Fishes

C H A P T E R
Fishes
24
• PHYLUM CHORDATA
• CLASS MYXINI
• CLASS PETROMYZONTIDA
• CLASS CHONDRICHTHYES
• CLASS ACTINOPTERYGII
• CLASS SARCOPTERYGII
Myxini
Hammerhead shark, Sphyrna, near the Galápagos Islands. Petromyzontida
Chondrichthyes
Actinopterygii
Sarcopterygii
Chordata
What Is a Fish?
In common (and especially older) usage, the term “fish” denotes a convenience, not as a taxonomic unit. Fishes are not a monophy-
mixed assortment of water-dwelling animals. We speak of jellyfish, letic group, because the ancestor of land vertebrates (tetrapods) is
cuttlefish, starfish, crayfish, and shellfish, knowing full well that when found within one group of fishes (the sarcopterygians). Thus, fishes
we use the word “fish” in such combinations, we are not referring to can be defined in an evolutionary sense as all vertebrates that are
a true fish. In earlier times, even biologists did not make such a dis- not tetrapods. Because fishes live in habitats that are less accessible
tinction. Sixteenth-century natural historians classified seals, whales, to humans than terrestrial habitats, people have rarely appreciated
amphibians, crocodiles, even hippopotamuses, as well as a host of the remarkable diversity of these vertebrates. Nevertheless, whether
aquatic invertebrates, as fishes. Later biologists were more discrimi- appreciated by humans or not, the world’s fishes have enjoyed an
nating, eliminating first the invertebrates and then the amphibians, effusive proliferation that has produced an estimated 28,000 living
reptiles, and mammals from the narrowing concept of a fish. Today species—more than all other species of vertebrates combined—with
we recognize a fish as an aquatic vertebrate with gills, appendages, adaptations that have fitted them to almost every conceivable aquatic
if present, in the form of fins, and usually a skin with scales of der- environment. No other animal group threatens their domination of
mal origin. Even this modern concept of the term “fish” is used for the world’s seas, lakes, and streams.
www.mhhe.com/hickmanipz14e CHAPTER 24 Fishes 515
T
he life of a fish is bound to its body form. Their mastery of are known. By the Devonian period, the Age of Fishes, several
stream, lake, and ocean is revealed in the many ways that distinct groups of jawed fishes were common. One of these,
fishes have harmonized their life design to the physical the placoderms (p. 512), became extinct in the following Car-
properties of their aquatic surroundings. Suspended in a medium boniferous period, leaving no descendants. A second group,
that is 800 times more dense than air, a trout or pike can remain cartilaginous fishes of class Chondrichthyes (sharks, rays,
motionless, varying its neutral buoyancy by adding or removing and chimaeras), lost the heavy dermal armor of early jawed
air from its swim bladder. It may dart forward or at angles, using fishes and adopted cartilage rather than bone for the skeleton.
its fins as brakes and tilting rudders. With excellent organs for salt Most chondrichthyans are active predators with sharklike or ray-
and water exchange, fishes can steady and finely tune their body like body forms that changed only slightly over the ages. As a
fluid composition in their freshwater or seawater environment. group, sharks and their kin flourished during the Devonian and
Their gills are the most effective respiratory devices in the animal Carboniferous periods of the Paleozoic era but declined dan-
kingdom for extracting oxygen from a medium that contains less gerously close to extinction at the end of the Paleozoic. They
than 1/20 as much oxygen as air. Fishes have excellent olfac- recovered in the early Mesozoic and radiated to form the mod-
tory and visual senses and a unique lateral-line system, which est but thoroughly successful assemblage of modern sharks and
has an exquisite sensitivity to water currents and vibrations. Thus rays (Figure 24.1).
in mastering the physical problems of their element, early fishes The other two groups of gnathostome fishes, acanthodi-
evolved a basic body plan and set of physiological strategies that ans (p. 512) and bony fishes, were abundant and diverse in
both shaped and constrained the evolution of their descendants. the Devonian period. Acanthodians resembled bony fishes but
were distinguished by having heavy spines on all fins except the
caudal fin. They became extinct in the lower Permian period.
ANCESTRY AND RELATIONSHIPS Although phylogenetic affinities of the acanthodians are much
OF MAJOR GROUPS OF FISHES debated, many authors consider them the sister group of bony
fishes. Bony fishes (Osteichthyes, Figure 24.2) are the most
Fishes are a vast array of distantly related gill-breathing aquatic
abundant fishes today. We can recognize two distinct groups
vertebrates with fins. They are the most ancient and most
of bony fishes. Of these two, by far the most diverse are ray-
diverse of the clade Vertebrata, constituting five of the nine
finned fishes (class Actinopterygii), which radiated to form
living vertebrate classes and about half of the approximately
most modern bony fishes. The other group, the lobe-finned
55,000 vertebrate species.
fishes (class Sarcopterygii), contains few fish species today but
Fishes are of ancient ancestry, having descended from an
includes the sister group of the tetrapods. Lobe-finned fishes are
unknown free-swimming protochordate ancestor about 550 mil-
represented today by lungfishes and coelacanths—meager
lion years ago (hypotheses of chordate and vertebrate origins are
remnants of important lineages that flourished in the Devonian
discussed in Chapter 23). The earliest vertebrates were an assem-
period (see Figure 24.1). A classification of the major fish taxa is
blage of jawless agnathan fishes, including the ostracoderms
on page 540.
(see Figure 23.14, p. 510). One group of ostracoderms gave rise
to the jawed gnathostomes.
LIVING JAWLESS FISHES
The use of fishes as the plural form of fish may sound odd to most
Living jawless fishes include approximately 108 species divided
people accustomed to using fish in both the singular and the plural.
between two classes: Myxini (hagfishes) with about 70 species
Fish refers to one or more individuals of the same species; fishes
and Petromyzontida (lampreys) with 38 species (Figures 24.3
refers to more than one species.
and 24.4). Members of both groups lack jaws, internal ossifica-
tion, scales, and paired fins, and both groups share porelike gill
The jawless agnathans include along with the extinct ostra- openings and an eel-like body form.
coderms the living hagfishes and lampreys, fishes adapted as Based on this morphological similarity, these two groups
scavengers or parasites. Although hagfishes have no vertebrae formerly were united under the name “Cyclostomata,” a group-
and lampreys have only rudimentary vertebrae, they neverthe- ing shown to be paraphyletic when morphological characters
less are included with the subphylum Vertebrata because they were analyzed with cladistic methods (p. 209). Lampreys have
have a cranium and many other vertebrate homologies. Although many characters, including vertebrae, extrinsic eye muscles,
hagfishes and lampreys superficially look much alike, they are in at least two semicircular canals, and cerebellum, which they
fact so different from each other that they have been assigned to uniquely share with gnathostomes. Interestingly, recent analysis
separate taxonomic classes by zoologists. of molecular characters shows hagfishes and lampreys forming
All remaining fishes have paired appendages and jaws and a monophyletic unit. This grouping, inconsistent with the mor-
are included, along with tetrapods (land vertebrates) in the phological data, is not supported by most zoologists, and this
monophyletic group of gnathostomes. They appear in the fossil “cyclostome” hypothesis needs further testing. Thus we take the
record in the late Silurian period with fully formed jaws, and view that hagfishes form the sister group of a clade that includes
no forms intermediate between agnathans and gnathostomes lampreys and gnathostomes (see Figure 24.2).
516 PART THREE Diversity of Animal Life
Lungfishes
Coelacanths
)
hes Sturgeons
d fis
Modern amphibians
e
finn
be-
Gars
ns
s (lo
ea
Amniotes
t
os
gian
dr
on
tery
Ch
s ts)
ian eos
cop
yg el
ter s (t
Sar
p n
eo gia
n ery
rly eopt
Ea rn n
de
Mo Modern
Early bony fishes
s
amphibians
istian
ians
pt eryg
Neo
Rhipid
s
es) dian
fish ntho
nne
d Aca
-fi hs
s (ray br anc
ian mo
ryg Ela
s
pte
ino Sharks, skates,
Act
Gnatho- rays
stomata
Placoderms Chondrichthyes Holocephalans

