ‘‘true’’ eels are from the teeth on their tongue that forms part of their distinguished from the approximately 45 other families bite. These freshwater fishes occur on all major conti- of ‘‘eel-like’’ fishes that have independently evolved an nents except Europe. The arapaima of South America elongate body. Anguillid eels are catadromous, spawn- is one of the world’s largest freshwater fishes, reaching ing at sea but spending most of their lives in fresh a length of 2.5 m. The African mormyrid elephant fishes water. Muraenid moray eels and their relatives are ma- produce and detect weak electric fields, have large cere- rine, tropical and warm temperate, predatory species. bellums, and have a brain size : body weight ratio com- Synaphobranchid cutthroat eels include an endopara- parable to that of humans (Fig. 5A). Two species, the sitic species, the snubnose parasitic eel, which has been mooneye and goldeye, occur in major river systems of found in the heart of a mako shark. The saccopharyngi- northern North America. form deep-sea gulper and swallower eels have giant mouths but lack many head bones, scales, and fins b. Subdivision Elopomorpha found in most other fishes.

Order Elopiformes: 2 families, 8 species, ladyfishes c. Subdivision Clupeomorpha and tarpons Order Albuliformes: 3 families, 29 species, including Order Clupeiformes: 5 families, 357 species, including bonefishes and spiny eels anchovies and herrings Order Anguilliformes: 15 families, 738 species, includ- ing freshwater, moray, cutthroat, and conger eels Clupeomorphs are small, schooling, silvery, pelagic ma- Order Saccopharyngiformes: 26 species, including swal- rine and occasionally freshwater feeders on zooplank- lower and gulper eels ton and phytoplankton. Herrings, round herrings, shads, alewives, sprats, sardines, pilchards, and menha- Elopomorphs all have ribbon-shaped ‘‘leptocepha- dens are extremely important commercial species. An- lus’’ larvae. The Atlantic tarpon is a highly prized game- chovies range in size from a 2-cm Brazilian species to fish that reaches a length of 2.5 m and a mass of a piscivorous, riverine, 37-cm New Guinea anchovy. 150 kg (Fig. 5B). Albuliform bonefishes are also popular The largest clupeids are the Indo-Pacific chirocentrid gamefishes that occupy sandy flats in shallow tropical wolf herrings, which reach a length of 1 m and have

