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Reis, Roberto E. and Helfman, Gene S. (2024) Fishes, Biodiversity of. In: Scheiner Samuel M. (eds.) Encyclopedia
of Biodiversity 3rd edition, vol. 2, pp. 341–368. Oxford: Elsevier.
http://dx.doi.org/10.1016/B978-0-12-822562-2.00092-X
© 2024 Elsevier Inc. All rights reserved.

Author's personal copy
Fishes, Biodiversity of
Roberto E Reis, Pontifical Catholic University of Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil
Gene S Helfman, University of Georgia, Athens, GA, United States
r 2024 Elsevier Inc. All rights reserved.
Introduction 342
Taxonomic Diversity 343
Overview 343
Agnathans 344
Hagfishes 344
Lampreys 344
Gnathostomes 347
Chondrichthyans 347
Holocephalans 347
Elasmobranchs 347
Sharks 347
Skates and Rays 348
Osteichthyans 349
Sarcopterygians 349
Coelacanths 350
Dipnoans 350
Lungfishes 350
Tetrapods 350
Actinopterygians 351
Bichirs and Reedfish 351
Sturgeons and Paddlefishes 351
Holosteans 352
Gars and Bowfins 352
Teleosts 352
Osteoglossomorphs 353
Elopomorphs 353
Otocephalans 353
Ostariophysans 354
Euteleosteans 355
Acanthomorphs 356
Acanthopterygians 357
Percomorphs 357
Geographic Diversity 361
Overview 361
Freshwater Diversity 362
The Nearctic Region 362
The Neotropical Region 364
The Palearctic Region 364
The African or Ethiopian Region 365
The Oriental Region 365
The Australian Region 365
Marine Diversity 365
The Indo-West Pacific Region 365
The Eastern Pacific Region 366
The Western Atlantic Region 366
The Eastern Atlantic Region 366
The Arctic Region 366
The Antarctic Region 366
Temperate Regions 366
Encyclopedia of Biodiversity 3rd edition, Volume 2 doi:10.1016/B978-0-12-822562-2.00092-X 341

342 Fishes, Biodiversity of
Pelagic Regions 367

The Deep Sea 367
Conclusion 367
References 368
Abstract
A fish is an aquatic vertebrate with scales, fins, and gills – most of the times! Over 36,400 fish species are known, divided into jawless
fishes (agnathans), cartilaginous fishes (chondrichthyans), ray-finned fishes (actinopterygians), and lobe-finned fishes (sarcopterygians).
Fishes occur wherever water of reasonable integrity exists, from sea depths of 8000 m to mountain streams and lakes at 5000 m
altitude. Marine fishes constitute 50% of all species, freshwater species 49%, with 1% of fishes moving regularly between sea and fresh
water. Tropical areas have the highest diversity. Fish biodiversity is threatened by many human activities, but habitat modification,
overharvest, and introduced species are particularly injurious.
Glossary
Adipose fin A small, fleshy fin usually without supporting spines or rays, set far back on the dorsal surface of many catfishes,
characins, salmons, and other groups.
Anadromy A form of diadromy where a fish is born in fresh water, matures in the ocean, and returns to fresh water to spawn
(lampreys, sturgeons, salmons). Contrast with catadromy.
Ancestral The taxon from which descendant species are derived, often synonymized as primitive. Ancestral traits or
conditions are those which appear in an ancestor.
Catadromy A form of diadromy where a fish is born in the ocean, matures in fresh water, and returns to the ocean to spawn
(freshwater eels, some mullets). Contrast with anadromy.
Clade (¼ monophyletic group) A complete group of organisms closely related phylogenetically; a group that shares a
common ancestor and all its descendants.
Depauperate Of low diversity, lacking in species; opposite of speciose.
Derived Later-appearing taxa within a lineage, often synonymized as advanced. Derived traits are those which appear in a
descendant species and are changed from the ancestral condition.
Diadromy Migration between marine and freshwater environments that some fish species undertake during specific periods
in their life cycle.
Endemic Restricted or native to a geographically defined area.
Extant Living; opposite of extinct.
Phylogeny A tree-like diagram depicting the evolutionary relationships among terminals.
Speciose Of high diversity, having many species in a group or area.
Taxon (plural ¼ taxa) A group of evolutionarily related species with a taxonomic name.
Key Points
• There are currently over 36,400 species of fishes and approximately 200–300 new species are described each year.
• 50% of all fishes are marine, 49% are freshwater, and 1% are diadromous.
• The highest diversity occurs in tropical waters both marine and freshwater.
• Fishes are not a natural (monophyletic) group and some fishes are more related to tetrapods than to other fishes.
• Fishes are classified in four different large group, jawless, cartilaginous, ray-finned, and lobe-finned fishes, the latter being
more closely related to us than to other fishes.
• Both marine and freshwater fishes are increasingly more threatened with extinction because of human activities. Over-
fishing and bad fishing practices are the main cause for marine stocks, while deforestation, hydropower dams, pollution,
and invasive species are the leading cause of extinction for freshwater species.
Introduction
More than 36,400 fish species occur in virtually all water habitats on Earth, from sea depths to mountain creeks. Marine habitats harbor
slightly more than half of all fish species, with tropical areas having the highest diversity. The different groups of fishes are classified in four
main groups, the jawless hagfishes and lampreys, the cartilaginous sharks, skates, rays, and chimaeras, the lobed-finned coelacanths and
Fishes, Biodiversity of 343
lung-fishes, and the ray-finned fishes, a large assemblage of roughly 35,000 species. Ray-finned fishes are highly diverse taxonomically and
ecologically, occupying every aquatic habitat type and niche imaginable. This amazing diversity is classified into seven taxonomic
subdivisions that reflect their evolutionary patterns. These seven main subdivisions are the polypteriforms (African bichirs), the aci-
penseriformes (sturgeons and paddlefishes), the holosteans (gars and bowfin), the osteoglossomorphs (bonytongues), the elopomorphs
(eels and tarpon), the otocephalans (herrings, minnow, tetras, catfishes, and relatives), containing most freshwater fishes worldwide, and
the euteleosts, containing the majority of modern bony marine fishes. The most important zoogeographic distinction splits marine and
freshwater fishes. Approximately 49% of all fishes are restricted to pure fresh water, while 50% occur only in normal oceanic salinity, with
the remaining 1% moving regularly between the two salinity designations. Most freshwater species are primary or obligatory freshwater
dwellers and are usually restricted to specific river basins. Among marine fishes, the vast majority live in shallow, warm areas such as coral
reefs. The remaining marine species are divided fairly evenly among shallow cold, open-ocean, and deep-bottom areas.
Taxonomic Diversity
Overview
Over 36,400 known fish species inhabit the oceans, estuaries, rivers, lakes, and streams of our planet. This incredible diversity
almost equals that of all other vertebrate groups combined (Box 1). It forms a wealth of biological wonder for ichthyologists (fish
scientists), but such large numbers can be overwhelming to someone unfamiliar with the many taxonomic groups and their
names. However, the different taxonomic groups are logically arranged according to well-studied evolutionary relationships, with
those more closely related to groups that diversified earlier in geologic time (so-called primitive taxa) placed earlier in lists and
those groups that evolved relatively recently (derived or advanced taxa) placed later in lists (Nelson, 2006).
The earliest derived of the living fishes are the jawless fishes. These include about 87 species of marine hagfishes and 49
freshwater or anadromous (migratory between freshwater and marine) lampreys. Cartilaginous sharks, skates, rays, and chimaeras
include approximately 14 orders, 70 families, and 1300 species of almost entirely marine, relatively large-bodied predators. Skates
and rays are more diverse than sharks, constituting about 55% of all cartilaginous fishes. Chimaeras are cartilaginous fishes closely
related to sharks and rays that consist of three families and 56 species.
Bony fishes make up the vast majority of living fish species, exceeding all other groups in species, habitat, reproductive, and
feeding diversity (Fig. 1). Bony fishes vary in length from the 8 mm pygmy gobies and Indonesian minnows to the 12 m long Oarfish
(Regalecus glesne), 900 kg marlins, and 1000 kg plus Ocean Sunfish (Mola mola). There are 51 orders and 543 families among the
approximately 35,000 species of bony fishes, ranging from the primitive bichirs, sturgeons, paddle-fish, gars, and Bowfin (Amia calva)
to the more derived teleostean (higher bony fish) groups that include bonytongues, eels and tarpons, herring-like fishes, the so-called
true teleostean groups of minnows, salmons, various deepsea taxa, and cods, in addition to those more closely related to tetrapods,
the coelacanths and lungfishes. The spine-finned teleosts are the most evolutionarily derived of the teleost fishes and include mullets,
silversides, scorpionfishes, perch-like fishes, tunas, flatfishes, puffers and many others (Helfman and Collette, 2011).
The classification presented below is mostly based on those of Betancur et al. (2017) and Dornburg and Near (2021) for bony
fishes and the website https://sharksrays.org/ by Gavin Naylor for chondrichthyans. Follow the classification of fishes presented
below in the phylogeny given in Fig. 2.
Phylum Chordata, Subphylum Craniata, Superclass Agnatha, Class Myxini, Order Myxiniformes: living hagfishes; one family,
87 species, temperate marine; Class Cephalaspidomorphi, Order Petromyzontiformes: living lampreys; three families, 49 species,
temperate fresh water and anadromous.
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. Exceptions to and variations in all traits are common (Paxton and Eschmeyer, 1998). Some sharks, tunas, and billfishes are
warm-blooded. Fins can be unsegmented and spiny or segmented and soft rayed (and soft rays can be hardened). Tail types include: (1)
the heterocercal tail of sharks and sturgeons, in which the notochord or vertebral axis extends considerably into the upper tail lobe; (2) the
abbreviate heterocercal tail of gars and Bowfin, in which the vertebral axis extends only slightly into the upper lobe; (3) the leptocercal or
diphycercal tail of lungfishes, the coelacanths, and rattails, in which vertebrae or tail rays extend through the middle of the tail, forming a
pointed tail; and (4) the homocercal tail of most derived bony fishes, in which the vertebral column ends at the tail base (the urostyle) and
fin rays form a symmetrical, two-lobed tail. Scales also vary in terms of the number of layers of bony material that constitute them and the
extent of spiny projections that cover their surface; more primitive fishes generally have heavier scales, and more derived fishes have
lighter scales, often with more projections. Scale types include the placoid scales of sharks, the ganoid scales of gars, bichirs, and
sturgeons, the cycloid scales of lower teleosts, and the ctenoid scales of higher teleosts. But unlike familiar groups of vertebrates such as
lizards, frogs, birds, or mammals that represent monophyletic groups or clades – those groups with one ancestral and all its descendant
species, fishes indeed have one ancestral, but not all descendants of that ancestral species are included. So, without the tetrapods, fishes
are not monophyletic or a natural group. Hence our definition can include hagfishes and lampreys, sharks, coelacanths, lungfishes, and
bony fishes, but it only becomes monophyletic when tetrapods are also included – in conclusion, we are all fishes! To ichthyologists, “fish”
refers to one or more individuals of a single species, whereas “fishes” refers to more than one species, regardless of how many individuals
are involved. Hence, this article is about fishes (Moyle and Cech, 2004; Barton, 2006; Helfman et al., 2009).
Soft-rayed dorsal fin (second dorsal)

Spinous dorsal fin (first dorsal)
Dorsal spines
Soft ray Caudal or tail fin
Operculum or gill cover