Chimaeras
Ostracoderms
Lampreys
Common Agnatha
chordate
ancestor Vertebrata (Craniata)
Hagfishes
Cambrian Ordovician Silurian Devonian Carboniferous Permian

MESOZOIC CENOZOIC
PALEOZOIC
570 245 66 0
Geologic time (My ago)
Figure 24.1
Graphic representation of the family tree of fishes, showing evolution of major groups through geological time. Numerous lineages of extinct
fishes are not shown. Widened areas in the lines of descent indicate periods of adaptive radiation and relative number of species in each group.
The lobe-finned fishes (sarcopterygians), for example, flourished in the Devonian period, but declined and are today represented by only four
surviving genera (lungfishes and coelacanths). Homologies shared by sarcopterygians and tetrapods suggest that they form a clade. Sharks and rays
radiated during the Carboniferous period, declined in the Permian, then radiated again in the Mesozoic era. Johnny-come-latelies in fish evolution
are the spectacularly diverse modern fishes, or teleosts, which make up most living fishes.
hic70049_ch24_514-542.indd 516 7/25/07 1:23:54 PM

Craniata = Vertebrata
Gnathostomata
Teleostomi
Agnatha Chondrichthyes Osteichthyes
Elasmobranchii Actinopterygii Sarcopterygii

Myxini Petromyzontida (sharks, skates, Holocephali (ray-finned (lobe-finned Tetrapods
(hagfishes) (lampreys) rays) (chimaeras) fishes) fishes)
Placoderms †
Acanthodii† Limbs used for

Loss of scales, terrestrial locomotion
Disclike "Ostracoderms"† teeth modified as
buccal funnel, grinding plates Unique supportive ele-
long larval stage ments in skeleton of
as ammocoete girdle and limbs
Lung or swim bladder derived
Placoid scales, from gut, endochondral bone
claspers
Gills not attached to interbranchial
septum (as they are in sharks), bony
operculum
Part of second visceral arch modified as

Naked skin with supporting element for jaws, teeth with dentine
slime glands,
accessory hearts Jaws from mandibular arch, 3 pairs semicircular canals, internal
pelvic and pectoral girdles
Mineralized tissue in dermis
2 or more pairs semicircular canals, vertebrae, extrinsic eye muscles
Distinct head, tripartite brain, specialized sense organs, neural crest tissue,
†Extinct
groups 1 or more pairs semicircular canals, cranium, well-developed pharyngeal skeleton
Figure 24.2
Cladogram of the fishes, showing the probable relationships of major monophyletic fish taxa. Several alternative relationships have been proposed.
Extinct groups are designated by a dagger (†). Some shared derived characters are shown to the right of branch points. The groups Agnatha and
Osteichthyes, although paraphyletic structural grades considered undesirable in cladistic classification, are conveniently recognized in systematics
because they share broad structural and functional patterns of organization.
Class Myxini: Hagfishes in its tail, then passes the knot forward along its body until it is
pressed securely against the side of its prey (see Figure 24.3D).
Hagfishes are an entirely marine group that feeds on annelids,
molluscs, crustaceans, and dead or dying fishes. Thus they
are not parasitic like lampreys but are scavengers and preda- While the strange features of hagfishes fascinate, hagfishes have not
tors. There are about 70 species of hagfishes, of which the best endeared themselves to commercial fishermen. In earlier days of com-
known in North America are the Atlantic hagfish, Myxine gluti- mercial fishing mainly by gill nets and set lines, hagfish often bit into
nosa (Gr. myxa, slime) (see Figure 24.3), and the Pacific hagfish, the bodies of captured fish and devoured the contents, leaving behind
Eptatretus stoutii (N. L. ept, Gr. hepta, seven, tretos, perfo- a useless sack of skin and bones. But as large and efficient trawls came
rated). Although almost completely blind, hagfishes are quickly into use, hagfishes ceased to be an important pest. Recently the com-
attracted to food, especially dead or dying fishes, by their keenly mercial fishing industry “turned the tables” and began targeting hag-
developed senses of smell and touch. A hagfish enters a dead fishes as a source of leather for golf bags and boots. Fishing pressure
or dying animal through an orifice or by digging into the body. has been so intense that some species have greatly declined.
Using two toothed, keratinized plates on its tongue that fold
together in a pincerlike action, the hagfish rasps bits of flesh Hagfishes are renowned for their ability to generate enor-
from its prey. For extra leverage, the hagfish often ties a knot mous quantities of slime. If disturbed or roughly handled, a
hic70049_ch24_514-542.indd 517 7/25/07 1:23:55 PM

Mouth
External gill
opening
Pores of
Mouth surrounded
slime glands Keratinized
by barbels
teeth on
tongue
Caudal fin
A B
Nostril Olfactory Brain Spinal Notochord Pharynx

sac cord
Barbels
Mouth
Internal
openings
to gill sacs
C Teeth on tongue Tongue D
Figure 24.3
Atlantic hagfish, Myxine glutinosa (class Myxini). A, External anatomy; B, Ventral view of head, showing keratinized teeth used to grasp food
during feeding; C, Sagittal section of head region (note retracted position of rasping tongue and internal openings into a row of gill sacs);
D, Hagfish knotting, showing how it obtains leverage to tear flesh from prey.
hagfish exudes a milky fluid from special glands positioned Unlike any other vertebrate, the body fluids of hagfishes
along its body. On contact with seawater, the fluid forms are in osmotic equilibrium with seawater, as are most marine
a slime so slippery that the animal is almost impossible to invertebrates. Hagfishes have several other anatomical and
grasp. physiological peculiarities, including a low-pressure circulatory
system served by three accessory hearts in addition to the main
heart positioned behind the gills.
The reproductive biology of hagfishes remains largely a
mystery, despite a still unclaimed prize offered more than 100
years ago by the Copenhagen Academy of Science for informa-
tion on the animal’s breeding habits. It is known that females,
which in some species outnumber males 100 to one, produce
small numbers of surprisingly large, yolky eggs 2 to 7 cm in
diameter depending on the species. There is no larval stage.
Class Petromyzontida: Lampreys

All lampreys of the Northern Hemisphere belong to family Pet-
romyzontidae (Gr. petros, stone, myzon, sucking). The group
name refers to a lamprey’s habit of grasping a stone with its mouth
to hold its position in current. The destructive marine lamprey, Pet-
romyzon marinus, occurs on both sides of the Atlantic Ocean (in
America and Europe) and may attain a length of 1 m (Figure 24.4).
Figure 24.4 Lampetra (L. lambo, to lick or lap up) also has a wide distribution
Sea lamprey, Petromyzon marinus, feeding on body fluids of a dying fish. in North America and Eurasia and ranges from 15 to 60 cm long.
hic70049_ch24_514-542.indd 518 7/25/07 1:23:59 PM