FIGURE 5 (A) Mormyrid elephant fish (reproduced with permission from Helfman et al., 1997); (B) Atlantic tarpon (reproduced with permission from Hildebrand, 1963). FISH, BIODIVERSITY OF 765 fang-like jaw teeth. Anadromous shads, alewives, and world’s smallest freshwater fish is a Burmese cyprinid, herrings occasionally establish landlocked populations Danionella translucida, which matures at 10 mm. The in rivers, lakes, and reservoirs. largest minnow in North America is the endangered, piscivorous Colorado squawfish, Ptychocheilus lucius. d. Subdivision Euteleostei Gyrinocheilid algae eaters scrape algae off rocks in Approximately 95% of the teleosts are placed in the swift, flowing waters. The catostomid suckers (i.e., buf- Euteleostei or ‘‘true’’ teleosts. This subdivision contains faloes, quillback, carpsuckers, blue sucker, redhorses, 391 families, 3795 genera, and 22,262 species—all in jumprocks, and the extinct harelip sucker) include 70 nine superorders. species of North American fishes, with 1 species in eastern China. Loaches (Cobitidae) are 110 species of i. Superorder Ostariophysi predominantly Eurasian stream fishes, including popu- lar aquarium fishes such as the kuhli, clown, and skunk Order Gonorhynchiformes: 4 families, 35 species, in- loaches, the weatherfishes, and the golden dojo. Weath- cluding milkfish erfishes (Misgurnus) become restless when barometric Order Cypriniformes: 5 families, 2660 species, includ- pressure decreases preceding a storm. ing minnows, barbs, algae eaters, suckers, and The characiforms are a speciose group of primarily loaches tropical ostariophysans characterized (usually) by a ray- Order Characiformes: 10 families, 1340 species, includ- less adipose fin and mouths armed with replacement ing freshwater hatchet fishes, tetras, and characins dentition (e.g., piranhas). Body size ranges from very Order Siluriformes: 34 families, 2405 species, including small (13 mm) tetras to large (1.5-m-long) tiger-fishes. North American freshwater, airbreathing, electric, Numerous aquarium fishes are included (headstanders, sea, upside-down, parasitic, callichthyid armored, freshwater hatchet fishes, blind characins, pencil fishes, and suckermouth armored catfishes tetras, and silver dollars), as well as important food Order Gymnotiformes: 6 families, 62 species, including fishes (Prochilodus, Colossoma, and Brycon). Most char- glass, ghost, and naked-back knifefishes and elec- acins (ca. 1150 species) are South American, about 200 tric eel are African, a few live in Central America, and 1 species, the Mexican tetra Astyanax mexicanus, extends naturally Rivers, lakes, and streams worldwide are dominated into southwestern Texas. numerically and ecologically by members of the super- Catfishes (Siluriformes) are surprisingly diverse, order Ostariophysi. Ostariophysans include milkfish, with 34 families and more than 2400 species (Fig. 6A). minnows, carps, barbs, suckers, loaches, piranhas, tet- Catfishes usually have barbels (‘‘whiskers’’) and some- ras, catfishes, and electric eels. Two distinctive traits times toxic spiny fins, and they are almost entirely unite this otherwise disparate assemblage: (i) the We- freshwater, nocturnal, and benthic. Catfishes are most berian apparatus, which is a series of modified anterior diverse in South America (e.g., loricariid suckermouth vertebrae that link the gas bladder to the inner ear armored catfishes, 550 species; pimelodid long-whisk- and aid in hearing, and (ii) production and reaction to ered catfishes, 300 species). Large species include the chemical alarm substances that are released when a fish European wels (5 m, 330 kg), the Asian Mekong catfish is injured and lead to a stereotyped escape response in (3 m, 300 kg), and a 3-m-long whiskered pimelodid of school members. South America. The largest catfishes in North America The gonorhynchiform milkfish, Chanos chanos, is an are the flathead and blue catfishes at about 1.5 m and important food fish in the Indo-Pacific region and is 50–68 kg. Some small catfishes are notable, such as the often cultured in brackish fishponds. The Cyprini- parasitic catfishes (Trichomycteridae) of South formes make up the largest order in the superorder. America, which normally parasitize the gills of fishes The Cyprinidae is the largest family of freshwater fishes but are known to swim up the urethra of bathers and and contains 2000 species of minnows, shiners, carps, lodge there, necessitating surgical removal. barbs, barbels, gudgeons, chubs, dace, squawfishes, The gymnotiform South American knifefishes are tench, rudd, bitterlings, bream, southeast Asian unusual ostariophysans that produce and receive weak ‘‘sharks’’ (redtail black shark and bala shark), goldfish, electric impulses. They have elongate, compressed bod- koi (domesticated common carp), danios, and rasboras. ies; an extremely long anal fin; and electrogenic tissue Cyprinids are most diverse in Southeast Asia, followed usually derived from modified muscle cells. Their elec- by Africa, North America (270 species), and Europe, trical output is constant at high frequencies, whereas but are missing from South America and Australia. The the osteoglossomorph mormyrids produce a pulsed, FIGURE 6 (A) Callichthyid armored catfish (reproduced with permission from Burgess, 1989); (B) chinook salmon (female above and male below) (reproduced with permission from Scott and Crossman, 1973); (C) tripodfish (reproduced with permission from Helfman et al., 1997); (D) lantern fish (reproduced with permission from Nafpaktitis et al., 1977). FISH, BIODIVERSITY OF 767 low-frequency output. Both groups detect objects that and salmons include golden, cutthroat, and gila trouts disrupt their electric fields. The 2-m-long electric eel and the spectacularly anadromous coho, chinook, (Electrophorus) produces a weak field for electroloca- chum, pink, and sockeye salmons, some of which un- tion and strong pulses for stunning prey or deterring dergo oceanic migrations of thousands of kilometers predators. before returning to their birth river to spawn and die. The actual number of genetically distinct races of Pacific ii. Superorder Protacanthopterygii salmons is unknown because many stocks are reproduc- tively isolated in small river systems. Evidence suggests Order Esociformes: 2 families, 10 species, pikes and that as many as 1000 stocks exist, 106 of which have mudminnows gone extinct and an additional 314 of which are im- Order Osmeriformes: 13 families, 236 species, includ- periled. ing barreleyes, smelts, salamander fish, and galaxiids Order Salmoniformes: 1 family, 66 species, whitefishes, iii. Superorder Stenopterygii graylings, chars, trouts, and salmons Order Stomiiformes: 4 families, 321 species, including Protacanthopterygians are a mixed agglomeration of bristlemouths, marine hatchet fishes, and barbeled marine, freshwater, and diadromous fishes. Esociform dragonfishes pikes, pickerels, and mudminnows are Northern Hemi- Order Ateleopodiformes: 1 family, 12 species, jelly- sphere predators; the northern pike has the largest geo- nose fishes graphical distribution of any Northern Hemisphere fish, occurring across the northern portions of North Stenopterygians are deep-sea fishes, often with long America, Europe, and Asia. Mudminnows can survive teeth and large mouths. Gonostomatid bristlemouths winters in high-latitude lakes by breathing from air may be the most abundant and widely distributed verte- bubbles trapped under the ice. Osmeriforms are small, brates on Earth. Idiacanthine black dragonfishes have silvery, elongate, water-column dwelling fishes. Osmer- a larva with eyes at the ends of elongate stalks. ids include commercially important species such as cap- elins, eulachons, Asian ayu, and smelts. The order also iv. Superorder Cyclosquamata includes the Southern Hemisphere lepidogalaxiid sala- mander fish and the galaxiids. Salamander fish inhabit Order Aulopiformes: 13 families, 219 species, including seasonal ponds of southwestern Australia, burying in telescope fishes, tripod fishes, lizard fishes, and lan- drying mud and reemerging with the next rains. Gala- cet fishes xiids have suffered numerous extirpations and extinc- tions as a result of the stocking of nonnative trouts. Cyclosquamates are also deep-sea forms, including Salmoniforms are important commercially, ecologi- the bizarre giganturid telescope fishes with large tubular cally, and aesthetically. Whitefishes and ciscoes are rela- eyes, a huge mouth, flexible teeth, and an expandable tively large-scaled, zooplanktivorous salmonids of high- stomach. Deep-sea tripod fishes have long pectoral, pel- latitude North American and Eurasian lakes. Several vic, and caudal rays that they use for resting on soft North American species have been decimated due to sediments of the deep ocean floor (Fig. 6C). Shallow introduced predators, competitors, and parasitic lam- representatives are the synodontid lizard fishes, which preys. Graylings are riverine fishes with a flowing dorsal are common benthic predators on coral reefs world- fin. At least one species, the Michigan grayling, is ex- wide. Alepisaurid lancet fishes are large (to 2 m) meso- tinct. The subfamily Salmoninae contains seven Eur- pelagic predators with a sail-like dorsal fin of un- asian and North American genera. The Siberian taimen, known function. Hucho taimen, is the world’s largest salmonid at 2 m and 70 kg. North American Salmoninae include the v. Superorder Scopelomorpha chars (lake, brook, and bull trout, Arctic char, and dolly varden). Arctic char live farther north than any other Order Myctophiformes: 2 families, 240 species, includ- freshwater fish. The remaining salmonines are the At- ing lantern fishes lantic basin salmon and trout (e.g., Atlantic salmon and European brown trout), and the 11 species of Pacific Scopelomorphs include the abundant, commercially basin trouts and salmons in the genus Oncorhynchus, important lantern fishes, which are identified based on 2 of which are endemic to Japan (Fig. 6B). Pacific trouts species-specific photophore (light organ) patterns. Lan- 768 FISH, BIODIVERSITY OF tern fishes occur at middle depths from the Arctic to and bythitid cusk eels and brotulas include blind cave the Antarctic. They are important in the diets of many species in freshwater systems of Caribbean and Galapa- fishes as well as of marine mammals (Fig. 6D). gos Islands as well as coral reef species that hide deep within crevices. The neobythitine cusk eel, Abyssobro- vi. Superorder Lampridiomorpha tula galatheae, holds the depth record for a fish at 8370 m in the Puerto Rico Trench. Order Lampridiformes: 7 families, 19 species, including The gadiforms include the cods, haddocks, hakes, opahs, tube-eye, ribbonfishes, and oarfishes pollocks, and whitings, which are some of the world’s most important commercial fishes. True cods (Gadinae) Lampridiforms are generally open-water, oceanic have three dorsal fins and two anal fins. Many species fishes. Opahs are relatively large (1.8 m, 70 kg), oval- have chin barbels. The burbot, Lota lota, of high-lati- shaped, pelagic predators on squids and other fishes. tude, Northern Hemisphere lakes is the only freshwater The 30-cm-long tube-eye (Stylephorus) can increase the species in the group. The commercially important At- volume of its mouth 40-fold during feeding—a record lantic cod is the largest species (1.8 m, 90 kg), but fish among vertebrates. The elongate oarfish, Regalecus, may more than 10 kg are rare due to drastic overfishing attain 12-m length and is the longest living teleost. It (Fig. 7A). One of the world’s largest food fisheries is has a bluish-silvery body, scarlet head crest, and deep for North Pacific walleye pollock. red fins. It is thought to be responsible for many ‘‘sea Batrachoidiforms include the midshipmen, which serpent’’ sightings. have hundreds of photophores, an unusual trait for a shallow dweller. Many batrachoids produce sounds by vibrating their gas bladders. Venomous toadfishes have vii. Superorder Polymixiomorpha dorsal and opercular spines which can inject a powerful toxin. Three South American toadfishes are restricted Order Polymixiiformes: 1 family, 5 species, beardfishes to fresh water. Lophiiforms are a diverse and often bizarre-looking viii. Superorder Paracanthopterygii order of marine fishes that include benthic, shallow- water forms as well as highly modified, open-water, Order Percopsiformes: 3 families, 9 species, trout- deep-sea forms. Many use a modified first dorsal spine perches, pirate perch, and cavefishes as a lure for catching smaller fish. The meter-long west- Order Ophidiiformes: 5 families, 355 species, including ern North Atlantic goosefish, Lophius americanus, has pearlfishes, cusk-eels, and viviparous brotulas a huge mouth with long, recurved teeth which it uses Order Gadiformes: 12 families, 482 species, including to catch fishes and even diving seabirds. Antennariid rattails, hakes, and cods frogfishes also rest on the bottom or walk over it with Order Batrachoidiformes: 1 family, 69 species, toad- their pectoral and pelvic fins (Fig. 7B). The ogcocepha- fishes lid batfishes walk on their pectorals, but they can also Order Lophiiformes: 16 families, 297 species, including swim via jet propulsion by shooting water out their goosefishes, frogfishes, handfishes, batfishes, and round, backward-facing opercular openings. The cera- deep-sea anglerfishes tioid anglerfishes include 11 families of strange- appearing bathypelagic predators, many of which Paracanthopterygians are primarily benthic, marine, have very small males that fuse to and become parasitic nocturnally active fishes; many live in the deep sea or on the larger females. The endemic Australian hand- in caves. Percopsiforms are small (Ͻ20 cm), freshwater fishes include a Tasmanian species, the spotted hand- fishes, most of which live in eastern North America. fish, that was once common but is now critically endan- The anus of the swamp-dwelling aphrododerid pirate gered due possibly to egg predation by an introduced perch is located in the throat region of adults for func- starfish. tionally mysterious reasons. Amblyopsid cavefishes are often blind and scaleless forms highly adapted for e. Superorder Acanthopterygii cave life. Most bony fishes belong to a single superorder, the Ophidiiforms often live in holes or even inside other Acanthopterygii, which contains about 13,500 species animals. Carapid pearlfishes live inside the body cavities in 251 families. Two small and one large taxonomic of starfishes, sea cucumbers, clams, and sea squirts; groupings, called series, are recognized, with the vast some feed on the internal organs of their hosts. Ophidiid majority in the third series, the Percomorpha. FIGURE 7 (A) Atlantic cod (reproduced with permission from Norman, 1905); (B) bloody frogfish (reproduced with permission from Heller and Snodgrass, 1903); (C) atherinomorph four-eyed fish (reproduced with permission from Norman, 1905); (D) lined seahorse (repro- duced with permission from Norman, 1905). 770 FISH, BIODIVERSITY OF

i. Series Mugilomorpha at an elevation of 4570 m in the Andes Mountains is the highest natural lake with fishes. Order Mugiliformes: 1 family, 66 species, mullets