Adipose fin
Dorsal surface
Snout
Caudal
Lateral line Peduncle

Pectoral fin
Ventral surface Finlets
Barbel Anal spines Anal fin

Pelvic fin
Fig. 1 Anatomical features of a hypothetical bony fish. Reproduced from Squire Jr., J. L. and Smith, S. E. (1977) Angler’s guide to the United
States Pacific coasts. Washington, DC: National Marine Fisheries Service.
Agnathans
The first fishes lacked jaws. Modern jawless fishes – hagfishes and lampreys – look approximately similar, with slippery, eel-like
bodies and jawless heads. Despite biologists disputed for decades whether lampreys and hagfishes are closely related or lampreys
are more closely related to gnathostomes (vertebrates with jaws), latest evidence suggest that hagfishes and lampreys do form a
clade (Cyclostomata) at the base of Vertebrates. Despite they are vertebrates, hagfishes lack vertebrae, relying on the notochord for
support. Lampreys, although having a well-developed, functional notochord, show simple vertebrae.
Hagfishes
Hagfishes, known also as slime eels or slime hags, produce copious mucus from many pairs of slime glands (Fig. 3). A couple of
disturbed 2-foot-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 knot in its tail and passes the knot forward until the slime is pushed off. Hagfishes are nocturnal
predators on small invertebrates but are better known for their scavenging behavior, which involves burrowing into a dead or
dying fish and consuming the prey from the inside.
Hagfishes occur almost worldwide in temperate and cold-temperate oceans, usually in water deeper than 30 m. Hagfishes can reach high
densities upwards of 0.5 m2 on soft-bottom marine areas, which is the most abundant habitat type in the world. Hence, hagfishes could be
ecologically important as predators and scavengers. They are also common in the diets of seals and sea lions. Hagfishes are commercially
important because their hides are the popular “eelskin” of wallets, purses, and briefcases. Overfishing has depleted hagfish stocks in Korea
and Japan, and new fisheries are being exploited, and probably overexploited, in the eastern Pacific and western Atlantic. This unfortunate
chain of events has characterized marine fisheries worldwide (Helfman, 2007). Drastic reductions in hagfish populations brought on by
overfishing could potentially disrupt a widespread ecosystem; not enough is known about hagfish ecology to predict these impacts.
Lampreys
Lampreys go through a long-lived larval phase; the free-living, blind, toothless larva called ammocoete, lives in silty streambeds
where it filters microscopic organisms from the water for up to seven years before transforming into an adult. In brook lampreys,
larvae transform into nonfeeding adults, live for few months, spawn, and die. Other species transform into feeding adults and live
for 1–3 years as parasites on other fishes. They rasp holes in their host’s skin and live off its body fluids (Fig. 4). Accidental
introduction of the parasitic Sea Lamprey (Petromyzon marinus) into the Great Lakes of North America has contributed to the
decline of Lake Trout, whitefishes, and Blue Pike.
Lampreys are cool-water species (301 north and south latitude or higher). Most lampreys live in fresh water, but some parasitic
species are anadromous. Brook lampreys typically live in headwater streams, an ecosystem type frequently disrupted by human
activities. Hence, several US brook lampreys are imperiled. North America’s smallest lamprey, the Miller Lake Lamprey
(Entosphenus minimus), was poisoned almost into extinction because it parasitized introduced trout in its only habitat – Miller
Lake, Oregon. The southern hemisphere is comparative poorer in lamprey diversity. Only six species of mordaciid and geotrid
lampreys occur in southern South America, Australia, and New Zealand, while 43 petromyzontid lampreys inhabit the northern
hemisphere.
Myxiniformes
Petromyzontiformes
Agnatha
Chimaeriformes Holocephali
Gnathostomata
Selachii Chondrichthyes
Elasmobranchii
Batoidea
Coelacanthiformes Sarcopterygii
Osteichthyes
Ceratodontiformes
Dipnoi
Tetrapoda
Polyteriformes
Acnopterygii
Acipenseriformes
Lepisosteiformes Holostei
Amiiformes
Osteoglossiformes
Osteoglossomorpha
Teleostei
Hiodontiformes
Elopiformes
Albuliformes Elopomorpha
Notocanthiformes
Anguilliformes
Clupeiformes Otocephala
Alepocephaliformes
Gonorhynchiformes Ostariophysi
Cypriniformes Otophysi
Gymnotiformes
Cithariniformes
Siluriformes
Characiformes
Lepidogalaxiiformes
Euteleostei
Argentiniformes
Esociformes
Salmoniformes
Stomiiformes
Osmeriformes
Galaxiiformes
Ateleopodiformes
Aulopiformes
Myctophiformes
Lampriformes
Acanthomorpha
Percopsiformes
Polymixiiformes
Zeiformes
Gadiformes
Stylephoriformes
Trachichthyiformes Acanthopterygii
Beryciformes
Ophidiiformes Percomorpha
Batrachoidiformes
Gobiiformes
Scombriformes
Syngnathiformes
Blanniiformes
Carangiformes
Synbranchiformes
Perciformes
Centrarchiformes
Labriformes
Acropomatiformes
Acanthuriformes
Fig. 2 Phylogeny of fishes at Order level, compiled from recent literature. The tree reflects the most recent understanding of fish relationships.
Fig. 3 Hagfishes, also called slime eels, produce prodigious amounts of mucus when disturbed. Photo by Meyer JL.
Fig. 4 Mouth region of a lamprey. Lampreys attach to the sides of larger fish and scrape a hole with their tongue, feeding on body fluids.
Gnathostomes
The appearance of mandibles was one of the most important evolutionary events that shaped vertebrate life. Mandibles and its
suspensorium (the connections to the skull), originated from the anterior branchial arches of the agnathans.
Chondrichthyans
Superclass Gnathostomata, Class Chondrichthyes, Subclass Holocephali (Chimaeras) and Subclass Elasmobranchii (sharks, skates,
and rays). Elasmobranchs include around 1300 species that live in all world’s oceans; a few live in fresh water (Carrier et al., 2004,
2010). Elasmobranchs are characterized by a cartilaginous skeleton hardened by calcium deposits and usually five (sometimes six
or seven) gill slits. They lack lungs or gas bladders but instead have large, oil-filled livers which may aid in maintain neutral
buoyancy. Their teeth and pedestal-like placoid scales develop from the same embryonic structures. Teeth are continually lost and
replaced, and a shark may produce as many as 30,000 teeth during its lifetime. All elasmobranchs have internal fertilization, and
many bear live young that are nourished by the mother via different strategies, including a complex umbilicus and placental
structure analogous to that found in mammals. Slow growth, late maturation, and low reproductive output make many elas-
mobranchs exceptionally vulnerable to human exploitation and prone to extinction.
Holocephalans
Class Chondrichthyes, Subclass Holocephali, Order Chimaeriformes: three families, 56 species, chimaeras. Chimaeras, also known
as rat- or rabbitfishes, share a cartilaginous skeleton and other features with elasmobranchs. They differ by having (1) the upper
jaw permanently attached to the braincase, (2) continually growing tooth plates in the jaws instead of replaceable teeth, (3) a
single gill flap instead of five or more gill slits, and (4) no scales. Chimaeras swim by flapping their pectoral fins (something sharks
never do) and by undulating their bodies (Fig. 7). All chimaeras are egg-layers, the egg being protected by a horny shell. Adult
chimaeras range in size from 60 to 200 cm. Chimaeras are cool-water, marine fishes that live at shallow to moderate depths
between 80 and 2600 m, where they usually swim just above the bottom. Chimaeras eat predominantly hard-bodied benthic
invertebrates, which they crush with their tooth plates. Surprisingly little is known about their general biology and natural history.
Elasmobranchs
Subclass Elasmobranchii: sharks and rays. Division Selachii: sharks. Order Heterodontiformes: one family, nine species, bullhead
and horn sharks. Order Orectolobiformes: seven families, 45 species, including wobbegongs, nurse, and whale sharks. Order
Carcharhiniformes: 10 families, 298 species, including catsharks, requiem, and hammerhead sharks. Order Lamniformes: eight
families, 16 species, including ragged tooth, Tiger, Megamouth, thresher, Basking, and mackerel sharks. Order Hexanchiformes:
two families, seven species, Frill and cow sharks. Order Echinorhiniformes: one family, two species. Order Squaliformes: six
families, 143 species, including sleeper and dogfish sharks. Order Squatiniformes: one family, 22 species, angel sharks. Order
Pristiophoriformes: one family, 10 species, saw-sharks. Division Batoidea skates and rays. Order Torpediniformes: five families, 68
species, torpedo electric rays and numbfishes. Order Rhinopristiformes: five families, 71 species, banjo rays, guitarfishes, wed-
gefishes, and sawfishes. Order Rajiformes: four families, 302 species, guitarfishes, skates. Order Myliobatiformes: 12 families, 244
species, stingrays, butterfly rays, eagle rays, and manta rays.
Sharks
Approximately 530 shark species are alive today. Sharks are generally large (41 m), predatory fishes. The diverse requiem or
ground sharks (carcharhiniforms) include the Tiger, Gray Reef, Bull, Blue, Lemon, and hammerhead sharks. Lamniform mackerel
sharks are primarily offshore, pelagic inhabitants, although White Sharks spend considerable time close to beaches and small
islands, especially where seals and sea lions abound. The squaliform dogfishes, the second largest shark order, are most successful
and abundant in the North Atlantic, North Pacific, and deepsea regions.
Sharks range in size from the 15 g, 16–20 cm Dwarf Lanternshark (Etmopterus perryi) to the 12,000 kg plus, 15 m long Whale
Shark (Rhincodon typus), the largest fish in the world. White Sharks (Carcharodon carcharias) as large as 6 m and over 3300 kg are
known; larger individuals are suspected. Other large sharks include Basking Sharks (Cetorhinus maximus, 9.8 m), Great Ham-
merheads (Sphyrna mokarran, 5.5 m), Greenland Sharks (Somniosus microcephalus, 6.4 m), Tiger Sharks (Galeocerdo cuvier, 5.9 m),
and Megamouth (Megachasma pelagios, 5.4 m).
Sharks inhabit all oceans except the Antarctic. Depth records for sharks are held by the Portuguese Shark (Centroscymnus
coelolepis) at 3690 m and an unidentified dogfish at 4050 m. A few carcharhinid sharks enter fresh water; Bull Sharks (Carcharhinus
leucas) have been captured 4200 km up the Amazon River in Peru and 1200 km up the Mississippi River. Large pelagic sharks may
cross entire ocean basins; Blue Sharks (Prionace glauca) have been tracked across the North Atlantic Ocean and back, a distance of
16,000 km, and a White Shark moved from South Africa to Western Australia and back to South Africa over a nine month period, a
distance of 22,000 km.
Most sharks are predatory on large prey, but three of the largest sharks – the Basking, Megamouth, and Whale sharks – feed on
zooplankton. A small, 40 cm long, midwater species, the Cookie-Cutter Shark (Isistius brasiliensis), is an ectoparasite on the sides of tunas,
dolphins, whales, an occasional Megamouth Shark, and even rubber sonar domes of nuclear submarines. Some sharks use structures other
than jaw teeth to capture prey. Thresher sharks use the long upper lobe of their tails to stun schooling prey. Saw-sharks (and rhino-
pristiform sawfishes) have elongate, blade-like snouts studded with lateral teeth which they slash laterally to disable prey. Hammerheads
use their broadened hammer-shaped head to pin stingrays against the bottom before biting chunks out of the rays’ wings.
Sharks are sensitive to chemicals, able to detect one part fish extract per 10 billion parts seawater. In addition, because their
nostrils are well separated, they can detect the direction a smell comes from, something humans cannot do. Sharks have good
vision, although they tend to be slightly myopic (farsighted). Sharks are also highly sensitive to sounds, including infrasonic sound
below 10 Hz. Sharks can also locate prey by detecting the weak electric fields their prey emit using their ampullae of Lorenzini. This
electro-sensitivity may also allow them to navigate using the earth’s geomagnetic fields. Many sharks have relatively large brains,
with brain to body weight ratios comparable to those of some birds and mammals.
Sharks grow slowly and live long: Spiny Dogfish live 70–100 years, Lemon Sharks (Negaprion brevirostris) 50–60 years, and the
Greenland Shark is thought to live for up to 400 year, the longest life-span of any vertebrate. Sharks also reproduce slowly. Lemon
Sharks may not mature until they are 24 years old, and Spiny Dogfish may not mature until they are 35 years old. Sharks produce
relatively few, large young with a long gestation period. Clutch size varies from one or two live young (Ragged Tooth, threshers,
and makos) to 300 in the Whale Shark. Gestation periods average 9–12 months but may be two years in Spiny Dogfish and 3.5
years in Basking Sharks. Bullhead and nurse sharks lay eggs, but most sharks bear live young. The requiem sharks have the most
complex developmental pattern, in which an umbilical cord connects mother and embryo, transporting nutrients and oxygen to
the embryo and carrying metabolic wastes to the mother.
Because many sharks mature late, reproduce at long intervals, and have low reproductive output, shark populations are easily
and frequently overfished (Fig. 5). Thresher sharks, School Sharks (Galeorhinus galeus), Spiny Dogfish, Porbeagles (Lamna nasus),
Basking Sharks, and Bull Sharks are all examples of shark stocks that have been overexploited. White Sharks are protected in
Australia, South Africa, and the US; capture and sale of White Sharks, Whale Sharks, and Basking Sharks is restricted under the
Convention on International Trade in Endangered Species (CITES).
Skates and Rays