There are 20 species of lampreys in North America. About half of

these belong to the nonparasitic brook type; the others are para-
Characteristics of Class Myxini sitic. The genus Ichthyomyzon (Gr. ichthyos, fish, myzon, suck-
1. Body slender, eel-like, rounded, with naked skin ing), which includes three parasitic and three nonparasitic species,
containing slime glands is restricted to eastern North America. On the west coast of North
2. No paired appendages, no dorsal fin (the caudal fin America the chief marine form is Lampetra tridentata, commonly
extends anteriorly along the dorsal surface)
sold as P. marinus by biological supply companies.
3. Fibrous and cartilaginous skeleton; notochord persistent
All lampreys ascend freshwater streams to breed. Marine
4. Biting mouth with two rows of eversible teeth, but no jaws
5. Heart with sinus venosus, atrium, and ventricle; accessory forms are anadromous (Gr. anadromos, running upward); that
hearts, aortic arches in gill region is, they leave the sea where they spend their adult lives to swim
6. Five to 16 pairs of gills with a variable number of gill up streams to spawn. In North America all lampreys spawn in
openings winter or spring. Males begin building a nest and are joined later
7. Pronephric and segmented mesonephric kidneys; by females. Using their oral discs to lift stones and pebbles and
marine, body fluids isosmotic with seawater vigorous body vibrations to sweep away light debris, they form
8. Digestive system without stomach; no spiral valve or cilia an oval depression (Figure 24.5). At spawning, with the female
in intestinal tract attached to a rock to maintain her position over the nest, the
9. Dorsal nerve cord with differentiated brain; no cerebellum; male attaches to the dorsal side of her head. As eggs are shed
10 pairs of cranial nerves; dorsal and ventral nerve roots
into the nest, they are fertilized by the male. The sticky eggs
united
adhere to pebbles in the nest and quickly become covered with
10. Sense organs of taste, smell, and hearing; eyes degenerate;
one pair semicircular canals sand. Adults die soon after spawning.
11. Sexes separate (ovaries and testes in same individual but Eggs hatch in about 2 weeks, releasing small larvae called
only one is functional); external fertilization; large yolky ammocoetes, which are so unlike their parents that early biolo-
eggs, no larval stage gists thought they were a separate species. The larva bears a
remarkable resemblance to amphioxus and possesses the basic
Migration Parasitic stage in lakes

towards
lakes Reproduction
in streams
Filter-feeding
ammocoete
larvae
Metamorphosis
Figure 24.5
Life cycle of the “landlocked” form of the sea lamprey Petromyzon marinus.
hic70049_ch24_514-542.indd 519 7/25/07 1:24:03 PM

Tongue
Characteristics of Class Petromyzontida Pharynx
1. Body slender, eel-like, rounded with naked skin Buccal funnel with
keratinized teeth
2. One or two dorsal fins, no paired appendages
3. Fibrous and cartilaginous skeleton; notochord persistent
4. Suckerlike oral disc and tongue with well-developed
keratinized teeth
Lingual
5. Heart with sinus venosus, atrium, and ventricle; aortic arches cartilage
in gill region
6. Seven pairs of gills each with external gill opening
Attachment to fish with
7. Opisthonephric kidney (p. 673); anadromous and fresh-
keratinized teeth and suction
water; body fluids osmotically and ionically regulated
8. Dorsal nerve cord with differentiated brain, small
cerebellum present; 10 pairs cranial nerves; dorsal and
ventral nerve roots separated
9. Digestive system without distinct stomach; intestine with
spiral fold
10. Sense organs of taste, smell, hearing; eyes well-developed
in adult; two pairs semicircular canals
11. Sexes separate; single gonad without duct; external
fertilization; long larval stage (ammocoete)
Tongue protruded for

rasping flesh
chordate characteristics in such simplified and easily visual-
ized form that it has been considered a chordate archetype
Figure 24.6
How a lamprey uses its keratinized tongue to feed. After firmly
(p. 509). After absorbing the remainder of their yolk supply, attaching to a fish by its buccal funnel, the protrusible tongue rapidly
young ammocoetes, now about 7 mm long, leave the nest gravel rasps an opening through the fish’s integument. Body fluid, abraded
and drift downstream to burrow in a suitable sandy, low-current skin, and muscle are eaten.
area. The larvae live as suspension feeders while growing slowly
for 3 to 7 or more years, then rapidly metamorphose into adults. lampreys began to decline, due in part to depletion of their food
This change involves the eruption of eyes, replacement of the and in part to expensive control measures (mainly chemical lar-
hood by the oral disc with keratinized teeth, enlargement of fins, vacides placed in selective spawning streams). Lake trout, aided
maturation of gonads, and modification of the gill openings. by a restocking program, are now recovering, but wounding
Parasitic lampreys either migrate to the sea, if marine, or rates are still high in some lakes.
remain in freshwater, where they attach themselves by their
suckerlike mouth to a fish and, with their sharp keratinized teeth,
rasp through the flesh and suck out body fluids (Figure 24.6). To CLASS CHONDRICHTHYES:
promote the flow of blood, the lamprey injects an anticoagulant
into the wound. When gorged, the lamprey releases its hold but
CARTILAGINOUS FISHES
leaves the fish with a large, gaping wound that can be fatal. There are about 970 living species in class Chondrichthyes, an
Parasitic freshwater adults live 1 to 2 years before spawning and ancient group that appeared in the Devonian period. Although a
then die; anadromous forms live 2 to 3 years. much smaller and less diverse assemblage than bony fishes, their
Nonparasitic lampreys do not feed after emerging as adults impressive combination of well-developed sense organs, powerful
and their digestive tract degenerates to a nonfunctional strand of jaws and swimming musculature, and predaceous habits ensures
tissue. Within a few months they spawn and die. them a secure and lasting place in the aquatic community. One
Invasion of the Great Lakes by the sea lamprey, Petromyzon of their distinctive features is a cartilaginous skeleton. Although
marinus, in this century has had a devastating effect on fisheries. calcification may be extensive in their skeletons, bone is entirely
No lampreys were present in the Great Lakes west of Niagara absent throughout the class—a curious evolutionary feature, since
Falls until the Welland Ship Canal was deepened between 1913 Chondrichthyes are derived from ancestors having well-developed
and 1918, allowing lampreys to bypass the Falls. Moving first bone. Although bone was lost in Chondrichthyes, possibly through
through Lake Erie to Lakes Huron, Michigan, and Superior, sea a process of neoteny (p. 508), phosphatized mineral tissues were
lampreys, accompanied by overfishing, caused a total collapse of retained in teeth, scales, and spines. Almost all chondrichthyans
a multimillion-dollar lake trout fishery in the early 1950s. Rain- are marine; only 28 species live primarily in freshwater.
bow trout, lake whitefish, lake herring, chubs, and other species Except for whales, sharks include the largest living verte-
were destroyed in turn. After reaching peak abundance in 1951 brates. The larger sharks may reach 12 m in length. Dogfish sharks
in Lakes Huron and Michigan and in 1961 in Lake Superior, sea commonly studied in zoological laboratories rarely exceed 1 m.
Subclass Elasmobranchii: Sharks,