The mullets are a family of nearshore, marine and iii. Series Percomorpha freshwater fishes of considerable economic importance. Many mullets feed on organic silt and minute plants, Order Stephanoberyciformes: 9 families, 86 species, in- an unusual food type among fishes. cluding whale fishes Order Beryciformes: 7 families, 123 species, including ii. Series Atherinomorpha flashlight fishes, roughies, and squirrelfishes Order Zeiformes: 6 families, 39 species, including dories Order Atheriniformes: 8 families, 285 species, including and boarfishes rainbow fishes and silversides Order Gasterosteiformes: 11 families, 257 species, in- Order Beloniformes: 5 families, 191 species, including cluding sticklebacks, pipefishes, seahorses, trumpet needlefishes, flying fishes, and halfbeaks fishes, and shrimp fishes Order Cyprinodontiformes: 8 families, 807 species, in- Order Synbranchiformes: 3 families, 87 species, includ- cluding topminnows, killifishes, livebearers, and ing swamp and tiretrack eels pupfishes Order Scorpaeniformes; 25 families, 1271 species, in- cluding scorpion fishes, rockfishes, sea robins, sable- Atherinomorphs are shallow-water, marine or fresh- fishes, greenlings, sculpins, Baikal oilfishes, and water fishes that live near the surface. Many atherino- lumpfishes morphs bear live young. Atheriniforms include the mel- Order Perciformes (9300 species) anotaeniid rainbow fishes of Australia and New Guinea, Suborder Percoidea: 71 families, 2860 species, includ- in which males have brighter colors and longer fins ing snooks, temperate basses, sea basses, centrarchid than females, traits that make them popular aquarium sunfishes, black basses, darters, perches, cardinal species. Silversides are widespread, freshwater and ma- fishes, bluefishes, remoras, dolphin fishes, jacks, rine schooling fishes and include the grunions of south- pompanos, snappers, grunts, croakers, drums, goat- ern and Baja California, which ride waves up beaches fishes, archerfishes, butterfly fishes, and angelfishes on dark nights to spawn in wet sand biweekly during Suborder Elassomatoidei: 1 family, 6 species, pygmy the summer. Beloniforms are predominantly silvery, sunfishes marine fishes active at and sometimes above the surface Suborder Labroidei : 6 families, 2200 species, including of the water. The lower lobe of the tail in flying fishes cichlids, surfperches, damselfishes, wrasses, and par- is relatively long and is used to scull rapidly during rotfishes takeoff. Many cyprinodontiforms, although basically Suborder Zoarcoidei: 9 families, 318 species, including freshwater fishes, tolerate considerable salinity and eel pouts, gunnels, and wolffishes hence occur in streams on isolated oceanic islands (Fig. Suborder Notothenioidei: 5 families, 122 species, in- 7C). The rivulines of South America and Africa live cluding icefishes and Antarctic dragonfishes only 1 year, laying eggs which survive in the dried Suborder Trachinoidei: 13 families, 212 species, includ- bottoms of pools and which hatch with the next season’s ing sand lances, weeverfishes, and stargazers rains. Rivulus marmoratus of south Florida and the West Suborder Blennioidei: 6 families, 732 species, including Indies is the only truly hermaphroditic fish, fertilizing clinids and blennies its own eggs. The livebearers include the mollies, platys, Suborder Icosteoidei: 1 family and species, ragfish guppies, and swordtails of the aquarium trade. Some Suborder Gobiesocoidei: 1 family, 120 species, species are all female. Many cyprinodontid pupfishes clingfishes are tolerant of extreme water conditions and conse- Suborder Callionymoidei: 2 families, 137 species, drag- quently can live in saltmarsh and desert conditions. onets However, they cannot tolerate total desiccation, which Suborder Gobioidei: 8 families, 2120 species, including has endangered many desert species that have to com- sleepers and gobies pete with humans for water. The Devil’s Hole pupfish Suborder Kurtoidei: 1 family, 2 species, nurseryfishes has the smallest known range of any fish species—one Suborder Acanthuroidei: 6 families, 125 species, includ- shallow shelf in a single spring in Death Valley, Califor- ing spadefishes, scats, rabbitfishes, moorish idol, nia. Other pupfish relatives inhabit Lake Titicaca, which and surgeonfishes FISH, BIODIVERSITY OF 771

Suborder Scombrolabracoidei: 1 family and species, the tant, long-lived, and overfished group of the temperate scombrolabracid North Pacific. Hexagrammid greenlings are littoral zone Suborder Scombroidei: 6 families, 136 species, includ- and kelp-associated fishes endemic to the North Pacific, ing barracudas, mackerels, tunas, swordfish, and bill- including the highly edible lingcod, Ophiodon elongatus. fishes The suborder Cottoidei includes many freshwater spe- Suborder Stromateoidei: 6 families, 65 species, includ- cies, including the cottid sculpins of North American ing driftfishes and butterfishes streams and the highly divergent (pelagic and livebear- Suborder Anabantoidei: 5 families, 81 species, gouramis ing) comephorid oilfishes of Lake Baikal in northern Suborder Channoidei: 1 family, 21 species, snakeheads Asia. The cabezon of the Pacific coast of North America Order Pleuronectiformes: 11 families, 570 species, in- is unusual in having toxic eggs, whereas the lumpfish cluding flounders and soles of the North Atlantic produces valuable caviar and has Order Tetraodontiformes: 9 families, 339 species, in- consequently been overfished. cluding triggerfishes, boxfishes, trunkfishes, cow- The Perciformes are the largest order of perco- fishes, puffers, porcupine fishes, burrfishes, and morphs, with 148 families and 9300 species, including ocean sunfishes most marine and freshwater fishes of littoral (inshore and shallow-water) zones. Perciforms reach their great- The percomorphs constitute by far the largest taxo- est diversity on coral reefs, but they are also highly nomic group of fishes, far too many to deal with in any diverse in rivers, streams, and lakes. Coral reef perci- detail. What follows is a very brief overview of some forms include six of the eight largest fish families (go- of the more interesting orders, suborders, and fam- bies, wrasses, sea basses, blennies, damselfishes, and ilies. cardinal fishes). Two other large families, cichlids and Beryciforms are shallow- to moderate-depth, often croakers, are characteristic of tropical lakes and near- red, almost always nocturnal fishes, including the reef- shore temperate marine habitats, respectively. dwelling squirrelfishes. Also included is the commer- The largest perciform suborder is the Percoidei. cially important orange roughy, Hoplostethus atlanticus, Many percoids are bass-like fishes. Centropomids are of high-latitude, southern ocean regions. Orange rough- large predatory fishes and include the snooks of tropical ies are being overexploited because they are slow grow- America, the barramundi of Australia, and the Nile ing and long-lived, taking more than 20 years to mature perch of Africa (Fig. 8A). Nile perch are an introduced and reaching ages more than 100 years. Zeiforms in- predator in Lake Victoria, in which they are thought clude commercial species such as the European John to have extinguished perhaps hundreds of endemic Dory, Zeus faber. cichlids. Moronid temperate basses include the striped Gasterosteiforms are small marine, estuarine, and and white bass of North America. The sea bass family freshwater fishes with dermal armor plating. Stickle- Serranidae is one of the largest families (450 species) backs are well-studied fishes that frequently form dis- that range in size from small (Ͻ5 cm) anthiines and tinct, isolated populations characterized by unusual hamlets to giant groupers and jewfish (3 m long, 400 spines, plates, and behavior. The suborder Syngna- kg). Sea basses also include commercially important thoidei includes the bizarre pegasid seamoths and syng- hinds, coneys, gag, and scamp (Fig. 8B). Many serranids nathid pipefishes, sea dragons, and seahorses. Syng- are hermaphroditic, usually starting as female and then nathid pipefishes and seahorses are the only vertebrates later becoming male, although some hamlets are both in which the female ‘‘impregnates’’ the male by laying male and female simultaneously. eggs in his brood pouch, which he then fertilizes and The Centrarchidae contains the sunfishes, crappies, raises until hatching. Seahorses are heavily overfished rockbasses, and black basses of North American fresh for medicinal uses and the aquarium trade (Fig. 7D). waters (Fig. 8C). Largemouth bass have been intro- Synbranchiforms are primarily freshwater, eel-like, duced extensively as a sport fish. At least 162 species often air-breathing fishes. Swamp eels have recently make up the family Percidae, 150 of which occur in been introduced into the southeastern United States North America. Percids include the yellow and Eurasian and are of major concern as potential invading predators perch, walleye and sauger (= pikeperches), and about in river systems. 150 species of small, stream-dwelling, spectacularly col- Scorpaeniforms are predominantly marine fishes, ored, and often imperiled darters. with spiny heads and sometimes venomous fin spines Apogonid cardinal fishes are diverse (320 species), (i.e., stonefishes, scorpion fishes, and lionfishes). The small (Ͻ10 cm), nocturnal coral reef fishes. Cardinal sebastine rockfishes are a diverse, commercially impor- fishes mouthbrood their eggs, an unusual trait among 772 FISH, BIODIVERSITY OF