The batoid skates and rays are 707 species of mostly benthic (bottom-living), mostly marine forms. In skates and rays, the pectoral
fins are fused to the sides of the head and the five gill slits are displaced to the ventral surface. Skates are most diverse in deep water
and at high latitudes, whereas stingrays are most diverse in tropical, inshore waters. Some batoids live much or all of their lives in
fresh water. Largetooth Sawfish frequently swim up rivers in Central and South America. Two stingray families contain entirely
freshwater species – the river stingrays of South America family Potamotrygonidae and several species in the large stingray family
Dasyatidae. Species of the latter inhabit African, Southeast Asian, and New Guinea rivers.
Skates and rays feed mostly on benthic invertebrates, except for the huge (up to 6-m wide) manta rays, which capture small crustaceans
and fishes in the water column. Torpedo rays stun prey with powerful electrical discharges (50 V and 50 A¼ 1 kW output). Batoids
Fig. 5 Ragged Tooth Sharks, also called gray nurse and sand tiger sharks, have suffered declines because of fishing and as by-catch in nets.
Ragged Tooth sharks are protected in several countries. “Raggies” are relatively docile, adapt well to captivity, and are often featured in public aquaria.
Fig. 6 A mermaid purse (egg case) of a Big Skate, Raja binoculata, from the North Pacific. Each case holds a single embryo, which can be seen
because part of the egg shell has been removed. Some skates are overfished because they are born at a size that is already too large to escape
through the mesh of drag nets.
reproduce by either laying eggs (skates) or bearing live young (rays). Embryonic skates develop inside “mermaid purse” egg cases for as
much as 15 weeks (Fig. 6).
Skates in some sites are actually increasing in number because of overexploitation of competing bony fishes, such as cod in the
North Atlantic. However, the giant Barndoor Skate of the northwest Atlantic and its relative, the ironically named Common Skate
of the northeast Atlantic, are caught incidental to bottom trawling for bony fishes; they have been seriously depleted and may face
extinction. Largetooth Sawfish have been drastically overfished and all five species are threatened with extinction.
Osteichthyans
Modern bony fishes, or Osteichthyes (literally “bony fishes”), consist of two major clades, the lobe-finned and the ray-finned
fishes. Many living members belong to groups that were much more diverse during Paleozoic and Mesozoic eras. Several of these
fishes are classified as “bony” even though they have cartilaginous skeletons. Their skeletal condition is actually a derived trait;
their immediate ancestors were bony. The classification of bony fishes has changed considerably in the later years, more so after the
widespread use of genomic information by systematists.
Sarcopterygians
Superclass Sarcopterygii, Class Coelacanthimorpha, Order Coelacanthiformes: one family and two species, the coelacanths. Class
Dipnotetrapodomorpha, Subclass Dipnomorpha, Order Ceratodontiformes: three families and six species, the Australian lungfish,
the South American lungfish, and four African lungfishes. Subclass Tetrapodomorpha (amphibians, mammals, and reptiles
[including birds]).
Fig. 7 A Pacific Ratfish, Hydrolagus colliei, showing the beak-like mouth, flexible pectoral fin, and lack of scales that set chimaeras apart from
sharks and rays. The large, reflective eye is characteristic of fishes that live in deep, dark waters.
Coelacanths
Coelacanths were thought to have gone extinct by the end of the Cretaceous, 65 million years ago, until a live one was trawled up
in 1938 off South Africa. Today, one small population of 200–600 coelacanths, Latimeria chalumnae, lives at 100–500 m depths off
two small volcanic islands in the Comoros Archipelago, between Madagascar and Mozambique. Other populations of the same
species have been discovered along Africa’s eastern coast from South Africa to Tanzania. In 1998, another species, L. manadoensis,
was discovered at similar depths in northern Indonesia. The living coelacanths have fleshy, lobed pectoral, pelvic, anal, and second
dorsal fins ( ¼ the lobed fins that define the class Sarcopterygii); a symmetrical, three-lobed tail with a central extension; hollow
neural spines (hence “coelacanth” or “hollow spines”); a unique unconstricted notochord; a joint in the dorsal braincase that aids
jaw opening; relatively large, thick, bony scales; and live young.
Coelacanths are large (to 180 cm, 95 kg), long-lived (probably past 100 years), and produce relatively few, live young (five to
26 young per clutch). The gestation period is about 13 months. Replacement rate in a population is therefore slow. Coelacanths
are captured primarily as by-catch in the hook-and-line fishery for Oilfish (Ruvettus pretiosus), and it is unlikely that the Comoran
population can sustain even the current by-catch rate of five to 10 animals per year. Counts from small submarines indicate the
Comoran species is declining. The Comoran government has outlawed its capture, and trade in coelacanths is outlawed by the
Convention on the International Trade in Endangered Species (CITES).
Dipnoans
Lungfishes
Lungfishes today are represented by three families and six species, all located on the former Gondwanan continents of Australia,
South America, and Africa. Lungfishes lack jaw teeth but have unusual tooth plates on the mouth roof and floor. The Australian
species, Neoceratodus forsteri, is limited to four river systems of northeastern Australia. It is large (1 m), with large scales, flipper-like
fins, a broad tail, and a single lung. It crushes benthic crustaceans, mollusks, and small fishes with its tooth plates. It can, but does
not have to, breathe atmospheric oxygen. Its young lack external gills. Neoceratodus populations have declined dramatically and the
fish is protected; species recovery efforts include transplantation into several Queensland reservoirs and rivers.
The one South American and four African lungfishes have eel-like bodies; slender, almost-filamentous paired fins; lack scales;
have paired lungs; have larvae with external gills; and must breathe air to survive. The four African species occur across central and
south Africa, often in swampy areas that frequently experience drought (Fig. 8). When a swamp dries up, African lungfishes dig a
burrow and can wait four years for rains to return. The South American species occurs in swampy regions of the Amazon and
Paraguay river basins. Comparatively little is known about its biology.
Tetrapods
Paleontologic, anatomic, and genetic evidence indicate that the 37,600 species of tetrapods (modern amphibians, reptiles
[including birds], and mammals) are in reality a subclass of the lobe-finned sarcopterygians. An exciting, confirming fossil
discovery, the “fishapod” Tiktaalik roseae, was unearthed in 2005 in Nunavut Territory of Arctic Canada. Tetrapods are therefore a
sideline taxon of fishes that moved onto land and into the air, with some groups such as turtles, whales, penguins, and seals
returning to an aquatic existence. We are all fish!.
Fig. 8 African lungfishes, Protopterus, estivate, burrowing into mud and remaining inactive for years while waiting for rains to return. Photo
courtesy of Chapman L and Chapman C.
Actinopterygians
Superclass Actinopterygii, Class Cladistia, Order Polypteriformes: one family, 14 species, bichirs and Reedfish; Subclass Chondrostei, Order
Acipenseriformes: two families, 29 species, sturgeons and paddlefishes; Subclass Neopterygii, Infraclass Holostei, Order Lepisosteiformes:
one family, seven species, gars; Order Amiiformes: one family and one species, the Bowfin, and the hyper diverse Teleostei.
Bichirs and Reedfish

The polypteriform bichirs and Reedfish of west and central tropical Africa have been difficult to assign to any particular taxonomic group.
Their gilled larvae, lobe-like fins, thick ganoid scales, and modified heterocercal tail suggest affinities with several living groups, particularly
the chondrostean sturgeons and paddlefishes. However, their unique dorsal, caudal, and paired fins and unusual chromosomes place them
apart from all extant groups. Their placement near the base of the bony fish lineage is justified by egg structure, genetics (mitochondrial,
nuclear, and other DNA and amino acid sequences), and head skeleton. Taxonomists now place them in their own Class, the Cladistia.
Thirteen species of polypteriforms are called bichirs (Polypterus) and an 14th, elongate species is the Reedfish or Ropefish,
Erpetoichthys calabaricus. Bichirs reach 120 cm in length and Reedfish 90 cm. All are predatory and inhabit shallow, swampy
regions, where they breathe atmospheric oxygen with their paired lungs. Bichirs, also called “flagfins”, have unique dorsal and
pectoral fins. Each dorsal finlet consists of a vertical spine with attached horizontal rays, looking like a flagpole with streaming
banners. In more usual ray-finned fishes, dorsal fin rays emerge vertically from the body of the fish. The polypteriform pectoral fin
has an internal arrangement that involves a wishbone-shaped, flattened plate, again unlike that in any other fish. Bichirs are not
particularly well studied; no evidence exists to suggest Polypterus species are imperiled. Erpetoichthys calabaricus is currently assessed
as Near Threatened globally, but is Endangered in parts of Africa.
Sturgeons and Paddlefishes

The next-most primitive actinopterygian fishes are the acipenseriform sturgeons and paddlefishes. All 27 species of sturgeons live
in the Northern Hemisphere. All spawn in fresh water, although some species move seasonally between marine and fresh water.
North American freshwater species include the Lake Sturgeon and three river sturgeons. Anadromous species include the Atlantic
and White sturgeons, the latter being an occupant of west coast bays and rivers. White Sturgeons attain the largest size of any North
American freshwater fish (3.8 m, 630 kg). The world’s largest freshwater fish is the Beluga Sturgeon of eastern Europe and Asia,
Huso huso, at 8.6 m and 1300 kg (Fig. 9).
Sturgeons have four barbels ahead of a ventrally located mouth, five rows of large bony shields on an otherwise scaleless body, and a
heterocercal tail. They are exceptionally long-lived (118 years for Beluga and 70–80 years for White Sturgeon), mature slowly (as late as 30
years old), and spawn infrequently (every 3–5 years). They migrate up rivers to spawn in clean sand and gravel areas; hence, dam building
and siltation of rivers impede their reproduction. A spawning female can be worth thousands of dollars for her caviar alone, and many
sturgeon stocks have been reduced 99% from historical levels. The Shortnose Sturgeon of Atlantic coastal rivers is listed as Vulnerable, and
Fig. 9 Beluga Sturgeon, Husa huso, may be the world’s largest and most valuable freshwater fish. The eggs or caviar from a single, large female
can be worth thousands of dollars.
Lake Sturgeon have been extirpated from a large part of their native range. Three species endemic to the Aral Sea may be extinct due to
extensive drying of that once huge water body, and several species in the former Soviet Union are fished mercilessly.
The North American Paddlefish, Polyodon spathula, occurs in large rivers of North America and also has a heterocercal tail, unconstricted
notochord, largely cartilaginous skeleton, and scaleless body. The most distinctive feature of the paddlefish is its paddle or spoonbill, which is
flat and rounded. Paddlefishes are water column swimmers that feed on zooplankton and fishes. The Paddlefish may help direct food into its
mouth, but abundant electroreceptor cells on the paddle suggest an additional sensorial function. North American Paddlefish may live 30
years and grow to be 2.2 m in length and 83 kg in weight. Late maturation (at 10 years of age), infrequent spawning (every 2–5 years), lack of
clean gravel spawning habitat, and overfishing have all contributed to population reductions and range contraction. The now considered to be
extinct Chinese Paddlefish, Psephurus gladius, was larger, reaching 7 m in length. This paddlefish was recently (2022) considered to be extinct
because of overfishing, habitat destruction, pollution, and dam construction that blocked spawning migrations in the Yangtze River basin
where it used to live. The construction of the Three Gorges Dam in China may have sealed the fate of the Chinese species.
Holosteans
Gars and Bowfins