The worldwide shark fishery is experiencing unprecedented pres-
Skates, and Rays sure, driven by the high price of shark fins for shark-fin soup, an
The 13 living orders of elasmobranchs number about 937 species. Asian delicacy (which commonly sells for up to $100.00 per bowl).
Coastal waters are dominated by ground sharks, order Carcha- Coastal shark populations in general have declined so rapidly that
rhiniformes, which contains typical-looking sharks such as tiger “finning” has been outlawed in the United States; other countries,
and bull sharks and more bizarre forms, including hammerheads too, are setting quotas to protect threatened shark populations.
(Figure 24.7). Order Lamniformes contains several large, pelagic Even in the Marine Resources Reserve of the Galápagos Islands,
sharks dangerous to humans, including great white and mako one of the world’s exceptional wild places, tens of thousands of
sharks. Dogfish sharks, familiar to generations of comparative sharks have been killed illegally for the Asian shark-fin market.
anatomy students, are in order Squaliformes. Skates belong to the Contributing to the threatened collapse of shark fisheries worldwide
order Rajiformes, and several groups of rays (stingrays, eagle rays, is the low fecundity of sharks and the long time required by most
manta rays, and devil rays) belong to the order Myliobatiformes. sharks to reach sexual maturity; some species take as long as
Although most sharks are by nature timid and cautious, 35 years.
some of them are dangerous to humans. There are numer-
ous authenticated cases of shark attacks by great white sharks,
Carcharodon (Gr. karcharos, sharp, odous, tooth) (reaching Form and Function
6 m); mako sharks, Isurus (Gr. is, equal, ouros, tail); tiger Although to most people sharks have a sinister appearance and
sharks, Galeocerdo (Gr. galeos, shark, kerdō, fox); bull sharks, fearsome reputation, they are at the same time among the most
Carcharhinus leucas (Gr. Karcharos, sharp, rhinos, nose); gracefully streamlined of all fishes. The body of a dogfish shark
and hammerhead sharks, Sphyrna (Gr. sphyra, hammer). More (Figure 24.8) is fusiform (spindle-shaped). The asymmetrical het-
shark casualties have been reported from tropical and temper- erocercal tail, in which the vertebral column turns upward and
ate waters of the Australian region than from any other. During extends into the dorsal lobe of the caudal fin, provides thrust and
World War II there were several reports of mass shark attacks on some lift as it sweeps back and forth. There are paired pectoral
victims of ship sinkings in tropical waters. and pelvic fins supported by appendicular skeletons, one or two
A Hammerhead
shark F Shortfin mako shark
H Basking shark
D Bull shark
B Angel shark
C Nurse shark
E Sawshark
G Great white shark
I Whale shark
J Tiger shark K Thresher shark
L Spined pygmy
shark M Megamouth shark
Figure 24.7
Diversity in sharks of the subclass Elasmobranchii: A, hammerhead shark, Sphyrna; B, angel shark, Squatina; C, nurse shark, Ginglymostoma
cirratum; D, bull shark, Carcharhinus leucas; E, sawshark, Pristiophorus; F, shortfin mako shark, Isurus oxyrinchus; G, great white shark, Carcharodon
carcharias; H, basking shark, Cetorhinus maximus; I, whale shark, Rhincodon typus; J, tiger shark, Galeocerdo cuvier; K, thresher shark, Alopias
vulpinus; L, spined pygmy shark, Squaliolus laticaudus; and M, megamouth shark, Megachasma pelagios.
hic70049_ch24_514-542.indd 521 7/25/07 1:24:06 PM

median dorsal fins (each with a spine in Squalus [L. a kind of sea
Characteristics of Class Chondrichthyes fish]), and a median caudal fin. A median anal fin is present in
most sharks, including the smooth dogfish, Mustelus (L. mustela,
1. Large (average about 2 m), body fusiform, or dorsoventrally weasel). In males, the medial part of the pelvic fin is modified to
depressed, with a heterocercal caudal fin (diphycercal in form a clasper, which is used in copulation. Paired nostrils (blind
chimaeras) (see Figure 24.16); paired pectoral and pelvic fins; pouches) are ventral and anterior to the mouth (Figure 24.9). The
pelvic fins in male modified as “claspers”
lateral eyes are lidless, and behind each eye a spiracle (remnant of
2. Mouth ventral; two olfactory sacs that do not open into the
mouth cavity in elasmobranchs; nostrils open into mouth
the first gill slit) is usually present. Five gill slits are found anterior
cavity in chimaeras; jaws present to each pectoral fin. The tough, leathery skin is covered with tooth-
3. Skin with placoid scales (see Figure 24.17) or naked; like, dermal placoid scales arranged to reduce the turbulence of
teeth of modified placoid scales and polyphyodont water flowing along the body surface during swimming.
in elasmobranchs; teeth modified as grinding plates in Sharks are well equipped for their predatory life. They track
chimaeras their prey using highly sensitive senses in an orderly sequence.
4. Endoskeleton entirely cartilaginous; notochord persistent Sharks may initially detect prey from a kilometer or more away
but reduced; vertebrae complete and separate (vertebrae with their large olfactory organs, capable of detecting chemicals
present but centra absent in chimaeras) as low as 1 part per 10 billion. The laterally placed nostrils of
5. Digestive system with J-shaped stomach (stomach absent in hammerhead sharks (see Figure 24.7) may enhance odor localiza-
chimaeras); intestine with spiral valve; often with large oil-
tion by improving stereo-olfaction. Prey also may be located from
filled liver for buoyancy
6. Circulatory system of several pairs of aortic arches; single
long distances by sensing low-frequency vibrations with mecha-
circulation; hepatic portal and renal portal systems; heart noreceptors in the lateral-line system. This system is composed
with sinus venosus, atrium, ventricle, and conus arteriosus of special receptor organs (neuromasts) in interconnected tubes
7. Respiration by means of five to seven pairs of gills leading and pores extending along the sides of the body and over the
to exposed gill slits in elasmobranchs; four pairs of gills head (Figure 24.10). At closer range a shark switches to vision as
covered by an operculum in chimaeras the primary method of tracking prey. Contrary to popular belief,
8. No swim bladder or lung most sharks have excellent vision, even in dimly lit waters. Dur-
9. Opisthonephric kidney and rectal gland; blood isosmotic or ing the final stage of attack, sharks are guided to their prey by the
slightly hyperosmotic to seawater; high concentrations of bioelectric fields that surround all animals. Electroreceptors, the
urea and trimethylamine oxide in blood ampullae of Lorenzini (see Figure 24.9), are located primarily
10. Brain of two olfactory lobes, two cerebral hemispheres,
on the shark’s head. In addition, sharks may use electroreception
two optic lobes, cerebellum, medulla oblongata; 10 pairs of
cranial nerves; three pairs of semicircular canals; senses
to find prey buried in the sand.
of smell, vibration reception (lateral-line system), vision, and Both upper and lower jaws of sharks are provided with
electroreception well-developed many sharp teeth. The front row of functional teeth on the edge
11. Sexes separate; gonads paired; reproductive ducts open into of the jaw is backed by rows of developing teeth that replace
cloaca (separate urogenital and anal openings in chimaeras); worn teeth throughout the life of the shark (see Figures 24.8
oviparous, ovoviviparous, or viviparous; direct development; and 24.9). The mouth cavity opens into a large pharynx, which
fertilization internal contains openings to separate gill slits and spiracles. A short,
wide esophagus runs to the J-shaped stomach. A liver and
pancreas open into a short, straight intestine, which contains
First Second
Nostril Spiracle Spine dorsal fin dorsal fin Caudal
fin
Rostrum
Clasper
Lateral line Pelvic fin