FIGURE 8 Representative percoids: (A) a centropomid snook; (B) a serranid black sea bass; (C) a centrarchid smallmouth black bass; (D) a chaetodontid four-eye butterfly fish (A, C, and D, reproduced with permission from Norman, 1905); B, reproduced with permission from Bigelow and Schroeder, 1953).

reef species. The voracious, schooling pomatomid blue- bass, corbinas, and the endangered Mexican totoaba, fish, Pomatomus saltatrix, occurs in most temperate and one of the few unquestionably imperiled marine fishes. semitropical oceans except the eastern Pacific. In echen- The range of the freshwater drum, Aplodinotus grun- eid remoras or sharksuckers, the modified first dorsal niens, includes much of eastern North America into fin forms a suction disk that is used to cling to various Central America. Another bottom-oriented family hosts. The coryphaenid dolphin fishes or mahi mahis is the tropical reef-dwelling mullid goatfishes, which are two species of open-water, pelagic predators that have movable, muscularized chin barbels. Toxotid often associate with floating structure. They are also archerfishes are Indo-Pacific, brackish-water fishes one of the few marine pelagic fishes that is successfully well-known for their ability to shoot droplets of water aquacultured. Carangid scads, jacks, pompanos, and that knock insects out of overhanging vegetation. amberjacks are a large family (140 species) of tropical Two closely related, colorful reef families are the nearshore predators. chaetodontid butterfly fishes (114 species) and the po- Lutjanid snappers include 125 species of generally macanthid angelfishes (74 species). Both families are carnivorous marine fishes. Snappers are usually near- most diverse in the Indo-Pacific region. Butterfly fishes bottom dwellers (i.e., gray snapper, red snapper, and feed on coral polyps, small invertebrates, tube worms, mangrove snapper), but some are water-column zoo- or zooplankton, whereas angelfishes eat attached inver- planktivores (vermilion snapper). tebrates such as sponges, tunicates, and anthozoans Haemulidae grunts are moderate-sized coral reef (Fig. 8D). fishes and are most diverse in the New World tropics. Elassomatoid pygmy sunfishes are a suborder of min- They are seen most often in their daytime resting iature (20- to 45-mm), colorful freshwater swamp schools around coral heads; at night they disperse to dwellers of the southeastern United States. feed in surrounding reef and grass areas. Sciaenid croak- Several very speciose families belong to the suborder ers (270 species) have chin barbels and a muscularized Labroidei. Tropical marine families include the po- gas bladder used for sound production. Sciaenids are a macentrid damselfishes (315 species), which are small, widespread family that is particularly diverse in the colorful, usually herbivorous, territorial reef dwellers. southeastern United States, but representatives occur Some are zooplanktivores (Chromis and ane- widely in tropical marine and freshwater habitats. Spe- monefishes). Several occupy temperate regions (e.g., cies include red drum (spot tail bass), black drum, the garibaldi of California). The largest labroid family croakers, weakfish, sea trouts, kingfishes, white sea is the mostly tropical, reef-dwelling wrasses, Labridae FISH, BIODIVERSITY OF 773

(500 species). Wrasses range in size from the 5-cm- reefs. They are live-bearers, feeding on zooplankton or long cleaner wrasses, slippery dicks, and blueheads to small invertebrates. The largest labroid family is the the 2.3-m-long Maori wrasse of the Indo-Pacific, which freshwater cichlids, with more than 1300 species. Cich- is hunted unmercifully for the live-fish restaurant trade. lids are chiefly tropical, South American and African Cool temperate species include the California sheep- fishes, with a few species that occur further north (the head (Fig. 9A) and senorita, western Atlantic tautog Rio Grande cichlid is found in south Texas). Central and cunner, and the eastern Atlantic cuckoo wrasse. and South American species include freshwater Many wrasses change sex from female to a more colorful angelfishes, discus, oscars, convict cichlids, and pea- male. The scarid parrotfishes (83 species) are almost cock bass (Fig. 9B). Most cichlids occur in Africa, where exclusively coral reef dwellers, best known for their they are particularly speciose in the African Great Lakes fused parrot-like teeth that are used for biting off algal and are threatened by introduced predators such as Nile and coral pieces, which are then crushed in the massive perch. African tilapias and other cichlids have been pharyngeal (throat) jaws. The embiotocid surfperches deliberately or accidentally introduced into Florida, are 24 species of deep-bodied, temperate (mostly east- California, and Hawaii as a by-product of aquaculture ern Pacific) fishes associated with kelp beds and rocky activities.

FIGURE 9 (A) California sheephead (reproduced with permission from Norman, 1905); (B) blue-eye cichlid (reproduced with permission from Greenfield and Thomerson, 1997); (C) Japanese goby (reproduced with permission from Norman, 1905); (D) four-spot flounder (reproduced with permission from Bigelow and Schroeder, 1953); (E) gray triggerfish (reproduced with permission from Bigelow and Schroeder, 1953). 774 FISH, BIODIVERSITY OF