Highly disputed in the last decades, the latest evidence supports he existence of the Infraclass Holostei, including the gars and the Bowfins.
The lepisosteiform gars are seven species of elongate, predaceous fishes that occur in eastern North America, Cuba, and Central America.
They typically inhabit backwater areas of lakes and rivers, such as oxbows and bayous, and breathe atmospheric oxygen using a highly
vascularized gas bladder. Gars have bony skeletons, but their vertebral centra are unique, being convex anteriorly and concave posteriorly.
In most fishes, the vertebrae are concave on both surfaces. Gars have an abbreviate heterocercal tail (vertebral column extends into the top
lobe of the tail) and hinged, diamond-shaped, interlocking ganoid (heavy and bony) scales. Gars are the only freshwater fish in North
America with poisonous eggs. Alligator Gars can be 3 m long and weigh 140 kg (Fig. 10). In recent years, Alligator Gars have come under
intense commercial fishing, and concern for their well-being is increasing. The Bowfins, Amia calva, and A. ocellicauda, are the only living
members of the genus, family and order. The Bowfins have the abbreviate heterocercal tail and spiral valve intestine of the gars but also has
teleost-like biconcave vertebrae as well as cycloid scales, a relatively light scale type also possessed by many teleosts. The Bowfins’ head is
exceptionally bony and the throat is covered by a distinctive large bone, the gular plate. Bowfin swim slowly forwards or backwards by
passing undulations back and forth along its elongate dorsal fin. Bowfin occur throughout much of eastern North America in backwater,
often swampy areas; they also have a highly vascularized gas bladder which functions as a lung. They are relatively large and robust (to 1 m
and 9 kg) and predatory on anything that moves. Bowfin males guard the young vigorously until they are relatively large (10 cm).
Teleosts
The Infraclass Teleostei (“perfect bone”) contains most living fishes. Teleosts are not only taxonomically diverse but also ecolo-
gically diverse, occupying every aquatic habitat type and niche imaginable. The 34,940 living teleostean species are placed in 4963
genera, 534 families, and 45 orders. This incredible diversity is generally organized into four taxonomic subdivisions that reflect
Fig. 10 An Alligator Gar, Atractosteus spatula, one of the largest freshwater fishes in North America. Photo courtesy of Jean-Francois Healias,
www.anglingthailand.com.
patterns of evolution that date back to the Mesozoic. These four main subdivisions are the osteoglossomorphs (bonytongues),
elopomorphs (eels and tarpon), otocephalans (herrings, minnow, tetras, catfishes, and relatives), containing most freshwater
fishes worldwide, and the euteleosts, containing the majority of modern bony marine fishes.
Osteoglossomorphs
Infraclass Teleostei, Supercohort Osteoglossomorpha, Order Hiodontiformes: one family, two species, mooneyes. Order Osteoglossi-
formes: six families, 251 species, including bonytongues, African knifefishes, and elephantfishes. Osteoglossomorphs derive their name
“bonytongue” from the teeth on their mouth floor that forms part of their bite. These freshwater fishes occur on all major continents
except Europe. The hiodontiforms comprise two species, the Mooneye and Goldeye, both of which occur in major river systems of
northern North America. The osteoglossiforms are much more diverse. The Asian Arowana or Golden Dragonfish, Scleropages formosus,
has been depleted in the wild due to overcollecting and is now protected under Appendix I of the Convention on International Trade in
Endangered Species https://cites.org/eng/app/appendices.php (Fig. 11). The Arapaima of South America is one of the world’s largest
freshwater fishes, reaching a length of 2.5 m. The African mormyrid elephantfishes produce and detect weak electric fields, have large
cerebellums, and have a brain size-body weight ratio comparable to that of humans.
Elopomorphs
Order Elopiformes: two families, nine species, ladyfishes and tarpons; Order Albuliformes: three families, 13 species, including
bonefishes and spiny eels; Order Notacanthiformes: two families, 27 species; Order Anguilliformes: 19 families, 1037 species,
including freshwater, moray, cutthroat, conger eels, swallower eel, and gulper eels.
Elopomorphs all have ribbon-shaped leptocephalus (“pointed head”) larvae. The Atlantic Tarpon is a highly prized gamefish
that reaches a length of 2.5 m and a mass of 150 kg. Albuliform bonefishes are also popular gamefishes that occupy sandy flats in
shallow tropical waters. The 15 families of anguilliform “true” or freshwater eels are distinguished from the approximately 45
other families of “eel-like” fishes that have independently evolved an elongate body. Anguillid eels are catadromous, spawning at
sea but spending most of their lives in fresh water. Muraenid moray eels and their relatives are marine, tropical and warm-
temperate, predatory species. Synaphobranchid cutthroat eels include an endoparasitic species, the Snubnose Parasitic Eel,
Simenchelys parasitica, which has been found in the heart of a mako shark. The saccopharyngiform deepsea gulper and swallower
eels have giant mouths but lack many head bones, scales, and fins found in most other fishes.
Otocephalans
Order Clupeiformes: seven families, 433 species, including anchovies and herrings.
Fig. 11 The Asian Arowana or Golden Dragonfish (Scleropages formosus, Osteoglossidae), an internationally protected species. Desirable color varieties
have sold for as much as US$ 5000. Photo by Marcel Burkhard, Wikimedia Commons, http://en.wikipedia.org/wiki/Image:Arowanacele4.jpgfile.
Clupeiforms are small, schooling, silvery, pelagic marine and occasionally freshwater feeders on zooplankton and phyto-
plankton. Herrings, round herrings, shads, alewives, sprats, sardines, pilchards, and menhadens are extremely important com-
mercial species. Anchovies range in size from a 2 cm Brazilian species to a piscivorous, riverine, 37 cm New Guinea anchovy. The
largest clupeids are the Indo-Pacific chirocentrid wolf herrings, which reach a length of 1 m and have fang-like jaw teeth.
Anadromous shads, Alewife, and herrings occasionally establish landlocked populations in rivers, lakes, and reservoirs.
Order Alepocephaliformes: two families, 142 species, including slickheads and smells. This order was previously classified as a suborder
of the Argentiniformes of the Euteleosteomorpha, but recent molecular evidence is now placing it within the Cohort Otocephala.
Ostariophysans
Order Gonorhynchiformes: four families, 37 species, including milkfish; Order Cypriniformes: 23 families, 4742 species, including
minnows, barbs, algae eaters, suckers, and loaches; Order Gymnotiformes: five families, 266 species, including glass, ghost, and naked-back
knifefishes and three species of Electric Eel; Order Cithariniformes: two families, 116 species; Order Siluriformes: 39 families, 4083 species,
including airbreathing, electric, marine, upside-down, parasitic, callichthyid armored, and suckermouth armored catfishes; Order Char-
aciformes: 22 families, 2201 species, including freshwater hatchetfishes, piranhas, pacus, piaus, tetras, and characins.
Rivers, lakes, and streams worldwide are dominated numerically and ecologically by members of the superorder Ostariophysi.
Ostariophysans include minnows, carps, barbs, suckers, loaches, piranhas, tetras, catfishes, and electric eels. Two distinctive traits
unite this otherwise disparate assemblage: (1) the Weberian apparatus, which is a series of modified anterior vertebrae that link the
gas bladder to the inner ear and aid in hearing, and (2) production and reaction to chemical alarm substances that are released
when a fish is injured and lead to a stereotyped escape response in school members.
The gonorhynchiform Milkfish, Chanos chanos, is an important food fish in the Indo–Pacific region and is often cultured in
brackish fishponds. The Cypriniformes make up the largest order in the superorder with 4742 species of minnows, carps, barbs,
barbels, loaches, suckers, gudgeons, chubs, dace, pikeminnows, Tench, Rudd, bitterlings, bream, southeast Asian “sharks” (Redtail
Black Shark and Bala Shark), Goldfish, Koi (domesticated common carp), danios, and rasboras. Cyprinids are most diverse in
Southeast Asia, followed by Africa, North America, and Europe, but are missing from South America and Australia. The world’s
smallest freshwater fish is an Indonesian paedocypridid, Paedocypris progenetica, that matures at 7.8 mm. The largest minnow in
North America is the endangered, piscivorous Colorado Pikeminnow, Ptychocheilus lucius.
Gyrinocheilid algae eaters scrape algae off rocks in swift, flowing waters. The catostomid suckers (i.e., buffaloes, Quillback,
carpsuckers, Blue Sucker, redhorses, jumprocks, and the extinct Harelip Sucker) include 83 species of North American fishes, with
one species in eastern China and another in Siberia. Loaches (Cobitidae) are 227 species of predominantly Eurasian stream fishes,
including popular aquarium fishes such as the kuhli, Clown, and skunk loaches, the weatherfishes, and the Golden Dojo.
Weatherfishes (Misgurnus) become restless when barometric pressure decreases preceding a storm.
The gymnotiform South American knifefishes are unusual ostariophysans that produce and detect weak electric impulses. They
have elongate, compressed bodies; an extremely long anal fin; and electrogenic tissue usually derived from modified muscle cells.
Their electrical output is constant at high frequencies, whereas the osteoglossomorph mormyrids produce a pulsed, low-frequency
output. Both groups detect objects that disrupt their electric fields. The three species of 2 m long electric eels (Electrophorus),
produce a weak field for electrolocation and strong pulses for stunning prey or deterring predators.
The cithariniforms have long been classified within the characiforms, but recent molecular evidence suggest they are sister-
group to siluriforms plus characiforms. There are two African families in this order, the Citharinidae, with eight species and the
Distichodonidae, with 108 species.
Fig. 12 Piranhas (subfamily Serrasalminae) belong to the incredibly diverse order of characiform fishes that dominate South American fresh
waters. Although certainly capable of dismembering prey, most reported attacks on humans involved drowning victims.
Siluriforms (catfishes) are surprisingly diverse, with 40 families and more than 4083 species. Catfishes usually have barbels (whiskers)
and sometimes toxic spiny fins, and they are mostly freshwater, nocturnal, and benthic (notable exceptions being the marine Ariidae with
157 species and Plotosidae with 42 species). Large species include the European Wels, Silurus glanis (5 m, 330 kg), the Asian Mekong Giant
Catfish, Pangasianodon gigas (3 m, 300 kg), and the 3 m long whiskered pimelodid of South America, the Piraíba, Brachyplatystoma
filamentosum. The largest catfishes in North America are the Flathead and Blue catfishes at about 1.5 m and 50–68 kg. Some small catfishes
are notable, such as the parasitic catfishes (Trichomycteridae) of South America, which normally parasitize the gills of fishes but species of
Candiru (Vandellia), are known to swim up the urethra of bathers and lodge there, necessitating surgical removal.
The characiforms are a speciose group of primarily tropical ostariophysans characterized (usually) by a rayless adipose fin and
mouths armed with replacement dentition (e.g., piranhas) (Fig. 12). Body size ranges from very small (13 mm) tetras to large
(1.5 m long) African tigerfishes. Numerous aquarium fishes are included (headstanders, freshwater hatchetfishes, blind characins,
pencilfishes, tetras, and silver dollars), as well as important food fishes (Prochilodus, Colossoma, and Brycon). Most characids (1252
species) are South American, a few live in Central America, and one species, the Mexican Tetra Astyanax mexicanus, extends
naturally into southwestern Texas.
Euteleosteans
Approximately 2/3 of the teleosts are placed in the Euteleostei, the true teleosts, a vast assemblage of more derived fishes. This
subdivision contains 397 families, 3311 genera, and 21,240 species.
Order Lepidogalaxiiformes: one family, one species only, Lepidogalaxias salamandroides. The southern hemisphere salamander
fish inhabits seasonal ponds of southwestern Australia, burying in drying mud and reemerging with the next rains.
Order Argentiniformes: four families, 99 species, including barreleyes, pencilsmelts, and slickheads; Order Salmoniformes: one
family, 247 species, including smelts, trouts, and salmons; Order Esociformes: two families, 13 species, pikes and mudminnows.
Protacanthopterygians, as these three orders are known, are a mixed agglomeration of marine, freshwater, and diadromous fishes.
Argentiniforms are mostly deepwater marine species. Salmoniforms are important commercially, ecologically, and esthetically.
Whitefishes and ciscoes are relatively large-scaled, zooplanktivorous salmonids of high-latitude North American and Eurasian lakes.
Several North American species have been decimated due to introduced predators, competitors, and parasitic lampreys. Graylings are
riverine fishes with a flowing dorsal fin. The subfamily Salmoninae contains seven Eurasian and North American genera. The Siberian
Taimen, Hucho taimen, is the world’s largest salmonid at 2 m and 70 kg. North American Salmoninae include the chars (Lake, Brook,
and Bull trout, Arctic Char, and Dolly Varden). Arctic Char live farther north than any other freshwater fish. The remaining salmonines
are the Atlantic basin salmon and trout (e.g., Atlantic Salmon and European Brown Trout), and the 11 species of Pacific basin trouts and
salmons, two of which are endemic to Japan. Pacific trouts and salmons include Golden, Cutthroat, and Gila trouts and the specta-
cularly anadromous Coho, Chinook, Chum, Pink, and Sockeye salmons, some of which undergo oceanic migrations of thousands of
kilometers before returning to their birth river to spawn and die (Fig. 13). The Rainbow Trout is a sixth species of Pacific salmon and is
called a Steelhead Salmon if it spends part of its life at sea. The actual number of genetically distinct races of Pacific salmons is unknown
because many stocks are reproductively isolated in small river systems. Evidence suggests that as many as 1000 stocks exist, more than
100 of which have gone extinct and an additional 300 þ of which are imperiled.
Esociform pikes, pickerels, and mudminnows are Northern Hemisphere predators, including the 1.4 m Muskellunge; the Northern Pike
has the largest geographical distribution of any Northern Hemisphere fish, occurring across the northern portions of North America,
Europe, and Asia. Mudminnows can survive winters in high-latitude lakes by breathing from air bubbles trapped under the ice.
Fig. 13 A spawned-out, dead Sockeye Salmon, Oncorhynchus nerka, in an Alaskan stream. Spawning runs of Sockeye and other Pacific salmon
species play a pivotal role in the ecosystems of the Pacific Northwest. Bears, eagles, etc. feed on spawning fish, and dead fish provide nutrients
for food webs, including promoting growth of the large trees that shade the streams essential for survival of young salmon.
Order Osmeriformes: Four families, 41 species. Stomiiformes: Four families, 454 species. Osmeriforms are small, silvery,
elongate, water column dwelling fishes. Osmerids include commercially important species such as capelins, eulachons, Asian Ayu,
and smelts. Stomiiforms include bristlemouths, marine hatchetfishes, and barbeled dragonfishes. They are deepsea fishes, often
with long teeth and large mouths. Gonostomatid bristlemouths may be the most abundant and widely distributed vertebrates on