External
gill openings
Pectoral fin
Figure 24.8
Tooth Male spiny dogfish shark, Squalus acanthias. Inset: section of lower
shed jaw shows new teeth developing inside the jaw. These move forward
to replace lost teeth. Rate of replacement varies among species.
hic70049_ch24_514-542.indd 522 7/25/07 1:24:09 PM

tendrils that wrap around the first firm object it contacts, much
like tendrils of grape vines. Embryos are nourished from the yolk
for a long period—6 to 9 months in some, as much as 2 years
in one species—before hatching as miniature replicas of adults.
Many sharks, however, retain embryos in their reproductive tract
for prolonged periods. Many are ovoviviparous (lecithotrophic
viviparous) species, which retain developing young in the uterus
while they are nourished by contents of their yolk sac until born.
Still other species have true viviparous reproduction. In these,
embryos receive nourishment from the maternal bloodstream
through a placenta, or from nutritive secretions, “uterine milk,”
produced by the mother. Some sharks, including sand tigers,
exhibit a grisly type of reproduction in which embryos receive
additional nutrition by eating eggs and siblings. The evolution of
retention of embryos by many elasmobranchs was an important
Figure 24.9 innovation that contributed to their success. Regardless of the
Head of sand tiger shark, Carcharias sp. Note the series of successional
teeth. Also visible below the eye are ampullae of Lorenzini. initial amount of maternal support, all parental care ends once
eggs are laid or young are born.
Marine elasmobranchs have developed an interesting solu-
the spiral valve that slows passage of food and increases the tion to the physiological problem of living in a salty medium. To
absorptive surface (Figure 24.11). Attached to a short rectum is a prevent water from being drawn out of the body osmotically, elas-
rectal gland, unique to chondrichthyans, which secretes a col- mobranchs retain nitrogenous compounds, especially urea and
orless fluid containing a high concentration of sodium chloride. trimethylamine oxide, in their extracellular fluid. These solutes,
The rectal gland assists the opisthonephric kidney in regulat- combined with the blood salts, raise the blood solute concentra-
ing salt concentration of the blood. Chambers of the heart are tion to exceed slightly that of seawater, eliminating an osmotic
arranged in tandem formation, and blood circulates in the same inequality between their bodies and surrounding seawater.
pattern seen in other gill-breathing vertebrates (Figure 24.11). More than half of all elasmobranchs are rays, a group that
Blood leaving the heart via the ventral aorta enters capillary includes skates, electric rays, sawfishes, stingrays, and manta rays.
beds in the gills where oxygen is absorbed, then circulated to Most are specialized for bottom dwelling, with a dorsoventrally
the rest of the body via the dorsal aorta, without first reentering flattened body and greatly enlarged pectoral fins, which they
the heart (see Figure 31.10A, p. 692). move in a wavelike fashion to propel themselves (Figure 24.12).
All chondrichthyans have internal fertilization, but maternal Gill openings are on the underside of the head, but the large spir-
support of embryos is highly variable. Some sharks and all skates acles are on top. Respiratory water enters through these spiracles
lay large, yolky eggs immediately after fertilization; these species to prevent clogging the gills, because the mouth is often buried in
are termed oviparous. Some deposit their eggs in a horny cap- sand. Teeth are adapted for crushing prey: molluscs, crustaceans,
sule called a “mermaid’s purse,” which often is provided with and an occasional small fish.
Lateral line Ampullae

of Lorenzini
Opening to surface Lateral-line canal
Pore
Jelly-filled
canal
Neuromast Ampulla of
cells Lorenzini
Nerve
Figure 24.10
Sensory canals and receptors in a shark. The ampullae of Lorenzini respond to weak electric fields, and possibly to temperature, water pressure, and
salinity. The lateral-line sensors, called neuromasts, are sensitive to disturbances in the water, enabling a shark to detect nearby objects by reflected
waves in the water.
hic70049_ch24_514-542.indd 523 7/25/07 1:24:10 PM

Figure 24.11 Celiac Kidney First

dorsal
Dorsal artery Stomach
Internal anatomy of a dogfish Efferent fin
aorta Testis
shark, Squalus acanthias. branchial
artery Spleen Caudal
Hindbrain Caudal vein
artery
Midbrain
Forebrain
Rostrum
Rectal
Afferent Heart Pelvic
Liver Intestine gland Cloaca
branchial Ventral fin
artery aorta Spiral
Pectoral Pancreas valve
fin
Stingrays have a slender and whiplike tail armed with one or

more saw-edged spines with venom glands at the base. Wounds
from the spines are excruciatingly painful, and may heal slowly
and with complications. Electric rays are sluggish fish with large
electric organs on each side of their head (Figure 24.13). Each
organ is composed of numerous vertical stacks of disclike cells
connected in parallel so that when all cells discharge simultane-
ously, a high-amperage current is produced that flows out into
the surrounding water. The voltage produced is relatively low
(50 volts) but power output may be almost one kilowatt—quite
sufficient to stun prey or discourage predators. Electric rays were
Olfactory lobes
Cerebellum
A
Left electric Cranial
organ nerves
Right electric
organ
Spinal cord
Figure 24.13
Electric ray, Torpedo, with electric
organs exposed. Organs are built
up of disclike, multinucleated cells
B called electrocytes. When all cells are
discharged simultaneously, a high-
Figure 24.12 amperage current flows into the
Skates and rays are specialized for life on the seafloor. Both clearnose surrounding water to stun prey or
skates, Raja eglanteria (A), and southern stingrays, Dasyatis americana discourage predators. Electrosensory
(B), are flattened dorsoventrally and move by undulations of winglike information is processed in the large
pectoral fins. This stingray (B) is followed by a pilot fish. cerebellum.
hic70049_ch24_514-542.indd 524 7/25/07 1:24:13 PM

term as a valid taxon. Rather, it is used as a term of convenience

to describe vertebrates with endochondral bone that are
conventionally termed “bony fishes.”
Fossils of the earliest bony fishes show similarities in several
craniopharyngeal structures, including a bony operculum and
branchiostegal rays, with acanthodians (p. 512 and Figure 23.17),
indicating they likely descended from a common ancestor. By
the middle Devonian bony fishes already had radiated exten-
sively into two major groups, with adaptations that fitted them
for every aquatic habitat except the most inhospitable. One of
these groups, ray-finned fishes (class Actinopterygii), includes
modern bony fishes (Figure 24.15), the most species-rich group
of living vertebrates. A second group, lobe-finned fishes (class
Sarcopterygii), is represented today by only eight fishlike ver-
Figure 24.14 tebrates, the lungfishes and coelacanths (see Figures 24.22 and
Spotted ratfish, Hydrolagus collei, of North American west coast. This 24.23); however, it includes the sister group of land vertebrates
species is one of the most handsome of chimaeras, which tend toward
bizarre appearances.
(tetrapods).
Several key adaptations contributed to the radiation of bony
fishes. They have an operculum over the gill composed of
used by ancient Egyptians for a form of electrotherapy in the bony plates and attached to a series of muscles. This feature
treatment of afflictions such as arthritis and gout. increases respiratory efficiency because outward rotation of the
operculum creates a negative pressure so that water would be
drawn across the gills, as well as pushed across by the mouth
Subclass Holocephali: Chimaeras pump (see Figure 31.20). A gas-filled derivative of the esophagus
Members of the small subclass Holocephali, distinguished by provides an additional means of gas exchange in hypoxic
such suggestive names as ratfish ( Figure 24.14 ), rabbitfish, waters and an efficient means for achieving neutral buoyancy.
spookfish, and ghostfish, are remnants of a line that diverged In fishes that use these pouches primarily for gas exchange, the
from the shark lineage at least 360 million years ago. Fossil pouches are called lungs, while in fishes that use these pouches
chimaeras (ky-meer-uz) first occurred in the Devonian period, primarily for buoyancy, the pouches are called swim bladders
reached their zenith in the Cretaceous and early Tertiary periods (pp. 531–532). Progressive specialization of jaw musculature and
(120 million to 50 million years ago), and have declined ever skeletal elements involved in feeding is another key feature of
since. Today there are only about 33 species extant. bony fish evolution.
Anatomically chimaeras have several features linking them
to elasmobranchs, but they possess a suite of unique charac-
ters, too. Instead of a toothed mouth, their jaws bear large flat Class Actinopterygii: Ray-Finned Fishes
plates. The upper jaw is completely fused to the cranium, a most Ray-finned fishes are an enormous assemblage containing all our
unusual feature in fishes. Their food includes seaweed, molluscs, familiar bony fishes—almost 27,000 species. Earliest actinopter-
echinoderms, crustaceans, and fishes—a surprisingly mixed diet ygians, known as palaeoniscids (paylee-o-nisids), were small
for such a specialized grinding dentition. Chimaeras are not fishes, with large eyes, heterocercal caudal fin (Figure 24.16),
commercial species and are seldom caught. Despite their bizarre and thick, interlocking scales with an outer layer of ganoin
shape, they are beautifully colored with a pearly iridescence. (Figure 24.17). These fishes had a single dorsal fin and numerous
bony rays derived from scales stacked end to end, distinctively
different in appearance from the lobe-finned fishes with which
OSTEICHTHYES: BONY FISHES they shared the Devonian waters. Palaeoniscids are represented
by fossil fragments as early as the late Silurian, and flourished
Origin, Evolution, and Diversity throughout the late Paleozoic era, during the same period that
In the early to middle Silurian, a lineage of fishes with bony ostracoderms, placoderms, and acanthodians disappeared and
endoskeletons gave rise to a clade of vertebrates that contains sarcopterygians declined in abundance (see Figure 24.1).
96% of living fishes and all living tetrapods. Fishes of this clade From those earliest ray-finned fishes arose several clades.
have traditionally been termed “bony fishes” (Osteichthyes); Bichirs, in the clade Cladistia, have lungs, heavy ganoid scales,
bony fishes and tetrapods are united by the presence of endo- and other characteristics similar to those of palaeoniscids
chondral bone (bone that replaces cartilage developmentally, (Figure 24.18). The 16 species of bichirs live in freshwaters of
p. 650), presence of lungs or a swim bladder derived from the Africa. A second group are the chondrosteans (Gr. chondros,
gut, and several cranial and dental characters. Because tradi- cartilage, osteon, bone), represented by 27 species of freshwa-
tional usage of Osteichthyes does not describe a monophyletic ter and anadromous sturgeons and paddlefishes (Figure 24.18).
(natural) group (see Figure 24.2), most recent classifications, Nearly all chondrosteans have experienced severe population
including the one presented on page 540, do not recognize this declines from dam construction, overfishing, and pollution.
Myomere
(muscle segment) Swim bladder
First dorsal fin