Fishes of the suborder Zoarcoidei are all eel-like, from feeding on the feces of other animals. Rabbitfishes bottom-living, marine, cool- to cold-water species. They are Indo-Pacific reef, grassbed, and estuarine herbivores range in size from the small, intertidal pricklebacks and that are convergent in many ways with some butterfly gunnels to the live-bearing eel pouts, some of which fishes, a description that also applies to the moorish live 3000 m below the surface. The large (to 2.5 m idol (Zanclidae). The 72 species of usually herbivorous long) anarhicadid wolffishes and wolf eels of shallow acanthurid surgeonfishes, unicornfishes, and tangs have North Pacific and Atlantic waters are anatomically and a knifeblade on the caudal peduncle. ecologically similar to moray eels. The suborder Scombroidei contains some of the Icefishes (suborder Notothenioidei) are mostly Ant- largest and most spectacular marine fishes. Twenty spe- arctic, mostly benthic fishes that live under the ice cies of barracudas inhabit tropical and subtropical and have antifreeze compounds in their blood. The oceans almost worldwide. The gempylid snake macker- crocodile icefishes lack red blood cells, hemoglobin, els (23 species) are pelagic and deep-water predators, and myoglobin, and hence have colorless blood and including the cosmopolitan oilfish, Ruvettus pretiosus, flesh. a large (1.8 m, 45 kg) predator of moderate depths. An Trachinoids are marine, generally benthic fishes with active fishery for oilfish in the Comoro Islands captures a tendency to bury themselves in sand. Ammodytid sand endangered coelacanths as by-catch. The scombrid lances are small, elongate, and abundant zooplankton mackerels and tunas are quintessential open-sea preda- feeders that spend their nights buried. Trachinid wee- tors, with streamlined bodies and a physiology geared verfishes occur in the eastern Atlantic and Mediterra- to a high-speed lifestyle. They range from relatively nean and have venomous opercular and dorsal spines. small, 50-cm mackerels to giant bluefin tuna (4 m, 500 Uranoscopid stargazers emit strong pulses of electricity kg). Most are schooling fishes of tremendous commer- (up to 50 V) from highly modified eye muscles. cial importance. The temperate and warm-temperate Blennioids are small marine fishes that usually asso- xiphiid swordfish and the more tropical istiophorid ciate with structure. Chaenopsid pikeblennies and tube- sailfishes, spearfishes, and marlins have an elongate blennies often live inside corals and worm tubes. upper jaw bone that forms the bill. It is used as a Combtooth blennies are diverse (345 species) small spear, a cutlass, or a billy. Swordfish grow to 530 kg, fishes in tropical and subtropical waters; they scrape whereas blue and black marlin grow to 900 kg. Sword- algae with their comb-like teeth. fish have been heavily overfished, particularly in the The suborder Icosteoidei consists of the peculiar Atlantic. North Pacific ragfish, Icosteus aenigmaticus, which has Labyrinth fishes (Suborder Anabantoidei) have an a largely uncalcified, cartilaginous skeleton and is a auxiliary breathing structure in the gill chamber for preferred food of sperm whales. The small, marine and aerial respiration. Anabantid climbing gouramis are Af- freshwater gobiesocoid clingfishes are shallow-water rican and Asian freshwater fishes that can move across and even amphibious fishes often found in high-energy wet ground and reportedly up wet tree trunks. The wave zones. Their pelvic fins are modified into a suck- kissing gourami is the sole member of the family Helo- ing disc. stomatidae. The belontiid gouramis, fighting fishes (bet- Gobioids are usually small, benthic, often abundant tas), and paradise fishes have elongate pelvic fin rays fishes. The eleotrid sleepers are small to medium (to that serve as feelers. Bettas (Siamese fighting fish) have 60 cm) estuarine and stream fishes in tropical and sub- been bred to battle like fighting cocks, placing them tropical areas, often on islands. The gobiid gobies are among the few fishes that have been cultured for pur- incredibly diverse (Ͼ1875 species). Many gobies have poses other than food, appearance, or research. a suction disk formed by fused pelvic fins (Fig. 9C). The Pleuronectiform flatfishes are distinctive, com- family includes the amphibious mudskippers. Gobies pressed, benthic fishes that have both eyes on the same range in size from tiny pygmy gobies (i.e., 8- to 10- side of the head (Fig. 9D). Many flatfishes are important mm adults) to a comparative giant, the western Atlantic commercially (e.g., dab, flounders, halibuts, plaice, violet goby, which is a purplish eel-like fish 50 cm long. sole, tonguefishes, turbots, and whiffs). Paralichthyids Round gobies have recently been introduced into North include the summer flounder and California halibut, American Great Lakes from southern Europe and are the latter reaching 1.5 m and 30 kg. The pleuronectid spreading rapidly; other introduced goby species are right-eye flounders include the Atlantic and Pacific hali- now extremely common in San Francisco Bay. buts. Pacific halibuts may live 40 years and attain The suborder Acanthuroidei includes the ephippid lengths of 3 m and masses of 200 kg. The fishery for spadefishes of Atlantic reefs and beaches as well as the Pacific halibut in the North Pacific is a well-regulated, Indo-Pacific scatophagid scats, which get their name sustainable enterprise. FISH, BIODIVERSITY OF 775

The most advanced bony fishes are in the order Tet- zoogeographic distinction can be made between marine raodontiformes, an almost entirely marine order of me- and freshwater fishes, with substantial overlap oc- dium-sized fishes with thick, leathery skin and with curring where intermediate salinities occur. Many fishes scales often modified into spines or bony plates. In are restricted to pure fresh water (little or no salinity), balistoid triggerfishes and filefishes, the long, rigid first many are restricted to normal oceanic salinity (about dorsal spine is locked erect by an interaction with the 35 parts per 1000 salt in water), some occur in both shorter, second spine (Fig. 9E). Ostraciid boxfishes are habitats at different times of their lives or of the year, encased in a triangular or rectangular bony box, with and some occur and are even restricted to areas of just the fins and caudal peduncle emerging. Puffers and intermediate salinity, such as estuaries. ocean sunfishes lack true teeth. Instead, the jaw bone In terms of numbers, about 58% of all fishes are has a cutting edge that looks like separated teeth or is marine and 41% live in fresh water, with the remaining fused into a parrot-like beak. Diodontid porcupine 1% moving regularly between the two salinity designa- fishes inflate their body by filling the stomach with tions (Fig. 10 and Box 2). Among the 10,250 freshwater water, a process that also helps erect and interlock species, 80% are primary or obligatory freshwater fishes their body spines. Tetraodontid puffers concentrate a and are intolerant of even moderate salinities. The re- powerful and potentially fatal toxin, tetraodotoxin, in maining 20% can tolerate some salinity and hence in- their viscera, which adds to the allure of eating puffers habit upper estuarine areas or can cross through near- in licensed Fugu restaurants in Japan. The ocean sun- shore ocean regions to move from one river basin to fish, Mola mola, is one of the world’s heaviest fishes at another. Among the 14,500 marine fishes, the vast ma- 1000–2,000 kg, producing as many as 300 million eggs. jority (69%) live in shallow, warm areas such as coral All this biomass is supported on a diet of jellyfishes. reefs. The remaining marine species are divided fairly evenly among shallow, cold, deep, open-ocean and deep-bottom areas (about 10% each). About 2% of ma- II. GEOGRAPHIC DIVERSITY rine fishes live in near-surface, open-sea (pelagic) habi- tats. The approximately 160 diadromous species that A. Overview live in different salinity regions at different times of their lives are divided among three groups. Anadromous Fishes occur just about everywhere water occurs as fishes (54%) live most of their lives in the ocean but then long as water is in its liquid state, is available through migrate to fresh water to spawn; this group includes most of the year, and remains below 40ЊC. A major lampreys, sturgeons, herrings, and salmons. Catadro-

FIGURE 10 The global biodiversity of marine, diadromous, and freshwater fishes. Numbers of species are given for each habitat type and its subdivisions. 776 FISH, BIODIVERSITY OF