Earth. Idiacanthine black dragonfishes have a larva with eyes at the ends of elongate stalks.
Order Galaxiiformes: One family, 66 species. Galaxiiforms have suffered numerous extirpations and extinctions as a result of
the stocking of nonnative trouts. Some species of high latitudes of the southern hemisphere live in freshwater as adults, but spawn
in the ocean. Circumpolar currents carry the eggs and newly hatched larvae, that eventually enter rivers around the globe, the same
genetically connected populations living in South America, south Africa, Australia and New Zealand.
Order Ateleopodiformes: One family, 11 species, jellynosefishes. The jellynosefishes are elongate deepwater marine fishes that
swim just above the bottom. Their large, pointed head and pointed tail are traits they share with other deepsea fishes such as
chimaeras, spiny eels, halosaurs, eucla cods, rattails, and grenadiers.
Order Aulopiformes: 16 families, 296 species, including telescopefishes, tripodfishes, lizardfishes, and lancetfishes. Aulopiforms
are also deepsea forms, including the bizarre giganturid telescopefishes with large tubular eyes, a huge mouth, flexible teeth, and an
expandable stomach. Deepsea tripodfishes have long pectoral, pelvic, and caudal rays that they use for resting on soft sediments of
the deep ocean floor. Shallow representatives are the synodontid lizardfishes, which are common benthic predators on coral reefs
worldwide. Alepisaurid lancetfishes are large (to 2 m) mesopelagic predators with a sail-like dorsal fin of unknown function.
Order Myctophiformes: Two families, 260 species, including lanternfishes. Myctophiforms include the abundant, commercially
important lanternfishes, which are identified based on species-specific photophore (light organ) patterns. Lanternfishes occur at
middle depths from the Arctic to the Antarctic. They are important in the diets of many fishes as well as of marine mammals.
Acanthomorphs
Order Lampriformes: six families, 29 species, including opahs, tube-eye, ribbonfishes, and Oarfishes.
Lampriforms are generally open-water, oceanic fishes. Opahs are relatively large (1.8 m, 70 kg), oval-shaped, pelagic predators
on squids and other fishes. The elongate Oarfish, Regalecus glesne, may attain 12-m length and is the longest living teleost. It has a
bluish-silvery body, scarlet head crest, and deep red fins. It is thought to be responsible for many “sea serpent” sightings (Fig. 14).
Paracanthopterygians, as the following five orders are known, include: Order Percopsiformes: three families, 12 species,
troutperches, pirate perch, and cavefishes. Order Polymixiiformes: one family, 11 species, beardfishes. Order Zeiformes: six
families, 33 species, zeiforms include the dories and boarfishes, as well commercial species such as the European John Dory, Zeus
faber. Order Stylephoriformes: one family, one species, the Tube-eyes, Stylephorus chordatus. Order Gadiformes: 17 families, 617
species, including rattails, hakes, and cods.
Paracanthopterygians are primarily benthic, marine, nocturnally active fishes; many live in the deep sea or in caves. Percopsiforms
are small (o20 cm), freshwater fishes, most of which live in eastern North America. The anus of the swamp-dwelling aphrododerid
Pirate Perch is located in the throat region of adults for functional reasons. Amblyopsid cavefishes are often blind and scaleless forms
highly adapted for cave life. Polymixiiforms as represented by 11 species of beardfishes in one family of deep-sea, bottom dwellers
from most oceans around the World. Polymixiiforms were classified as sister-group to the Acantopterygii by Betancur et al. (2017).
Fig. 14 The Oarfish, Regalecus glesne, is probably the longest bony fish in the world and is responsible for many reports of sea monsters. This
individual died at a shoreline in Mexico. Photo courtesy of Michael Kanzler.
The gadiforms include the cods, haddocks, hakes, pollocks, and whitings, which are some of the world’s most important
commercial fishes. True cods (Gadinae) have three dorsal fins and two anal fins. Many species have chin barbels. The
Burbot, Lota lota, of high-latitude, Northern Hemisphere lakes is the only freshwater species in the group. The commercially
important Atlantic Cod, Gadus morhua, is the largest species (1.8 m, 90 kg), but fish more than 10 kg are rare due to drastic
overfishing. One of the world’s largest food fisheries is for North Pacific Walleye Pollock, Gadus chalcogrammus. The 30 cm
long tube-eye (Stylephorus chordates – only stylephoriform) can increase the volume of its mouth 40-fold during feeding – a
record among vertebrates.
Acanthopterygians
Most bony fishes belong to a single Division, the Acanthopterygii or spine-finned fishes, which contains about 19,050 species in
327 families. Two orders and one large taxonomic grouping, the Percomorpha, are recognized, with the vast majority in the latter.
Order Trachichthyifomes: five families, 66 species. The trachichthyiforms include the commercially important Orange Roughy,
Hoplostethus atlanticus, of high-latitude, southern ocean regions. Orange Roughies are being overexploited because they are slow
growing and long-lived, taking more than 20 years to mature and reaching ages of more than 100 years.
Order Beryciformes: nine families, 211 species, including flashlight fishes, roughies, and squirrelfishes. Beryciforms are shallow-
to moderate-depth, often red, almost always nocturnal fishes, including the reef-dwelling squirrelfishes.
Percomorphs
The Percomorpha constitute by far the largest taxonomic group of fishes, with around 18,780 species in 313 families. They are far
too many to deal with in any detail. What follows is a very brief overview of the orders and more interesting families. Percomorphs
are classified in 13 Orders by Dornburg and Near (2021), the classification followed here, and nine Series with 36 Orders by
Betancur et al. (2017) in a more traditional order-level classification.
Ophidiiformes, with four families and 558 species, include pearlfishes, cuskeels, and viviparous brotula. They often live in
holes or even inside other animals. Carapid pearlfishes live inside the body cavities of starfishes, sea cucumbers, clams, and sea
squirts; some feed on the internal organs of their hosts. Ophidiid and bythitid cuskeels and brotulas include blind cave species in
freshwater systems of Caribbean and Galapagos Islands as well as coral reef species that hide deep within crevices. The neo-
bythitine Cusk Eel, Abyssobrotula galatheae, holds the depth record for a fish at 8370 m in the Puerto Rico Trench.
Batrachoidiformes, with one family and 84 species, include the midshipmen, which have hundreds of photophores, an
unusual trait for a shallow dweller. Many batrachoidids produce sounds by vibrating their gas bladders. Venomous toadfishes have
dorsal and opercular spines that can inject a powerful toxin. Three South American toadfishes are restricted to fresh water.
Gobiiformes (¼ Gobiaria) includes 11 families and 2695 species. The apogonid cardinalfishes, a diverse (385 species), small
(o10 cm), and nocturnal group of coral reef fishes. Cardinalfishes mouthbrood their eggs, an unusual trait among reef species.
Gobiiforms are usually small, benthic, often abundant fishes. The eleotrid sleepers are small to medium (to 60 cm) estuarine and stream
fishes in tropical and subtropical areas, often on islands. The gobiid gobies are incredibly diverse (about 1970 species). Most gobies have
a suction disk formed by fused pelvic fins. The family includes the amphibious mudskippers of the Indian and Pacific oceans. Gobies
range in size from tiny pygmy gobies (8–10 mm adults) to a comparative giant, the western Atlantic Violet Goby, Gobioides broussonnetii,
which is a purplish eel-like fish 50 cm long. Round Gobies have recently been introduced into North American Great Lakes from
southern Europe and are spreading rapidly; other introduced goby species are now extremely common in San Francisco Bay.
The Order Scombriformes (¼ Pelagiaria) and the Order Syngnathiformes ( ¼ Syngnatharia) are sister-group to each other.
Scombriformes, with 16 families and 284 species, contain some of the largest and most spectacular marine fishes. Twenty species
of barracudas inhabit tropical and subtropical oceans almost worldwide. The gempylid snake mackerels (26 species) are pelagic and
deepwater predators, including the cosmopolitan Oilfish, Ruvettus pretiosus, a large (1.8 m, 45 kg) predator of moderate depths. An
active fishery for Oilfish in the Comoro Islands captures endangered Coelacanths as by-catch. The scombrid mackerels and tunas are
quintessential open-sea predators, with streamlined bodies and a physiology geared to a high-speed lifestyle. They range from
relatively small, 50-cm mackerels to giant Bluefin Tuna, Thunnus thynnus (4 m, 500 kg). Most are schooling fishes of tremendous
commercial importance; a single, 340 kg Bluefin Tuna sold for US$ 400,000 in Tokyo in 2011. The temperate and warm-temperate
xiphiid Swordfish, Xiphias gladius, and the more tropical istiophorid sailfishes, spearfishes, and marlins have an elongate upper jaw
bone that forms the bill. It is used as a spear, a cutlass, or a billy. Swordfish grow to 530 kg, whereas Blue and Black marlins grow to
900 kg. Swordfish have been heavily overfished, particularly in the Atlantic, although revised management plans have reduced fishing
pressure and the species has shown signs of a comeback. The voracious, schooling pomatomid Bluefish, Pomatomus saltatrix, occurs in
most temperate and semitropical oceans except the eastern Pacific. The icosteid consists of the peculiar North Pacific Ragfish, Icosteus
aenigmaticus, which has a largely uncalcified, cartilaginous skeleton and is a preferred food of sperm whales.
The Syngnathiformes, with 10 families and 679 species, includes the bizarre pegasid seamoths and syngnathid pipefishes, seadragons,
and seahorses. Syngnathid pipefishes and seahorses are the only vertebrates in which the female impregnates the male by laying eggs in his
brood pouch, which he then fertilizes and raises until hatching. Seahorses are heavily overfished for medicinal uses and the aquarium
trade. Another bottom-oriented family is the tropical reef-dwelling mullid goatfishes, which have movable, muscularized chin barbels.
Blenniiformes (¼ Ovalentaria) includes 49 families and 5866 species. The largest family is Cichlidae, with around than 1800 species.
Cichlids are chiefly tropical, Central and South American, up to the Rio Grande in south Texas (approx. 600 species) and African fishes
(approx. 1200 species), with a few species in India and Sri Lanka. Central and South American species include freshwater angelfishes,
discus, oscars, dwarf cichlids, and peacock basses. Most cichlids occur in Africa, where they are particularly speciose in the African Great
Lakes and are threatened by introduced predators such as the Nile Perch, Lates niloticus (Fig. 17). African tilapias and other cichlids have
been deliberately or accidentally introduced in many parts of the world, as a byproduct of aquaculture activities. Other important
blenniiforms are pomacentrid damselfishes (425 species), which are small, colorful, usually herbivorous, territorial reef dwellers. Some
damselfishes are zooplanktivores and some maintain a symbiotic mutualistic relation with anemones (anemonefishes or clownfishes –
Nemo!). Several occupy temperate regions (e.g., the Garibaldi of California, see Fig. 20). The embiotocid surfperches are 23 species of
deep-bodied, temperate (mostly eastern Pacific) fishes associated with kelp beds and rocky reefs. They are livebearers, feeding on
zooplankton or small invertebrates. The Superfamily Atherinoidea are shallow-water, marine or freshwater fishes that live near the
surface, and include the melanotaeniid rainbowfishes of Australia and New Guinea, in which males have brighter colors and longer fins
than females, traits that make them popular aquarium species. Silversides are widespread, freshwater and marine schooling fishes and
include the grunions of southern and Baja California that ride waves up beaches on dark nights to spawn in wet sand biweekly during
the summer. Superfamily Belonoidea are predominantly silvery, marine fishes active at and sometimes above the surface of the water,
including needlefishes, flyingfishes, and halfbeaks. The lower lobe of the tail in flyingfishes is relatively long and is used to scull rapidly
during takeoff. Superfamily Cyprinodontoidea also comprise a large group, with approximately 1430 species of topminnows, killifishes,
livebearers, and pupfishes. Many cyprinodontiforms, although basically freshwater fishes, tolerate considerable salinity and hence occur
in streams on isolated oceanic islands. The rivulids of South America and Africa live only one year, laying eggs that survive in the dried
bottoms of pools and that hatch with the next season’s rains. Kryptolebias marmoratus of south Florida and the West Indies is the only
truly hermaphroditic fish, fertilizing its own eggs. The poeciliid livebearers include the mollies, platys, guppies, and swordtails of the
aquarium trade. Two species are all female and only reproduce parthenogenetically. Many cyprinodontid pupfishes are tolerant of
extreme water conditions and consequently can live in saltmarsh and desert conditions. However, they cannot tolerate total desiccation,
which has endangered many desert species that have to compete with humans for water. The Devil’s Hole Pupfish, Cyprinodon diabolis,
has the smallest known range of any fish species – one shallow shelf in a single spring in Nevada, near the Death Valley. Other pupfish
group inhabit Lake Titicaca, which at an elevation of 3812 m in the Andes Mountains is the highest natural lake with fishes. These are
severely threatened by the introduction of trout in the Titicaca and nearby lakes.
The mugilids include the mullets, and are a family of near-shore, marine and freshwater fishes of considerable economic
importance. Many mullets feed on organic silt and minute plants, an unusual food type among fishes. Gobiesocids are small,
marine and freshwater clingfishes, that live in shallow-water and sometimes are amphibious fishes, often found in high-energy
wave zones. Their pelvic fins are modified into a strong sucking disc. Blennioidei are small marine fishes that usually associate with
shore structure. Chaenopsid pikeblennies and tubeblennies often live inside corals and worm tubes. Bleniid combtooth blennies
are diverse (405 species) small fishes in tropical and subtropical waters; they scrape algae with their comb-like teeth.
The Order Synbranchiformes (¼ Anabantaria) and the Order Carangiformes ( ¼ Carangiria) are sister-group to each other.
Synbranchiformes includes the 12 families and 409 species. Suborder Synbranchoidei are primarily freshwater, eel-like, often
airbreathing fishes, including synbranchid swamp eels and mastacembelidae tiretrack eels. Swamp eels have recently been
introduced into the southeastern United States and are of major concern as potential invading predators in river systems. Fishes of
suborder Anabantoidei have an auxiliary breathing structure in the gill chamber for aerial respiration, the labyrinth. Anabantid
climbing gouramies are African and Asian freshwater fishes that can move across wet ground and reportedly up wet tree trunks. The
Kissing Gourami is the sole member of the family Helostomatidae. The osphronemid gouramies, fighting fishes (Bettas), and
paradise fishes have elongate pelvic fin rays that serve as feelers. Bettas (Siamese Fighting Fish, Betta splendens) have been bred to
battle like fighting cocks, placing them among the few fishes that have been cultured for purposes other than food, appearance, or
research. The channid snakeheads are highly predatory – and sometimes highly invasive – African and Asian fishes. The nandid