Kidney Second dorsal fin
Fin ray support (opisthonephros)
Vertebra Lateral line

Spinal cord
Brain
Efferent
branchial artery
Olfactory Caudal
bulb fin
Afferent
branchial artery
Bulbus arteriosus
Anal fin
Ventricle Liver Ovary
Stomach Urinary
Pyloric ceca Anus bladder Figure 24.15
Spleen
Urogenital Anatomy of a yellow perch, Perca
Pelvic fin Intestine opening flavescens, a freshwater teleost fish.
The major clade of neopterygians are teleosts (Gr. teleos,

perfect, osteon, bone ), the modern bony fishes (see Fig-
ure 24.15). Teleost diversity is astounding, with almost 27,000
described species, representing about 96% of all living fishes or
about half of all vertebrates (Figure 24.20). In addition, it has
been estimated that there are an additional 5,000 to 10,000 unde-
scribed species. Although most of the 200 or so new species
Heterocercal Homocercal of teleosts described each year are from poorly sampled areas
(shark) (perch) such as South America or deep oceanic waters, several new spe-
cies are described each year from areas as well known as the
freshwaters of North America! Teleosts range in size from 7 mm
Diphycercal
(lungfish) adult minnows to 17 m oarfish and 900 kg, 4.5 m blue marlin
(Figure 24.20). These fishes occupy almost every conceivable
Figure 24.16 habitat, from elevations up to 5200 m in Tibet to 8000 m below
Types of caudal fins among fishes. the surface of the ocean. Some species live in hot springs at
44 C, while others live under the Antarctic ice at 2 C. They
The third major group of ray-finned fishes to emerge from may live in lakes with salt concentrations three times that of
palaeoniscid stock were the neopterygians (Gr. neos, new, seawater, caves of total darkness, swamps devoid of oxygen, or
pteryx, fin). Neopterygians appeared in the late Permian and even make extended excursions onto land, as do mudskippers
radiated extensively during the Mesozoic era (see Figure 24.1). (Figure 24.20).
During the Mesozoic one lineage gave rise to a secondary radi- Several morphological trends in the teleost lineage allowed
ation that led to modern bony fishes, the teleosts. There are it to diversify into this truly incredible variety of habitats and
two surviving genera of early neopterygians, the bowfin, Amia forms. Heavy dermal armor of primitive ray-finned fishes was
(Gr. tunalike fish), of shallow, weedy waters of the Great Lakes replaced by light, thin, flexible cycloid and ctenoid scales
and Mississippi River basin, and gars, Lepisosteus (Gr. lepidos, (see Figure 24.17). Some teleosts, such as most eels and cat-
scale, osteon, bone), of eastern and southern North America fishes, completely lack scales. Increased mobility and speed that
(Figure 24.19). The seven species of gars are large, ambush resulted from loss of heavy armor improved predator avoidance
predators with elongate bodies and jaws filled with needlelike and feeding efficiency. Changes in the fins of teleosts increased
teeth. Gars and bowfin may surface to gulp air, filling their maneuverability and speed and allowed fins to serve a variety
vascularized swim bladder with air to supplement oxygen of other functions. The symmetrical shape of the homocer-
obtained in the gills. cal tail (see Figure 24.16) of most teleosts focused musculature
hic70049_ch24_514-542.indd 526 7/25/07 1:24:24 PM

Dentine Enamel
Epidermis
Basal plate Pulp cavity
Ganoid scales Cycloid scales Ctenoid scales

(nonteleost bony fishes) (teleost fishes) (teleost fishes)
Placoid scales
(cartilaginous fishes)
Figure 24.17
Types of fish scales. Placoid scales are small, conical toothlike structures characteristic of Chondrichthyes. Diamond-shaped ganoid scales, present
in early bony fishes such as the gar, are composed of layers of silvery enamel (ganoin) on the upper surface and bone on the lower. Teleosts have
either cycloid or ctenoid scales. These are thin and flexible and are arranged in overlapping rows.
Figure 24.18
Primitive ray-finned
fishes of class
Actinopterygii. A Bichir
A, Bichir, Polypterus
bichir, of equatorial
West Africa. It is a
nocturnal predator.
B, Atlantic sturgeon,
Acipenser oxyrhynchus
(now uncommon),
of Atlantic coastal
rivers. C, Paddlefish,
B Atlantic sturgeon
Polyodon spathula, of
the Mississippi River
basin reaches 2 m and
A
80 kg.
C Paddlefish
contractions on the tail, permitting greater speed. The dorsal fin

shifted from a fixed keel that primarily prevented rolling, to a
flexible and highly specialized structure in derived teleosts (see
Figure 24.15). These changes in the morphology of fins were
useful for camouflage, braking and other complex movements, B
streamlining, and social communication. Bizarre modifications of
the dorsal fin include the lure of anglerfishes, venom-delivering
Figure 24.19
Nonteleost neopterygian fishes. A, Bowfin, Amia calva. B, Longnose
spines of scorpionfishes, and suctorial disc of sharksuckers (Fig- gar, Lepisosteus osseus. They frequent slow-moving streams and
ure 24.20). Teleost lineages demonstrated an increasingly fine swamps of eastern North America where they may hang motionless in
control of gas resorption and secretion in the swim bladder. Con- the water, ready to snatch passing fishes.
trol of buoyancy likely coevolved with fin modifications to refine
locomotion. Several anatomical modifications improved feeding increased final attack velocity. Gill arches of many teleosts diver-
efficiency. Changes in jaw suspension enabled the orobranchial sified into powerful pharyngeal jaws for chewing, grinding,
cavity to expand rapidly, creating a highly sophisticated suction and crushing. With so many innovations teleosts have become
device. Rapid jaw protrusion by sliding the upper jaw forward the most diverse of fishes.
hic70049_ch24_514-542.indd 527 7/25/07 1:24:27 PM