Box 2 long before the fishes actually spawn; examples include Why Are There so Many Freshwater Fishes? gobies, sleepers, and galaxiids. The high global diversity of freshwater fishes (Fig. 10) is at first surprising. Fresh waters comprise B. Freshwater Diversity only about 0.009% of the earth’s water, which The world’s freshwater habitats occur in six major zoo- means that almost half of all fish species live in geographic regions or realms that correspond approxi- less than 1% of the world’s water. This 7500-fold mately to continental distributions, with important ex- discrepancy in biodiversity per unit volume is ceptions (Fig. 11). Each region has a fairly distinct fish probably best explained by the relative productiv- fauna (again with some exceptions and shared ele- ity and isolation of freshwater bodies. Most fresh- ments). water habitats are relatively shallow and receive ample sunlight and nutrients running off from 1. The Nearctic Region adjacent land. Hence, freshwater habitats are rela- The Nearctic region consists of subtropical, tropical, tively productive and capable of sustaining abun- temperate, and arctic North America. The region dant life. Most of the ocean, in contrast, is deep, stretches from the Mexican Plateau to northern Canada dark, and nutrient poor. Given that 81% of marine and Alaska. The Nearctic contains 14 families of pri- diversity occurs in shallow regions, a relationship mary freshwater fishes, with about 950 species. The between water depth and diversity in fresh waters most diverse families are minnows, suckers, North is not surprising. In addition to the influence of American catfishes, perches (and darters), and sun- available food is the comparative isolation of most fishes. Other important families include the lampreys, freshwater habitats. Lakes are often created and gars, salmons (many of which are anadromous), and affected by climatic and geologic forces (e.g., whitefishes; sculpins, which are freshwater species in drought, floods, landslides, earthquakes, and up- a primarily marine family (= ‘‘marine derivatives’’); pick- lifts) that separate them from other systems. Every erels and mudminnows; killifishes; and livebearers. The lake can be relatively isolated from other lakes, Nearctic is further subdivided into three subregions: the which means that genetically distinct populations Arctic-Atlantic (with six provinces), the Pacific (with can evolve into new species and that little genetic seven provinces), and the Mexican Transition subdivi- mixing occurs between lakes. Small streams are sion. Eleven major river systems drain the region; major separated from each other by larger rivers, which lakes are abundant, the largest being the five Laurentian are barriers to the movement of small fishes, and Great Lakes (Ontario, Erie, Huron, Michigan, and large rivers are separated by oceans. As a result, Superior). freshwater habitats are perfect for the speciation process. Oceans, in contrast, are largely continu- 2. The Neotropical Region ous habitats that are connected by currents, and The Neotropical region contains South America and ocean fishes typically produce larvae that float Middle America. It is the most speciose region of the for several weeks or months on these currents. world in terms of freshwater fishes, with 32 families Hence, genetic exchange is common and opportu- and more than 2500 species. Particularly diverse groups nities for speciation are not as great. Oceanic ba- include the colorful characiforms (1200 species of tet- sins have relatively distinct faunas, but connected- ras, piranhas, characins, and freshwater hatchet fishes), ness within basins discourages the kind of genetic 13 families and 1300 species of catfishes, 6 families isolation needed for speciation of the sort seen in and 62 species of gymnotiform South American electric lakes and rivers. knifefishes, and 150 species of cichlids. Several second- ary freshwater and marine derivative groups are in- cluded: freshwater stingrays, herrings, silversides, nee- dlefishes, killifishes, and croakers. Many species remain to be discovered and described, particularly in South mous fishes (25%) spend most of their lives in fresh America. The Neotropical region has been further di- water and migrate to the sea to spawn; included here vided into eight subdivisions with fairly distinctive fau- are river eels, mullets, and temperate basses. Amphi- nas. Eight major river systems drain the region; major dromous fishes (21%) move between fresh and salt lakes include Lake Titicaca, the world’s highest fish- water, but migration to the spawning habitat occurs containing lake. FISH, BIODIVERSITY OF 777

FIGURE 11 Major freshwater realms. Numbers indicate large river drainages, and letters show locations of larger lakes and inland seas. Neotropical, Ethiopian, Oriental, and Australian realms are shaded (reproduced with permission from Bond, 1996).

3. The Palearctic Region of the fishes are in the superorder Ostariophysi, includ- The Palearctic region encompasses Eurasia, including ing 300 minnows, 190 characiforms, and 360 catfishes Europe, northern Africa, and Asia north of the Oriental in six families. Other diverse groups include killifishes region. Twenty-seven families and about 550 species of and topminnows, elephant fishes and other osteoglossi- temperate freshwater fishes occur in the region, domi- forms, and cichlids. As many as 1000 cichlid fishes may nated by minnows and loaches but also perches, picker- occur in the three African Great Lakes of Lake Victoria, els, sturgeons, salmons, sculpins (including the Lake Lake Tanganyika, and Lake Malawi, with more cichlids Baikal endemics), and 10 species of catfishes in four in smaller surrounding lakes and rivers. Four lungfishes families. Diversity is greater in the southeastern than and all 11 polypteriform bichirs occur in Africa. Ten other parts of the region and also increases to the south, to 12 zoogeographic provinces are recognized, with six as is also the case in the Nearctic region. The Palearctic major river drainages, and numerous lakes including and Nearctic regions share numerous families and gen- the African Great Lakes. era (sturgeons, paddlefishes, minnows, smelts, salmons, pikes, mudminnows, and perches) but only a few spe- 5. The Oriental Region cies (i.e., northern pike, longnose sucker, burbot, The Oriental region includes eastern Iran, India and Sri threespine stickleback, and fourhorn sculpin). The re- Lanka, China south of the Yangtze River, Southeast gion is sometimes subdivided into six subregions based Asia, and the large island regions of Taiwan, the Philip- on faunal groupings. Ten major river systems drain the pines, and the East Indies/Indo-Malayan Archipelago. region; major lakes include the Black and Caspian Seas The Oriental region contains 43 families of primary and Lake Baikal, the world’s oldest and deepest lake. and secondary freshwater fishes. Most diverse are the minnows, loaches, and 12 families of catfishes; clariid 4. The African or Ethiopian Region walking catfishes and bagrid catfishes are particularly The African or Ethiopian region is second to the Neo- diverse. Other important groups include algae eaters, tropics in freshwater fish diversity, with 47 families river loaches, snakeheads, spiny eels, labyrinth fishes and more than 2000 species of primary and secondary and gouramis, a few cichlids, and archerfishes. The freshwater fishes. The African region includes all the Oriental region shares many families with the Palearctic African continent south of the Sahara Desert, plus the to the north and the Ethiopian to the west but few with large island of Madagascar with its endemic fauna. Half the Australian region to the southeast. The Oriental is 778 FISH, BIODIVERSITY OF often subdivided into two major subregions: Peninsular C. Marine Diversity India with more than 700 species and Southeast Asia with more than 1000 species. Each subregion has two Delimiting zoogeographic regions in the world’s oceans major river drainage systems; large lakes are uncom- is complicated by depth, currents, and geographic lo- mon. Southeast Asia is sometimes divided further into cales; different faunal breaks occur depending on near- five zoogeographic regions. shore, pelagic, or deep-sea environments. The greatest fish diversity and the greatest geographic differentiation occur in nearshore, continental shelf (to about 100-m 6. The Australian Region depth) regions. These regions are separated by conti- The Australian region (New Guinea, Australia, New nents, by large expanses of open ocean, and by currents Zealand, and Oceania) is relatively depauperate in true that differ in temperature from that of the region in freshwater fishes, and in fact all but three of the freshwa- question. Temperature zones divide the seas into tropi- ter fishes in the region are members of families obvi- cal, temperate, boreal, and polar regions (Fig. 12). In ously derived from marine groups. The northwestern addition, different faunal groupings apply to pelagic border of the region, and the practical limit of primary fishes and to fishes of the deep sea. freshwater fishes, is dramatically delineated by an ocean boundary that lies southeast of Java, Borneo, and Su- 1. The Indo-West Pacific Region lawesi and is known as Wallace’s or Weber’s Line. Nine- The Indo-West Pacific region includes shallow tropical teen families and about 210 species occur primarily in seas that extend from South Africa and the Red Sea fresh water in the region, but only the Australian lung- eastwards through the Indo-Malayan area and Australia fish and 2 species of bonytongue saratogas are true to Hawaii and Easter Island; it also includes Micronesia, freshwater fishes (another 33 families and 150 species of Melanesia, and Polynesia. The Indo-West Pacific is by marine fishes frequently enter fresh waters in Australia). far the most species-rich marine area, containing 3000 Other important families, many with species endemic to 4000 tropical fish species, and is similarly diverse in to specific regions, include lampreys, river eels, her- sea snakes and many invertebrate taxa such as reef- rings, two families of catfishes, southern smelts and building and soft corals, mollusks, tube worms, and graylings, salamander fish, galaxiids, silversides, rain- echinoderms (Box 3). It is considered the center of bow fishes, barramundi, grunters, glassfishes, temperate evolution for many of the common coral reef fish fami- perches or basses, sleepers and gobies, and torrentfishes lies that occur in other tropical regions. Only a few (in New Zealand). One major river system occurs on families are endemic to the Indo-West Pacific (e.g., New Guinea (the Fly) and two on Australia (Darling sillaginid whitings and rabbitfishes). Common families and Murray); permanent, large lakes are rare. include moray eels, squirrelfishes, sea basses, grunts,