leaffishes and badid chameleonfishes are small Asian families with important ornamental interest.
Carangiformes (¼ Carangiaria) includes 31 families and 1100 species, and is composed of the former orders Istiophoriformes,
Carangiformes, and Pleuronectiformes, of previous classifications. In echeneid remoras or sharksuckers, the modified first dorsal fin forms
a suction disk that is used to cling to various hosts. The coryphaenid dolphinfishes or mahi mahis are two species of open-water, pelagic
predators that often associate with floating structures. They are also one of the few marine pelagic fishes that is successfully aquacultured.
Carangid scads, jacks, pompanos, and amberjacks are a large family (151 species) of tropical near-shore predators. Centropomids and
latids are large predatory fishes, the former includes the snooks of tropical America, and the latter the Barramundi of Australia, Lates
calcarifer, and the Nile Perch of Africa, Lates niloticus. Nile Perch are an introduced predator in Lake Victoria, in which they are thought to
have extinguished perhaps hundreds of endemic cichlids. Toxotid archerfishes are Indo–Pacific, brackish-water fishes well-known for their
ability to shoot droplets of water that knock insects out of overhanging vegetation. Flatfishes (Suborder Pleuronectoidei) are distinctive,
compressed, benthic fishes that lay in one body side and have both eyes on the same side of the head. Many flatfishes are important
commercially (e.g., dab, flounders, halibuts, plaice, sole, tonguefishes, turbots, and whiffs). Paralichthyids include the Summer Flounder
and California Halibut, the latter reaching 1.5 m and 30 kg. The pleuronectid righteye flounders include the Atlantic and Pacific halibuts.
Pacific Halibuts may live 40 years and attain lengths of 3 m and masses of 200 kg. The fishery for Pacific Halibut in the North Pacific is a
well-regulated, sustainable enterprise. The achirid, citharid, cynoglossid, and soleid flatfish families have several freshwater representatives.
The Perciformes includes 52 families and 3200 species. The perciforms comprise many marine and freshwater fishes of littoral zones.
Perciforms reach their greatest diversity on coral reefs, but they are also highly diverse in rivers, streams, and lakes. At least 242 species make
up the family Percidae, most of which occur in North America. Percids include the Yellow and Eurasian perch, Walleye and Sauger
(¼ pikeperches), and about 190 species of small, stream-dwelling, spectacularly colored, and often imperiled darters. Fishes of the family
Zoarcidae are all eel-like, bottom-living, marine, cool- to cold-water species. They range in size from the small, intertidal pricklebacks and
gunnels to the live-bearing eelpouts, some of which live 3000 m below the surface. The large (to 2.5 m long) anarhichadid wolffishes and
wolf eels of shallow North Pacific and Atlantic waters are anatomically and ecologically similar to moray eels. Notothenid icefishes and
relatives are mostly Antarctic, mostly benthic fishes that live under the ice and have antifreeze compounds in their blood. The channichthyid
crocodile icefishes lack red blood cells, hemoglobin, and myoglobin, and hence have colorless blood and flesh. Trachinids weeverfishes are
marine, generally benthic fishes that tend to bury themselves in sand. They occur in the eastern Atlantic and Mediterranean and have
venomous opercular and dorsal spines. Gasterosteids are small marine, estuarine, and freshwater fishes with dermal armor plating.
Sticklebacks are well-studied fishes that frequently form distinct, isolated populations characterized by unusual spines, plates, and behavior.
Scorpaenids are marine fishes, with spiny heads and sometimes venomous fin spines (i.e., stonefishes, scorpionfishes, and lion-
fishes) (Fig. 15). The Lionfish (Pterois volitans) is native to the Indo-Pacific, but have been introduced and is now established along the
southeast coast of the US, the Caribbean, and the northeastern coast of South America up to southeastern Brazil. The lionfish is a
voracious predator and threatens native fish and the environment. The sebastine rockfishes are a diverse, commercially important, long-
lived, and overfished group of the temperate North Pacific, some of which live to be over 200 years old. Hexagrammid greenlings are
littoral zone (inshore and shallow water) and kelp-associated fishes endemic to the North Pacific, including the highly edible Lingcod,
Ophiodon elongatus. The cottids contains many freshwater species, including the sculpins of North American streams and the highly
divergent (pelagic and livebearing) oilfishes of Lake Baikal in northern Asia. The scorpaenichthyid Cabezon of the Pacific coast of North
America is unusual in having toxic eggs, whereas the cyclopterid lumpfish of the North Atlantic produces valuable caviar and has
Fig. 15 Lionfishes, Pterois volitans, belong to the order of scorpionfishes, many of which have toxic spines. Lionfishes are native to Indo–Pacific
regions. They have few natural predators, which has contributed to their success as an invasive introduced predatory species on tropical western
Atlantic reefs. Photo by Jens Petersen, from Wikimedia Commons.
Fig. 16 Large groupers, such as this Giant Grouper, Epinephelus lanceolatus from the Indo–Pacific, grow as large as 2.7 m long and can weigh over 400 kg.
consequently been overfished. The Serranidae range in size from small hamlets (10–15 cm) to giant groupers (3 m long, 400 kg)
(Fig. 16). Sea basses also include commercially important hinds, coneys, Gag, and Scamp. Many serranids are hermaphroditic, usually
maturing first as female and then later becoming male, although some hamlets are both male and female simultaneously.
The Centrarchiformes includes 17 families and 300 species. Centrarchiforms are mostly marine, but the freshwater centrarchids
contains the sunfishes, crappies, rockbasses, and black basses of North America. Largemouth Bass, Micropterus salmoides, have been
introduced extensively as a sport fish in many parts of the world. Elassomatid pygmy sunfishes are a family of miniature
(20–45 mm), colorful freshwater swamp dwellers of the southeastern United States.
The Labriformes includes seven families and 760 species of mostly tropical, reef-dwelling wrasses and parrotfishes. Labrid wrasses
range in size from the 5 cm long cleaner wrasses, Slippery Dicks, and Blueheads to the 2.3 m long Maori or Humphead Wrasse of the
Indo–Pacific, which is hunted unmercifully for the live fish restaurant trade. Cool temperate species include the California Sheephead
and Señorita, western Atlantic Tautog and Cunner, and the eastern Atlantic Cuckoo Wrasse. Many wrasses change sex from female to
a more colorful male. The scarin parrotfishes (100 species) are almost exclusively coral reef dwellers, best known for their fused
parrot-like teeth that are used for biting off algal and coral pieces, which are then crushed in the massive pharyngeal (throat) jaws.
Uranoscopid stargazers that emit strong pulses of electricity (up to 50 V) from highly modified eye muscles. Ammodytid sand lances
are small, elongate, and abundant zooplankton feeders that spend their nights buried.
The Acropomatiformes is composed of 21 small families and 308 species, of marine or brackish water, and some are capable of
producing bioluminescence.
The Acanthuriformes is composed of 55 families and 2360 species. It includes the luvarid and zanclid families with one species each,
and the usually herbivorous acanthurid surgeonfishes, unicornfishes, and tangs (85 species), which have a knife blade on the caudal
peduncle. Caproidae is represented by a single species of boarfish, occurring between 40 and 700 m deep in the northeastern Atlantic
Ocean and Mediterranean sea. Chaetodontids include the colorful reef butterflyfishes (134 species), related to the leiognathid pony-
fishes (53 species). Pomacanthid angelfishes (90 species) are also very conspicuous in coral reefs. These families are most diverse in the
Indo-Pacific region. Butterflyfishes feed on coral polyps, small invertebrates, tube worms, or zooplankton. Ephippids and drepaneids are
small families with 18 species of spadefishes and sicklefishes. Gerreids include 54 species of silver biddies and mojarras that live in
tropical to temperate seas, usually on sandy or muddy bottoms. Lobotids are composed by 15 species of tripletails, which occur in
tropical and subtropical oceans. Some species of tiger perch or freshwater tripletails occur in southeast Asia and New Guinea.
Suborder Lophoidei includes goosefishes, frogfishes, handfishes, batfishes, and deepsea anglerfishes. This is a diverse and often
bizarre-looking clade of marine fishes that include benthic, shallow-water forms as well as highly modified, open-water, deepsea forms.
Many use a modified first dorsal-fin spine as a lure for catching smaller fish (see Fig. 21). The meter-long western North Atlantic
Goosefish, Lophius americanus, has a huge mouth with long, recurved teeth that it uses to catch fishes and even diving seabirds.
Antennariid frogfishes also rest on the bottom or walk over it with their pectoral and pelvic fins. The ogcocephalid batfishes walk on their
pectorals, but they can also swim by means of jet propulsion by shooting water out their round, backward-facing opercular openings. The
ceratiid anglerfishes include strange appearing bathypelagic predators, many of which have very small males that fuse to and become
parasitic on the larger females. The endemic Australian handfishes include a Tasmanian species, the Spotted Handfish, Brachionichthys
hirsutus, that was once common but is now Critically Endangered due possibly to egg predation by an introduced starfish. Haemulid gruts
Fig. 17 Cichlid fishes in the Great Lakes of Africa are highly diverse ecologically and anatomically, with more than 1000 species in the major
lakes. These cichlids at the Georgia Aquarium in Atlanta are representative of what one might see diving in Lake Malawi, Africa.
and sweetlips (136 species), and lutjanid snappers, with 138 species are generally carnivorous marine fishes. Snappers are usually near
bottom dwellers (i.e., Gray Snapper, Red Snapper, and Mangrove Snapper), but some are water column zooplanktivores (Vermilion
Snapper). Haemulid grunts are moderate-sized coral reef fishes and are most diverse in the New World tropics. They are seen most often
in their daytime resting schools around coral heads; at night they disperse to feed in surrounding reef and grass areas. Moronid temperate
basses include the Striped and White bass of North America. Cepolid bandfishes have the dorsal and anal fins continuous with the
lanceolate caudal fin and fossorial habits, while priacanthid bigeyes are typically bright red. Sciaenid croakers (297 species) have chin
barbels and a muscularized gas bladder used for sound production. Sciaenids are a widespread family that is particularly diverse in the
southeastern North America, but representatives occur widely in tropical marine and freshwater habitats. Species include Red Drum (spot
tail bass), Black Drum, croakers, weakfish, kingfishes, White Sea Bass, corbinas, and the endangered Mexican Totoaba, Totoaba macdonaldi,
one of the many unquestionably imperiled marine fishes. The range of the Freshwater Drum, Aplodinotus grunniens, includes much of
eastern North America into Central America. Four genera and 22 species of freshwater drums form an important component of South
American fish fauna. Scientists have demonstrated that South American sciaenids invaded and differentiated in freshwaters two times.
Sparids include 282 species of seabreams and porgies, nemipterid has 95 species of threadfin breams and whiptail breams, and lethrinid
comprises 43 species of emperor breams and pigface breams. Most are bottom-dwelling carnivores inhabiting shallow temperate and
tropical marine waters, but few species are planktivores. Finally, Suborder Tetraodontoidei is currently classified in 10 families (435
species) including puffers, porcupine fishes, boxfishes, triggerfishes, and the Sunfish. Some of the most derived bony fishes are in this
suborder, an almost entirely marine clade of medium-sized fishes with thick, leathery skin and with scales often modified into spines or
bony plates. In balistid triggerfishes and monacanthid filefishes, the long, rigid first dorsal-fin spine is locked erect by an interaction with
the shorter, second spine. Ostraciidae boxfishes are encased in a triangular or rectangular bony box, with just the fins and caudal peduncle
emerging. Puffers and ocean sunfishes lack true teeth. Instead, the jaw bone has a cutting edge that looks like separated teeth or is fused
into a parrot-like beak. Diodontid porcupine fishes inflate their body by filling the stomach with water, a process that also helps erect and
interlock their body spines. Tetraodontid puffers concentrate a powerful and potentially fatal toxin, tetrodotoxin, in their viscera, which
adds to the allure of eating puffers in licensed Fugu restaurants in Japan. The Ocean Sunfish, Mola mola, is one of the world’s heaviest
fishes at 1000–2000 kg, producing as many as 300 million eggs. All this biomass is supported on a diet of jellyfishes (Fig. 18).
Geographic Diversity
Overview
Fishes occur just about everywhere water occurs in its liquid state, is available through most of the year, and remains below 401C. A major
zoogeographic distinction can be made between marine and freshwater fishes, with substantial overlap happening where intermediate
salinities occur. Many fishes are restricted to pure fresh water (little or no salinity), many are restricted to normal oceanic salinity (about 35
parts per 1000 salt in water), some occur in both habitats at different times of their lives or of the year, and some occur and are even restricted
Fig. 18 A Giant Ocean Sunfish, Mola mola, being cleaned by an adult Emperor Angelfish, Pomacanthus imperator, on a reef in Bali, Indonesia.
Photo courtesy of Ali Watters, www.mytb.org/Ali.
to areas of intermediate salinity, such as estuaries. In terms of numbers, about 50% of all fishes are marine and 49% live in fresh water, with
the remaining 1% moving regularly between the two salinity designations (Fig. 19 and Box 2). Among the 18,050 freshwater species, 80% are
primary or obligatory freshwater fishes and are intolerant of even moderate salinities. The remaining 20% can tolerate some salinity and
hence inhabit upper estuarine areas or can cross through near-shore ocean regions to move from one river basin to another. Among the
18,200 marine fishes, the vast majority (69%) live in shallow, warm areas such as coral reefs. The remaining marine species are divided fairly
evenly among shallow cold, open-ocean, and deep-bottom areas (about 10% each). About 2% of marine fishes live in near-surface, open-sea
(pelagic) habitats. The approximately 185 diadromous species that live in different salinity regions at different times of their lives are divided
among three groups. Anadromous fishes (54%) live most of their lives in the ocean but then migrate to fresh water to spawn; this group
includes lampreys, sturgeons, shad, and salmons. Catadromous fishes (25%) spend most of their lives in fresh water and migrate to the sea to
spawn; included here are freshwater eels, mullets, and temperate basses. Amphidromous fishes (21%) move between fresh and salt water, but
migration to the spawning habitat occurs long before the fishes actually spawn; examples include gobies, sleepers, and galaxiids.
Freshwater Diversity
The world’s freshwater habitats occur in six major zoogeographic regions or realms that correspond approximately to continental
distributions, with important exceptions. Each region has a fairly distinct fish fauna (again with some exceptions and shared
elements) (Berra, 2007).
The Nearctic Region