Class Sarcopterygii: Lobe-Finned Fishes

Characteristics of Class Actinopterygii The ancestor of tetrapods is found within a group of extinct sar-
1. Skeleton with bone of endochondral origin; caudal copterygian fishes called rhipidistians, which included several
fin heterocercal in ancestral forms, usually homocercal in lineages that flourished in freshwaters and shallow coastal areas
descendant forms (Figure 24.16); skin with mucous glands in the late Paleozoic. Rhipidistians, such as Eusthenopteron (see
and embedded dermal scales (Figure 24.21); scales ganoid in Figure 25.2, p. 546), were cylindrical, large-headed fishes with
ancestral forms, scales cycloid, ctenoid or absent in derived fleshy fins, and presumably lungs. The evolution of tetrapods
forms (Figure 24.17) from rhipidistians is discussed in Chapter 25.
2. Paired and median fins present, supported by long dermal All early sarcopterygians had lungs as well as gills, and a
rays (lepidotrichia); muscles controlling fin movement tail of the heterocercal type. However, during the Paleozoic
within body
the orientation of the vertebral column changed so that the tail
3. Jaws present; teeth usually present with enamaloid covering;
olfactory sacs do not open into mouth; spiral valve present in
became a symmetrical diphycercal tail (see Figure 24.16). These
ancestral forms, absent in derived forms sarcopterygians had powerful jaws; heavy, enameled scales with
4. Respiration primarily by gills supported by arches and a dentinelike material called cosmine; and strong, fleshy, paired
covered with an operculum lobed fins that may have been used to clamber over benthic sub-
5. Swim bladder often present with or without a duct strates filled with woody debris. The sarcopterygian clade today
connecting to esophagus, usually functioning in buoyancy is represented by only eight fish species: six species of lungfishes
6. Circulation consisting of a heart with a sinus venosus, and two species of coelacanths (Figures 24.22 and 24.23).
an undivided atrium, and an undivided ventricle; single Of the three surviving genera of lungfishes, most simi-
circulation; typically four aortic arches; nucleated lar to early forms is Neoceratodus (Gr. neos, new, keratos,
erythrocytes horn, odes, form), the living Australian lungfish, which may
7. Excretory system of paired opisthonephric kidneys; sexes
attain a length of 1.5 m (Figure 24.22). This lungfish, unlike its
usually separate; fertilization usually external; larval forms
may differ greatly from adults
relatives, normally relies on gill respiration, and cannot survive
8. Nervous system of a brain with small cerebrum, optic lobes, long out of water. The South American lungfish, Lepidosiren
and cerebellum; 10 pairs of cranial nerves; three pairs of (L. lepidus, pretty, siren, mythical mermaid), and the African
semicircular canals lungfish, Protopterus (Gr. prōtos, first, pteron, wing), can live
out of water for long periods of time. Protopterus lives in African
streams and ponds that may dry during the dry season, with
A B
Figure 24.20
Diversity among teleosts. A, Blue marlin,
Makaira nigricans, one of the largest
teleosts. B, Mudskippers, Periophthalmus sp.,
make extensive excursions on land to graze
on algae and capture insects; they build nests
in which the young hatch and are guarded
by the mother. C, Protective coloration of
the flamboyant lionfish, Pterois sp., advises
caution; the dorsal spines are venomous.
D, The sucking disc on the sharksucker,
Echeneis naucrates, is a modification of the
dorsal fin. C D
hic70049_ch24_514-542.indd 528 7/25/07 1:24:33 PM

Bony part Mucous Epidermis Dermis oxygen include a lung for respiratory gas exchange and partially
of scale glands separated pulmonary and systemic cardiovascular circuits.
Coelacanths also arose in the Devonian period, radiated
somewhat, and reached their peak of diversity in the Mesozoic
era. At the end of the Mesozoic era they nearly disappeared but
left one remarkable surviving genus, Latimeria (Figure 24.23).
Because the last coelacanths were believed to have become
extinct 70 million years ago, the scientific world was astonished
when the remains of a coelacanth were found on a dredge off the
Figure 24.21 coast of South Africa in 1938. An intensive search to locate more
Section through the skin of a bony fish, showing the overlapping scales
(yellow). The scales lie in the dermis and are covered by epidermis.
Australian lungfish African lungfishes South American lungfish
Figure 24.22
Lungfishes are lobe-finned fishes of class Sarcopterygii. The Australian lungfish, Neoceratodus forsteri, is the least specialized of three lungfish
genera. The African lungfishes, Protopterus sp., are best adapted of the three for remaining dormant in mucus-lined cocoons breathing air during
prolonged periods of drought.
specimens was successful off the coast of the Comoro Islands.

There fishermen occasionally catch them at great depths with
hand lines, providing specimens for research. This was the only
known population of Latimeria until 1998, when the scientific
world again was surprised by the capture of a new species of
coelacanth in Sulawesi, Indonesia, 10,000 km from the Comoros!
The “modern” marine coelacanths are descendants of
the Devonian freshwater stock. The tail is diphycercal (see
Figure 24.16) but possesses a small lobe between the upper
and lower caudal lobes, producing a three-pronged structure
(Figure 24.23).
Coelacanths are a deep metallic blue with irregular white or
AFRICA Philippine brassy flecks, providing camouflage against the dark lava-cave
Islands reefs they inhabit. Young are born fully formed after hatching
Comoro
Islands
internally from eggs 9 cm in diameter—the largest among bony
fishes.
AUSTRALIA
Madagascar STRUCTURAL AND FUNCTIONAL
ADAPTATIONS OF FISHES
Figure 24.23
The coelacanth genus Latimeria is a surviving marine relict of a group Locomotion in Water
of lobe-finned fishes that flourished some 350 million years ago. To the human eye, some fishes appear capable of swimming at
extremely high speeds. But our judgment is unconsciously tem-
their mud beds baked hard by the hot tropical sun. The fish pered by our own experience that water is a highly resistant
burrows down at the approach of the dry season and secretes a medium through which to move. Most fishes, such as a trout or a
copious slime mixed with mud to form a hard cocoon in which minnow, can swim maximally about 10 body lengths per second,
it estivates until rains return. Adaptations for using atmospheric obviously an impressive performance by human standards. When
waves of contraction moving backward along its body by alter-