FIGURE 12 Major oceanic temperature regions and surface current patterns (reproduced with permission from Bond, 1996). FISH, BIODIVERSITY OF 779

Box 3 The Deep Reef Coral reef fishes—those associated with tropical, shallow waters in which coral reefs grow—are the most diverse group of fishes on the planet. Fish diversity is generally correlated with coral species diversity and coral coverage; 2000 fish species, and many other taxa, occur on the Great Barrier Reef of Australia, on which 500 FIGURE 13 The ‘‘Dr. Seuss fish’’—Belonoperca pylei, a 6-cm long, species of coral also occur. However, most esti- brilliantly colored new species of sea bass from the deep reef zone mates of diversity of coral reef fishes are based of Rarotonga. The head and back are bright yellow, the body is pink, on species found only in shallow water less than and the spots are orange (photo by Richard Pyle). 50 m deep, where reef-building corals are abun- dant and where ichthyologists equipped with scuba gear can collect. An underappreciated and 2. The Eastern Pacific Region barely explored component of the diversity of The Eastern Pacific region, with approximately 800 fish reef fishes involves species that live in the deeper species, runs from southern Baja California to Ecuador, (50–150 m) ‘‘twilight zone’’ portion of the reef its northern and southern limits defined by the cold- face, which has only recently been made accessi- water California and Peru currents. Despite its location ble by the use of specialized, mixed-gas rebreath- in the Pacific Ocean, the Eastern Pacific is faunistically ing equipment. The fish fauna of this region con- more similar to the tropical Atlantic, containing many sists of species that seldom occur above 75 m. species that are almost indistinguishable from Atlantic Many new species and even a few new genera forms. The two oceans mixed before the Panamanian have been discovered recently from the deep reef Isthmus formed and the two areas still share 12 species, environment in families that are common in shal- despite 3 million years of physical separation. Most lower water (e.g., wrasses, sea basses, dam- families are less diverse here than in the Western Atlan- selfishes, angelfishes, and gobies). Most of the tic, with the exceptions of sea catfishes, croakers, and recent collecting using advanced diving tech- herrings. Dactyloscopid sand stargazers occur here and niques has occurred in deep reef areas of only in the Atlantic, but not in the Indo-West Pacific. Sixty- Indo-Polynesia (e.g., Rarotonga, Palau, and New two Indo-West Pacific species have managed to cross Guinea), but nearly 100 new species have already the Eastern Pacific Barrier. Three provinces—Mexican, been found (Fig. 13). If we can extrapolate from Panamanian, and Galapagos—are recognized. the few deep reef areas that have been surveyed, our estimates of the number of fishes inhabiting 3. The Western Atlantic Region ‘‘coral reefs’’ will have to be increased consider- The Western Atlantic region is the second most diverse ably. Also, fishes are just part of this barely ex- oceanic area, containing 1200 fish species. It includes plored, intermediate depth zone. How much of Bermuda (which, although at 32ЊN, sits in the tropical marine biodiversity are we missing? Gulf Stream), southern Florida, the Bahamas Bank, the Caribbean Sea, and tropical and temperate portions of South America. Most of the families that occur in the Indo-West Pacific also occur in the Western Atlantic; a few families are more diverse here, such as grunts snappers, cardinal fishes, butterfly fishes, angelfishes, and toadfishes. Strong currents of warm water separate damselfishes, wrasses, parrotfishes, surgeonfishes, go- the Western Atlantic fauna from colder waters along bies, triggerfishes, and boxfishes. Barriers to movement much of its boundaries. It is subdivided into Caribbean, of Indo-West Pacific fishes are cool waters to the north Brazilian, and West Indian provinces. and south and a vast expanse of open, deep water to the east known as the Eastern Pacific Barrier. Eight 4. The Eastern Atlantic Region different provinces have been recognized in the Indo- The Eastern Atlantic region is a relatively small region West Pacific, separating it into a huge Indo-Polynesian along the west coast of Africa from Senegal to Angola region and seven smaller areas. and extending out to oceanic islands such as Ascension 780 FISH, BIODIVERSITY OF and St. Helena. Tropical marine fishes here are limited This area can be divided into four fairly distinct regions by cool-water currents impinging from both the north according to location and temperature: Japanese warm and the south as well as by substantial freshwater run- temperate and Californian warm temperate regions and off and sediments from several major west African riv- Eastern and Western boreal regions. The warm temper- ers, all factors which discourage coral reef growth. The ate areas (from about Hong Kong to Tokyo in the west region contains ‘‘only’’ 500 shore fishes; most coral reef and from lower Baja California to central California in families occur but are represented by only a few species. the east) contain a fauna that fluctuates seasonally, as Porgies are particularly diverse. No subdivisions are tropical species move north in the summer and boreal recognized, with the possible exception of the warm- species move south in the winter. Notable families to temperate Mediterranean. The Mediterranean Sea con- the west include lizard-fishes, flying fishes, mullets, tains 540 species, with many species in the same families jacks, sea basses, and croakers and to the east include as those in the Eastern Atlantic. The Mediterranean endemic silversides, sea basses, croakers, damselfish, has the dubious distinction of being the most heavily wrasses, and flatfishes. The more northerly boreal re- invaded tropical marine area in the world, with at least gions (approximately north from central California in 52 alien fish species having moved in from the Red Sea the east and Korea in the west) contain similar families (an Indo-West Pacific subregion) via the human-made but different species. Important families include migra- Suez Canal. tory salmonids, sculpins, rockfishes, snailfishes, green- lings, gunnels, pricklebacks, and right-eyed flounders. 5. The Arctic Region In the southern Pacific, cold water currents create at The Arctic region encompasses high-latitude (above least three temperate faunal regions, with centers of 60ЊN) waters of both the Pacific and Atlantic Oceans. distribution around Australia, New Zealand, and It reaches from Nunivak Island, Alaska, northward and South America. across the polar region to Newfoundland and Norway In the Atlantic, three northern temperate areas occur: in the northern Atlantic. Of the two polar areas, the the western and eastern Atlantic boreal regions and Arctic is more diverse. Successful groups include skates, the Atlantic warm temperate or Carolinian region. The herrings, salmons, smelts, cods, eelpouts, greenlings, boreal regions (Newfoundland to Cape Hatteras in the sculpins, poachers, snailfishes, pricklebacks, wolffishes, west and British Isles to northern Europe and Scandina- gunnels, and right-eye flounders. Diversity within many via in the east) share a fauna of salmonids, cods, stickle- of these groups is greater in the Pacific than in the backs, poachers, sculpins, wolf-fishes, and right-eyed Atlantic portions of the region. A total of 415 species flounders, with occasional strays from more southerly occur here. Distribution of many of these families ap- waters during warm months. The Carolinian extends pears to be limited by temperature, with warmer waters from Cape Hatteras south to Florida and also to the and currents to the south determining species’ bound- Gulf of Mexico, with southern Florida housing tropical aries. species. Common groups are clupeids, sea robins, pipe- fishes, silversides, needlefishes, killifishes, croakers, 6. The Antarctic Region left-eyed flounders, and puffers. Temperate faunas also The Antarctic region (above 60ЊS) has its own distinc- occur in the southern Atlantic, but their areas and diver- tive fauna that is restricted to Antarctic waters and the sity are less than those of the southern temperate faunas surrounding Southern Ocean, including the cold waters of the Pacific Ocean. Two recognized regions are the of Australia, New Zealand, and nearby oceanic islands. eastern South American and southern African warm Forty-nine families and 274 species occur here, 13 fami- temperate regions. The former region’s fauna includes lies and 174 species of which are identified as Antarctic sea catfishes, croakers, herrings, gobies, scorpion fishes, continent species. A particularly successful group is the and sea basses; the latter area has many colder water notothenioid icefishes and relatives, which account for members of Indo-West Pacific families. 55% of antarctic species. Families include bovichthyids, cod and crocodile icefishes, plunderfishes, and drag- 8. Pelagic Regions onfishes. Non-notothenioids include skates, snailfishes, The 350 species of pelagic fishes occur in ocean surface eelpouts, lantern fishes, eel cods, deep-sea cods, and waters to a depth of 200 m. This habitat type can be southern flounders. divided into 10 different regions based on faunal differ- ences, with more joining of Southern Hemisphere areas 7. Temperate Regions because of the relative lack of large land masses. These To the north of the Indo-West Pacific lie cooler temper- regions are Arctic and Antarctic polar, North Pacific ate waters with their own characteristic fish faunas. cold temperate, North Pacific warm temperate, tropical FISH, BIODIVERSITY OF 781