The Nearctic region consists of subtropical, tropical, temperate, and arctic North America. The region stretches from the Mexican
Plateau to northern Canada and Alaska. The Nearctic contains 14 families of primary freshwater fishes, with about 1200 species.
The most diverse families are minnows, suckers, North American catfishes, perches (and darters), and sunfishes. Other important
families include the lampreys, gars, salmons (many of which are anadromous), and whitefishes; sculpins, which are freshwater
Deep benthic 2100

Meso/bathypelagic 1800
Epipelagic 380
Shallow cold 1820
Shallow warm 12,100 18,200
Habitat type
Total marine
Amphidromous 40
Catadromous 45
Anadromous 100
Total diadromous 185
Secondary 3600
Primary 14,450
Total freshwater 18,050
0 5000 10,000 15,000 20,000

Number of species
marine diadromous freshwater

Fig. 19 The global biodiversity of marine, diadromous, and freshwater fishes. Approximate numbers of species are given for each habitat type
and its subdivisions. Black: marine; Gray: diadromous; White: freshwater.
Fig. 20 The Garibaldi, Hypsypops rubicundus, is a temperate member of the otherwise largely tropical family of pomacentrid damselfishes. It is
common in kelp beds of southern California and is the state marine fish.
species in a primarily marine family (¼ marine derivatives); pickerels and mudminnows; killifishes; and livebearers. The Nearctic is
further subdivided into three subregions: the Arctic–Atlantic (with six provinces), the Pacific (with seven provinces), and the
Mexican Transition subdivision. Eleven major river systems drain the region; major lakes are abundant, the largest being the five
Laurentian Great Lakes (Ontario, Erie, Huron, Michigan, and Superior).
Fig. 21 A 15 cm long female Wolftrap Anglerfish, Lasiognathus amphirhamphus. Anglerfishes are among the more diverse groups in the
bathypelagic region. Many anglerfishes possess a lure at the tip of their modified first dorsal spine used to bring prey within striking distance of a
large, expandable mouth. Photo courtesy of Pietsch TW.
Box 2 Why are there so many freshwater fishes?
The high global diversity of freshwater fishes (Fig. 19) is at first surprising. Fresh waters make up only about 0.009% of the earth’s
water, which means that almost half of all fish species live in a small fraction of the world’s water. This 7,500-fold discrepancy in
biodiversity per unit volume is probably best explained by the relative productivity and isolation of freshwater bodies. Most freshwater
habitats are relatively shallow and receive ample sunlight as well as nutrients running off from adjacent land. Hence freshwater
habitats are relatively productive and capable of sustaining abundant life. Most of the ocean, in contrast, is deep, dark, and nutrient-
poor. Given that 81% of marine diversity occurs in shallow regions, a relationship between water depth and diversity in fresh waters is
not surprising. Adding to the influence of available food is the comparative isolation of most freshwater habitats. River basins and lakes
are often created and affected by climatic and geologic forces (e.g., drought, floods, landslides, earthquakes, uplifts) that separate
them from other systems. Every basin then can be relatively isolated from other basins, which means that genetically distinct
populations can evolve into new species and that little genetic mixing occurs between basins. Small streams are separated from each
other by larger rivers which are barriers to the movement of small fishes, and large rivers are separated by oceans or mountain chains.
As a result, freshwater habitats are perfect habitats for the speciation process. Oceans in contrast are largely continuous habitats that
are connected by currents, and ocean fishes typically produce larvae that float for several weeks or months on these currents. Hence
genetic exchange is common and opportunities for speciation are not as great. Oceanic basins have relatively distinct faunas, but
connectedness within basins decreases the kind of genetic isolation needed for speciation of the sort seen in freshwaters.
The Neotropical Region

The Neotropical region contains South America and Middle America. It is the most speciose region of the world in terms of freshwater
fishes, with 63 families, more than 6400 described species, and current estimated of around 9000 species (Reis et al., 2016), with many
species remaining to be discovered and described. Particularly diverse groups include the colorful characiforms (2201 species of tetras,
piranhas, characins, and freshwater hatchet fishes), 15 families and 2427 species of catfishes, five families and 266 species of gymnotiform
electric knifefishes, and around 600 species of cichlids. Several secondary freshwater and marine derivative groups are included: freshwater
stingrays, herrings, silversides, needlefishes, killifishes, croakers, and others. The Neotropical region has been further divided into eight
subdivisions with fairly distinctive faunas. Six major river systems drain the region, including the largest river on Earth; major lakes include
Lake Titicaca, the world’s highest fish-containing lake at 3812 m above sea level, and colossal permanently or seasonally flooded areas.
The Palearctic Region

The Palearctic region encompasses Eurasia, including Europe, northern Africa, and Asia north of the Oriental region. Twenty-seven
families and about 700 species of temperate freshwater fishes occur in the region, dominated by minnows and loaches but also
perches, pickerels, sturgeons, salmons, sculpins (including the Lake Baikal endemics), and 12 species of catfishes in four families.
Endemism increases to the south, as is also the case in the Nearctic region (Kottelat and Freyhof, 2007). The Palearctic and Nearctic
regions share numerous families and genera (sturgeons, paddlefishes, minnows, smelts, salmons, pikes, mudminnows, and
perches) but only a few species occur in both (i.e., Northern Pike, Longnose Sucker, Burbot, Threespine Stickleback, and Fourhorn
Sculpin). The region is sometimes subdivided into six subregions based on faunal groupings. Ten major river systems drain the
region; major lakes include the Black and Caspian Seas and Lake Baikal, the world’s oldest and deepest lake.
The African or Ethiopian Region

The African or Ethiopian region is second to the Neotropics in freshwater fish diversity, with 48 families and more than 3600
species of primary and secondary freshwater fishes. The African region includes all the African continent south of the Sahara Desert,
plus the large island of Madagascar with its endemic fauna. One third of the fishes are in the superorder Ostariophysi, including
400 minnows, over 200 characiforms, 116 cithariniforms, and 400 catfishes in six families. Other diverse groups include killifishes
and topminnows, elephantfishes and other osteoglossiforms, and cichlids. As many as 1200 cichlid fishes may occur in the three
African Great Lakes of Lake Victoria, Lake Tanganyika, and Lake Malawi, with more cichlids in smaller surrounding lakes and rivers
(see Fig. 17). Four lungfishes and all 14 polypteriform bichirs occur in Africa. Ten to 12 zoogeographic provinces are recognized,
with six major river drainages and numerous lakes including the African Great Lakes of Victoria, Malawi, and Tanganyika.
The Oriental Region

The Oriental region includes eastern Iran, India and Sri Lanka, China south of the Yangtze River, Southeast Asia, and the large island
regions of Taiwan, the Philippines, and the East Indies/Indo–Malayan Archipelago. The Oriental region contains 48 families of primary
and secondary freshwater fishes. Most diverse are the minnows, loaches, and 12 families of catfishes; clariid walking catfishes and bagrid
catfishes are particularly diverse. Other important groups include algae eaters, river loaches, snakeheads, spiny eels, labyrinth fishes and
gouramis, a few cichlids, and archerfishes. The Oriental region shares many families with the Palearctic to the north and the Ethiopian to
the west but shares few families with the Australian region to the southeast. The Oriental is often subdivided into two major subregions:
Peninsular India with more than 1050 species and Southeast Asia with more than 1650 species. Each subregion has two major river
drainage systems; large lakes are uncommon. Southeast Asia is sometimes divided further into five zoogeographic regions.
The Australian Region

The Australian region (New Guinea, Australia, New Zealand, and Oceania) is relatively depauperate in true freshwater fishes, and in fact all
but three of the freshwater fishes in the region are members of families obviously derived from marine groups. The northwestern border of
the region, and the practical limit of primary freshwater fishes, is dramatically delineated by an ocean boundary that lies southeast of Java,
Borneo, and Sulawesi and is known as Weber’s Line. Nineteen families and about 225 species occur primarily in fresh water in the region,
but only the Australian lungfish and two species of bonytongue saratogas are true freshwater fishes (another 33 families and 160 species of
marine fishes frequently enter fresh waters in Australia). Other important families, many with species endemic to specific regions, include
lampreys, river eels, herrings, catfishes, southern smelts, and graylings, Salamander fish and other galaxiids, silversides, rainbow fishes,
Barramundi, grunters, glassfishes, temperate perches or basses, sleepers and gobies, and Torrentfish (in New Zealand). One major river
system occurs on New Guinea (the Fly) and two on Australia (Darling and Murray); permanent, large lakes are rare.
Marine Diversity
Delimiting zoogeographic regions in the world’s oceans is complicated by depth, currents, and geographic locales; different faunal breaks
occur depending on near-shore, pelagic, or deepsea environments. The greatest fish diversity and the greatest geographic differentiation occur
in near-shore, continental shelf (to about 100 m depth) regions. These regions are separated by continents, by large expanses of open ocean,
and by currents that differ in temperature from that of the region in question. Temperature zones divide the seas into tropical, temperate,
boreal, and polar regions. In addition, different faunal groupings apply to pelagic fishes and to fishes of the deep sea (Briggs, 1995).
The Indo-West Pacific Region

The Indo-West Pacific region includes shallow tropical seas that extend from South Africa and the Red Sea eastwards through the Indo-
Malayan area and Australia to Hawaii and Easter Island; it also includes Micronesia, Melanesia, and Polynesia. The Indo-West Pacific is by
far the most species-rich marine area, containing 4000–5000 tropical fish species. It is also diverse in sea snakes and many invertebrate taxa
such as reef-building and soft corals, mollusks, tube worms, and echinoderms, although these numbers describe life in the upper 50–75 m
of the reef that are accessible to scuba divers. As new exploration technologies allow deeper exploration, hundreds more fish species have
been discovered. Regardless, the Indo-West Pacific is considered the center of diversity for many of the common coral reef fish families that
occur in other tropical regions. Only a few families are endemic to the Indo-West Pacific (e.g., sillaginid whitings and rabbitfishes).
Common families include moray eels, squirrelfishes, sea basses, grunts, snappers, cardinalfishes, butterflyfishes, angelfishes, damselfishes,
wrasses, parrotfishes, surgeonfishes, gobies, triggerfishes, and boxfishes. Barriers to movement of Indo-West Pacific fishes are cool waters to
the north and south and a vast expanse of open, deep water to the east known as the Eastern Pacific Barrier. Eight different provinces have
been recognized in the Indo-West Pacific, separating it into a huge Indo-Polynesian region and seven smaller areas.
The Eastern Pacific Region