Indo-Pacific, North Atlantic boreal, North Atlantic ties with increasing depth, which select for an increasing warm temperate, Atlantic tropical, southern warm tem- need to conserve energy and to be able to take advantage perate, and southern cold temperate. Many of the of feeding opportunities. Again, bathypelagic fishes world’s most important fisheries species occur in pelagic have names indicative of their adaptations: sawtooth regions, including numerous sharks, sardines and her- eels, gulper eels, swallower eels, dragonfishes, angler- rings, salmons, codfishes, pollocks, hakes, haddocks, fishes, seadevils, and fangtooths. sauries, mackerels, and tunas. In addition, about 100 Benthal fishes include about 1000 bony fish species species of mostly pelagic fishes have a worldwide distri- in four superorders and nine orders, plus chimaeras bution. This group includes pelagic sharks (white, and squaloid sharks. Different families inhabit bottom whale, tiger, and perhaps megamouth), swordfish, and compared to open-water regions. Benthal fishes include ocean sunfish. greeneyes, tripod fishes, hakes, grenadiers, cusk-eels, batfishes, snailfishes, and eelpouts. Although diversity 9. The Deep Sea decreases below 1000 m, grenadiers and rattails live Waters deeper than 200 m are as much a zoogeographic between 1000 and 4000 m, tripod fish have been found as a habitat entity. The deep sea is generally divided to 6000 m and snailfishes to 7000 m, and some cusk- into three major regions based on depth: the open- eels have been found as deep as 8000 m. water mesopelagic (200–1000 m) and bathypelagic Although some differences in species composition (1000–4000 m) regions and the bottom-associated ben- occur in different ocean basins or in association with thal (200–1000 m) region. Benthal fishes are further different water masses, deep-sea species are relatively divided into benthopelagic fishes that hover just above cosmopolitan, occurring in several different oceans. the bottom and benthic fishes that rest in contact with One trend is for fishes to occur deeper at lower latitudes, the bottom. Deep-sea fishes are most common in these such that species that are bathypelagic near the equator regions between 40ЊN and 40ЊS latitude, approximately may be mesopelagic at middle latitudes and even epipel- between San Francisco and Melbourne in the Pacific agic at the poles. and between New York City and the Cape of Good Hope in the Atlantic. Abyssal and hadal (trench) regions Websites deeper than 4000 m are relatively depauperate. Each region has a characteristic and relatively diverse www.fishbase.org (a remarkable site with information bony fish fauna consisting of fishes from many different on fishes and fisheries worldwide) taxonomic groups. The mesopelagic region worldwide www.fisheries.org/ (website of the American Fisher- contains about 750 species in seven different superor- ies Society) ders and nine orders. Despite their lack of relatedness, www.fishlinkcentral.com (a clearinghouse for the deep-sea fishes share many anatomical and physiologi- aquarium trade) cal features, suggesting independent, convergent evolu- www.fws.gov/r9endspp/endspp.html (U.S. Fish and tion of adaptations to deep-water existence. Mesope- Wildlife Service, Endangered Species homepage) lagic fishes typically have photophores (light organs) on www.utexas.edu/depts/asih/ (website of the American their silvery bodies; have relatively large, often-tubular Society of Ichthyologists and Herpetologists) eyes; undergo daily migrations to surface waters to feed www.wcmc.org.uk (World Conservation Monitoring at night; and have large mouths and long teeth. Com- Centre; lists information on imperiled species world- mon names of mesopelagic fishes reflect these traits: wide) barreleyes, bristlemouths, dragonfishes, sabertooth fishes, lantern fishes, tube-eyes, and swallowers. The bathypelagic region is the largest habitat space See Also the Following Articles on the earth, accounting for 88% of oceanic volume. • • The five superorders, nine orders, and 200 species in FISH CONSERVATION FISH STOCKS LAKE AND POND ECOSYSTEMS • OCEAN ECOSYSTEMS • PELAGIC ECOSYSTEMS the cold, dark bathypelagic region share some traits with mesopelagic fishes but possess them in the ex- treme. Photophores are concentrated on lures used to attract prey; eyes are often small; mouths are extremely Bibliography large and teeth very long; stomachs are expandable; Berra, T. M. (1981). An Atlas of Distribution of the Freshwater Fish bodies are black; and body musculature, bones, and Families of the World. Univ. of Nebraska Press, Lincoln. scales are greatly reduced. These traits reflect greater Bigelow and Schroeder (1953). Fishes of the Gulf of Maine. Fish Bull. habitat space and increasingly rare feeding opportuni- Fish Wildlife Serv. 53 (Fish Bull. 74). 782 FISH, BIODIVERSITY OF

Bond, C. E. (1996). Biology of Fishes, 2nd ed. Saunders, Fort Atlantic, Part 7, Salmon, trouts, and others. Mem. Sears Foundation Worth, TX. Mar. Res. 1, 111–145. Bone, Q., Marshall, N. B., and Blaxter, J. H. S. (1995). Biology of Miller et al. (1989). Extinctions of North American Fishes during Fishes, 2nd ed. Blackie, London. the past century. Fisheries 14(6), 22–38. Briggs, J. C. (1995). Global Biogeography. Elsevier, Amsterdam. Moyle, P. B., and Cech, J. J., Jr. (1996). Fishes: An Introduction to Burgess (1989). An Atlas of Freshwater and Marine Catfishes. TFH Ichthyology, 3rd ed. Prentice-Hall, Upper Saddle River, NJ. Publishers, Neptune City, New Jersey. Musick et al. (1991). The biology of Latimeria chalumnae and evolu- Compagno, L. J. V. (1984). FAO species catalogue. Vol. 4: Sharks of tion of coelacanths. Environ. Biol. Fish 32, 1–435. the world. An annotated and illustrated catalogue of shark species Nafpaktitis et al. (1977). Family Myctophidae. In: Fishes of the West- known to date. FAO Fisheries Synopsis No. 125. Food and Agri- ern North Atlantic. Mem. Sears Foundation Mar. Res. 1(7), 13–265. culture Organization, Rome. Nelson, J. S. (1994). Fishes of the World, 3rd ed. Wiley, New York. Compagno, L. J. V. (1990). Environ. Biol. Fish 28, 33–75. Norman, J. R. (1905). A history of fishes. Eschmeyer, W. N. (Ed.) (1998). Catalog of Fishes. California Academy Paxton, J. R., and Eschmeyer, W. N. (Ed.) (1998). Encyclopedia of of Sciences, San Francisco. Fishes, 2nd ed. Academic Press, San Diego. Greenfield, and Thomerson (1997). Fishes of the Continental Waters Scott and Crossman (1973). Freshwater fishes of Canada. Fish Res. of Belize. Univ Press of Florida, Gainesville, Florida. Board Can. Bull. 184, 1–920. Helfman, G. S., Collette, B. B., and Facey, D. E. (1997). The Diversity Suttkus (1963). Order Lepisostei. In: Fishes of the Western North of Fishes. Blackwell Science, Malden, MA. Atlantic, Part 3, Soft-rayed bony fishes. Mem. Sears Foundation Heller and Snodgrass (1903). Papers from the Hopkins Stanford Gala- Mar. Res. 1, 61–88. pagos Expedition, 1898–1899. XV. New fishes. Proc. Wash. Acad. Vladykov, and Greeley (1963). Order Acipenseroidei. In: Fishes of Sci. 5, 189–229. the Western North Atlantic, Part 3, Soft-rayed bony fishes. Mem. Hildebrand (1963). Family Elopidae. In: Fishes of the Western North Sears Foundation Mar. Res. 1(3), 24–60.