The Eastern Pacific region, with approximately 800 fish species, runs from southern Baja California to Ecuador, its northern and
southern limits defined by the coldwater California and Peru currents. Despite its location in the Pacific Ocean, the Eastern Pacific
is faunistically more similar to the tropical Atlantic, containing many species that are almost indistinguishable from Atlantic forms.
The two oceans mixed before the Panamanian Isthmus formed and the two areas still share some species, despite three million
years of physical separation. Most families are less diverse here than in the Western Atlantic, with the exceptions of sea catfishes,
croakers, and herrings. Dactyloscopid sand stargazers occur here and in the Atlantic, but not in the Indo-West Pacific, a distribution
pattern common to other families like Chaenopsidae and many genera. Sixty two Indo-West Pacific species have managed to cross
the Eastern Pacific Barrier. Three provinces – Mexican, Panamanian, and Galapagos – are recognized.
The Western Atlantic Region

The Western Atlantic region is the second most diverse oceanic area, containing 1200 fish species. It includes Bermuda (which,
although at 321 N, sits in the tropical 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 and toadfishes. Strong currents of warm water separate the Western Atlantic
fauna from colder waters along much of its boundaries. It is subdivided into Caribbean, Brazilian, and West Indian provinces.
The Eastern Atlantic Region

The Eastern Atlantic region is a relatively small area along the west coast of Africa from Senegal to Angola and extending out to
oceanic islands such as Ascension and St. Helena. Tropical marine fishes here are limited by cool-water currents impinging from both
the north and the south as well as by substantial freshwater runoff and sediments from several major west African rivers, all factors
that discourage coral reef growth. The region contains “only” 500 shore fishes; most coral reef families occur but are represented by
only a few species. Porgies are particularly diverse. No subdivisions are recognized, with the possible exception of the warm-
temperate Mediterranean. The Mediterranean Sea contains 760 species, with many species in the same families as those in the Eastern
Atlantic. The Mediterranean has the dubious distinction of being the most heavily invaded tropical marine area in the world, with
more than 50 alien fish species having moved in from the Red Sea (an Indo-West Pacific subregion) via the human-made Suez Canal.
The Arctic Region

The Arctic region encompasses high-latitude (above 601N) waters of both the Pacific and Atlantic Oceans. It reaches from Nunivak
Island, Alaska, northward and across the polar region to Newfoundland and Norway in the northern Atlantic. Of the two polar
areas, the Arctic is more diverse. Successful groups include skates, herrings, salmons, smelts, cods, eelpouts, greenlings, sculpins,
poachers, snailfishes, pricklebacks, wolffishes, gunnels, and righteye flounders. Diversity within many of these groups is greater in
the Pacific than in the Atlantic portions of the region. Over 415 species occur here. Distribution of many of these families appears
to be limited by temperature, with warmer waters and currents to the south determining species’ boundaries.
The Antarctic Region

The Antarctic region (above 601 S) has its own distinctive fauna that is restricted to Antarctic waters and the surrounding Southern Ocean,
including the cold waters of Australia, New Zealand, and nearby oceanic islands. Forty-nine families and over 280 species occur here, 13
families and 180 species of which are identified as Antarctic continent species. A particularly successful group is the notothenid icefishes
and relatives, which account for 55% of Antarctic species. Families include bovichthyids, cod and crocodile icefishes, plunderfishes, and
dragonfishes. Non-notothenids include skates, snailfishes, eelpouts, lantern fishes, eel cods, deepsea cods, and southern flounders.
Temperate Regions
To the north of the Indo-West Pacific lie cooler temperate waters with their own characteristic fish faunas. This area can be divided
into four fairly distinct regions according to location and temperature: Japanese warm-temperate and Californian warm-temperate
regions and Eastern and Western boreal regions. The warm-temperate areas (from about Hong Kong to Tokyo in the west and from
lower Baja California to central California in the east) contain a fauna that fluctuates seasonally, as tropical species move north in
the summer and boreal species move south in the winter. Notable families to the west include lizardfishes, flyingfishes, mullets,
jacks, sea basses, and croakers and to the east include endemic silversides, sea basses, croakers, damselfishes, wrasses, and flatfishes
(Fig. 20). The more northerly boreal regions (approximately north from central California in the east and Korea in the west)
contain similar families but different species. Important families include migratory salmonids, sculpins, rockfishes, snailfishes,
greenlings, gunnels, pricklebacks, and righteyed flounders. In the southern Pacific, coldwater currents create at least three temperate
faunal regions, with centers of distribution around Australia, New Zealand, and South America.
In the Atlantic, three northern temperate areas occur: the western and eastern Atlantic boreal regions and the Atlantic warm-
temperate or Carolinian region. The boreal regions (Newfoundland to Cape Hatteras in the west and British Isles to northern Europe
and Scandinavia in the east) share a fauna of salmonids, cods, sticklebacks, poachers, sculpins, wolffishes, and righteyed flounders,
with occasional strays from more southerly waters during warm months. The Carolinian extends from Cape Hatteras south to Florida
and also to the Gulf of Mexico, with southern Florida housing tropical species. Common groups are clupeids, sea robins, pipefishes,
silversides, needlefishes, killifishes, croakers, lefteye flounders, and puffers. Temperate faunas also occur in the southern Atlantic, but
their areas and diversity are less than those of the southern temperate faunas of the Pacific Ocean. Two recognized regions are the
eastern South American and southern African warm-temperate regions. The former region’s fauna includes sea catfishes, croakers,
herrings, gobies, scorpionfishes, and sea basses; the latter area has many colder water members of Indo-West Pacific families.
Pelagic Regions
More than 400 species of pelagic fishes occur in ocean surface waters to a depth of 200 m. This habitat type can be divided into 10
different regions based on faunal differences, with more joining of Southern Hemisphere areas because of the relative lack of large
land masses. These regions are Arctic and Antarctic polar, North Pacific cold temperate, North Pacific warm-temperate, tropical
Indo-Pacific, North Atlantic boreal, North Atlantic warm-temperate, Atlantic tropical, southern warm-temperate, and southern
cold temperate. Many of the world’s most important fisheries species occur in pelagic regions, including numerous sharks, sardines
and herrings, salmons, codfishes, pollocks, hakes, haddocks, sauries, mackerels, and tunas. In addition, about 100 species of
mostly pelagic fishes have a worldwide distribution. This group includes pelagic sharks (White, Whale, Tiger, and perhaps
Megamouth), Swordfish, and ocean sunfishes.
The Deep Sea

Waters deeper than 200 m are as much a zoogeographic as a habitat entity. The deepsea is generally divided into three major
regions based on depth: the open-water mesopelagic (200–1000 m) and bathypelagic (1000–4000 m) regions and the bottom-
associated bathyal (200–4000 m) region. Bathyal fishes are further divided into benthopelagic fishes that hover just above the
bottom and benthic fishes that rest in contact with the bottom. Deepsea fishes are most common in these regions between 401N
and 401 S latitude, approximately between San Francisco and Melbourne in the Pacific and between New York City and the Cape
of Good Hope in the Atlantic. Abyssal and hadal (trench) regions deeper than 4000 m are relatively depauperate.
Each region has a characteristic and relatively diverse bony fish fauna consisting of fishes from many different taxonomic
groups. The mesopelagic region worldwide contains about 850 species in nine different orders. Despite their lack of relatedness,
deepsea fishes share many anatomical and physiological features, suggesting independent, convergent evolution of adaptations to
deepwater existence. Mesopelagic fishes typically have photophores (light organs) on their silvery bodies; have relatively large,
often-tubular eyes; undergo daily migrations to surface waters to feed at night; and have large mouths and long teeth. Common
names of mesopelagic fishes reflect these traits: barreleyes, bristlemouths, dragonfishes, sabertoothfishes, lanternfishes, tubeeyes,
and swallowers.
The bathypelagic region is the largest habitat space on the Earth, accounting for 88% of oceanic volume. The nine orders and
over 250 species in the cold, dark bathypelagic region share some traits with mesopelagic fishes but possess them in the extreme.
Photophores are concentrated on lures used to attract prey; eyes are often small; mouths are extremely large and teeth very long;
stomachs are expandable; bodies are black; and body musculature, bones, and scales are greatly reduced. These traits reflect greater
habitat space and increasingly rare feeding opportunities with increasing depth, which select for an increasing need to conserve
energy and to be able to take advantage of feeding opportunities. Again, bathypelagic fishes have names indicative of their
adaptations: sawtooth eels, gulper eels, swallower eels, dragonfishes, anglerfishes, seadevils, and fangtooths (Fig. 21. Anglerfish).
Bathyal fishes include over 1000 bony fish species in nine orders plus chimaeras and squaloid sharks. Different families inhabit
bottom compared to open-water regions. Bathyal fishes include greeneyes, tripodfishes, hakes, grenadiers, cuskeels, batfishes,
snailfishes, and eelpouts. Although diversity decreases below 1000 m, grenadiers and rattails live between 1000 and 4000 m,
tripodfish have been found to 6000 m and snailfishes to 7000 m, and some cuskeels have been found as deep as 8000 m.
Although some differences in species composition occur in different ocean basins or in association with different water masses,
deepsea species are relatively cosmopolitan, occurring in several different oceans. One trend is for fishes to occur deeper at lower
latitudes, such that species that are bathypelagic near the equator may be mesopelagic at middle latitudes and even epipelagic at
the poles.
Conclusion
Fishes represent an astonishing font of biodiversity information and a vast resource of food, recreation, and enjoyment. This group
is as diverse as all other vertebrates together, but human activities in the last century have driven many species to the verge of
extinction, some being gone forever. Besides studying their diversity, evolution, ecology, and natural history, humanity must make
every effort to find alternatives to our way of life in order to save many fishes from extinction.
References
Barton, M. (2006) Bond’s biology of fishes, third ed. Stamford, CT: Thomson Brooks/Cole.
Berra, T. M. (2007) Freshwater fish distribution. Chicago: University of Chicago Press.
Betancur, R., R., Wiley, E. O. and Arratia, G. et al. (2017) Phylogenetic classification of bony fishes. BMC Evolutionary Biology 17, 1–40.
Briggs, J. C. (1995) Global biogeography. Amsterdam: Elsevier.
Carrier, J., Musick, J., Heithaus, M. (eds.) 2004. The biology of sharks and their relatives. Boca Raton, FL: CRC Press.
Carrier, J., Musick, J., Heithaus, M. (eds.) 2010. Sharks and their relatives II. Biodiversity, adaptive physiology and conservation. Boca Raton, FL: CRC Press.
Dornburg, A. and Near, T. J. (2021) The emerging phylogenetic perspective on the evolution of Actinopterygian fishes. Annual Review of Ecology, Evolution, and Systematics 52, 427–52.
Helfman, G. S. (2007) Fish conservation: A guide to understanding and restoring global aquatic biodiversity and fishery resources. Washington, DC: Island Press.
Helfman, G. S. and Collette, B. B. (2011) Fishes: The animal answer guide. Baltimore, MD: Johns Hopkins University Press.
Helfman, G. S., Collette, B. B., Facey, D. E. and Bowen, B. (2009) The diversity of fishes: Biology, evolution, and ecology, second ed. Oxford: Wiley-Blackwell.
Kottelat, M. and Freyhof, J. (2007) Handbook of european freshwater fishes. Cornol: Publications Kottelat.
Moyle, P. B. and Cech, J. J. (2004) Fishes: An introduction to ichthyology, fifth ed. Upper Saddle River, NJ: Prentice-Hall.
Nelson, J. S. (2006) Fishes of the world, fourth ed. Hoboken, NJ: John Wiley & Sons.
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Encyclopedia of Biodiversity 3rd edition, Volume 2 doi:10.1016/B978-0-12-822562-2.00092-X 341

Pelagic Regions 367

Box 1 What is a ﬁsh and what are ﬁshes?

Soft-rayed dorsal fin (second dorsal)

Operculum or gill cover

Lateral line Peduncle

Ventral surface Finlets

Barbel Anal spines Anal fin

Skates and Rays

Bichirs and Reedﬁsh

Sturgeons and Paddleﬁshes

Gars and Bowﬁns

The Nearctic Region

Deep benthic 2100

0 5000 10,000 15,000 20,000

marine diadromous freshwater

Box 2 Why are there so many freshwater ﬁshes?

The Neotropical Region

The Palearctic Region

The African or Ethiopian Region

The Oriental Region

The Australian Region

The Indo-West Paciﬁc Region

The Eastern Paciﬁc Region

The Western Atlantic Region

The Eastern Atlantic Region

The Arctic Region

The Antarctic Region

The Deep Sea

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