nate contraction of myomeres on either side. The anterior end of
Characteristics of Class Sarcopterygii the body bends less than the posterior end, so that each undula-
1. Skeleton with bone of endochondral origin; caudal tion increases in amplitude as it travels along the body. While
fin diphycercal in living representatives, heterocercal undulations move backward, bending of the body pushes later-
in ancestral forms; skin with embedded dermal scales ally against the water, producing a reactive force that is directed
(Figure 24.21) with a layer of dentinelike material, cosmine,
forward, but at an angle. It can be analyzed as having two com-
in ancestral forms
ponents: thrust, which is used to overcome drag and propels the
2. Paired and median fins present; paired fins with a single
basal skeletal element and short dermal rays; muscles that fish forward, and lateral force, which tends to make the fish’s
move paired fins located on appendage head “yaw,” or deviate from the course in the same direction
3. Jaws present; teeth are covered with true enamel and as its tail. This side-to-side head movement is very obvious in a
typically are crushing plates restricted to palate; olfactory swimming eel or shark, but many fishes have a large, rigid head
sacs paired, may or may not open into mouth; intestine with with enough surface resistance to minimize yaw.
spiral valve The movement of an eel is reasonably efficient at low
4. Gills supported by bony arches and covered with an speed, but its body shape generates too much frictional drag
operculum for rapid swimming. Fishes that swim rapidly, such as trout, are
5. Swim bladder vascularized and used for respiration and less flexible and limit body undulations mostly to the caudal
buoyancy (fat-filled in coelacanths)
region (Figure 24.25). Muscle force generated in the large ante-
6. Circulation consisting of heart with a sinus venosus, two
rior muscle mass is transferred through tendons to the relatively
atria, a partly divided ventricle, and a conus arteriosus;
double circulation with pulmonary and systemic circuits; nonmuscular caudal peduncle and tail where thrust is gener-
characteristically five aortic arches ated. This form of swimming reaches its highest development
7. Nervous system with a cerebrum, a cerebellum, and optic in tunas, whose bodies do not flex at all. Virtually all thrust is
lobes; 10 pairs of cranial nerves; three pairs of semicircular derived from powerful beats of the caudal fin (Figure 24.26).
canals Many fast oceanic fishes such as marlin, swordfish, amberjacks,
8. Sexes separate; fertilization external or internal and wahoo have swept-back caudal fins shaped like a sickle.
Such fins are the aquatic counterpart of the high aspect ratio
wings of the swiftest birds (p. 601).
these speeds are translated into kilometers per hour it means that
a 30 cm (1 foot) trout can swim only about 10.4 km (6.5 miles) per
The body temperature of most fishes is the same as their envi-
hour. As a general rule, the larger the fish the faster it can swim.
ronment, because any heat generated internally quickly is lost
into the surrounding water. However, some fishes, such as tunas
Measuring fish cruising speeds accurately is best done in a “fish (Figure 24.26) and mako sharks, maintain a high temperature in
wheel,” a large ring-shaped channel filled with water that is turned their swimming muscles and viscera—as much as 10 C warmer
at a speed equal and opposite to that of the fish. Much more dif- than surrounding water. Marlins (see Figure 24.20A) and other bill-
ficult to measure are the sudden bursts of speed that most fish can fishes elevate the temperature of their brain and retina. Research by
make to capture prey or to avoid being captured. A hooked bluefin F. G. Carey and others explains how these fishes accomplish this
tuna was once “clocked” at 66 km per hour (41 mph); swordfish kind of thermoregulation, called regional endothermy. Heat is
and marlin are thought to be capable of incredible bursts of speed generated as a by-product of various activities, including digestion
approaching, or even exceeding, 110 km per hour (68 mph). Such and swimming, or for billfishes, by a specialized heat-generating
high speeds can be sustained for no more than 1 to 5 seconds. organ beneath the brain. This heat is conserved with a rete mira-
bile, a parallel bundle of blood vessels arranged to provide a coun-
tercurrent flow of blood (p. 682). High temperatures apparently
The propulsive mechanism of a fish is its trunk and tail mus-
promote powerful swimming and enhance digestive and nervous
culature. The axial, locomotory musculature is composed of zig-
system activity. Fishes with regional endothermy are the fastest in
zag bands, called myomeres. Muscle fibers in each myomere
the world.
are relatively short and connect the tough connective tissue par-
titions that separate each myomere from the next. On the surface
myomeres take the shape of a W lying on its side (Figure 24.24) Swimming is the most economical form of animal locomo-
but internally the bands are complexly folded and nested so that tion, largely because aquatic animals are almost perfectly sup-
the pull of each myomere extends over several vertebrae. This ported by their medium and need to expend little energy to
arrangement produces more power and finer control of move- overcome the force of gravity. If we compare the energy cost per
ment since many myomeres are involved in bending a given kilogram of body weight of traveling 1 km by different forms of
segment of the body. locomotion, we find swimming costs only 0.39 kcal (salmon) as
Understanding how fishes swim can be approached by study- compared with 1.45 kcal for flying (gull) and 5.43 kcal for walk-
ing the motion of a very flexible fish such as an eel (Figure 24.25). ing (ground squirrel). However, part of the unfinished business
The movement is serpentine, not unlike that of a snake, with of biology is understanding how fish and aquatic mammals are
hic70049_ch24_514-542.indd 530 7/25/07 1:24:47 PM

able to move through water while creating almost

Myomeres no turbulence. The secret lies in the way aquatic
animals bend their bodies and fins (or flukes) to
swim and in the friction-reducing properties of
the body surface.
Neutral Buoyancy and the

Swim Bladder
All fishes are slightly heavier than water because
Figure 24.24 their skeletons and other tissues contain heavy
Trunk musculature of a teleost fish, partly dissected to show internal arrangement of the elements present only in trace amounts in natu-
muscle bands (myomeres). The myomeres are folded into a complex, nested grouping, an ral waters. To keep from sinking, sharks, which
arrangement that favors stronger and more controlled swimming.
lack a swim bladder, must always keep moving
forward in the water. The asymmetrical (hetero-
cercal) tail of a shark provides lift as it sweeps through the water,
Eel Trout and its broad head and flat pectoral fins (see Figure 24.8) act as
angled planes to provide additional lift. Sharks also are aided in
buoyancy by having very large livers containing a special fatty
hydrocarbon called squalene with a density of only 0.86 grams
per milliliter. The liver thus acts like a large sack of buoyant oil
that helps to compensate the shark’s heavy body.
Thrust Reactive By far the most efficient flotation device is a gas-filled
force space. The swim bladder serves this purpose in bony fishes
(Figure 24.27). It arose from the paired lungs of primitive Devo-
nian bony fishes. Lungs were probably a ubiquitous feature of
Lateral Thrust
90˚ Devonian freshwater bony fishes when, as we have seen, warm,
force
Reactive swampy habitats would have made such an accessory respira-
force tory structure advantageous. Swim bladders are present in most
Push Lateral pelagic bony fishes but are absent in tunas, most abyssal fishes,
force 90˚
and most bottom dwellers, such as flounders and sculpins.
Without a swim bladder, bony fishes sink because their tis-
Push sues are denser than water. To achieve neutral buoyancy, they
displace additional water by a volume of gas in a swim bladder,
Figure 24.25 thus adjusting their total density to match that of the surround-
Movements of swimming fishes, showing the forces developed by an
ing water. This adjustment allows fishes with a swim bladder to
eel-shaped and spindle-shaped fish. remain suspended indefinitely at any depth with no muscular
Source: From Vertebrate Life, by Pough et al., 1996. Reprinted by effort. Unlike bone, blood, and other tissues, gas is compress-
permission of Prentice-Hall, Inc., Upper Saddle River, NJ. ible and changes volume as a fish changes its depth. If a fish
swims to a greater depth, the greater pressure exerted by the
surrounding water compresses the gas in the swim bladder, so
that the fish becomes less buoyant and begins to sink. The vol-
Body torpedo-shaped Caudal peduncle
ume of gas in the swim bladder must be increased to establish a
narrow
new equilibrium buoyancy. When a fish swims upward, gas in
the bladder expands because of the reduced surrounding water
Caudal fin pressure, making the fish more buoyant. Unless gas is removed,
sickle-shaped, the fish will continue to ascend with increasing speed as the
stiff swim bladder continues to expand.
Gas may be removed from the swim bladder in two ways.
The more primitive physostomous (Gr., phys, bladder,
stoma, mouth) fishes (trout, for example) have a pneumatic duct
Figure 24.26 that connects the swim bladder to the esophagus, through which
Bluefin tuna, Thunnus thynnus, showing adaptations for fast
swimming. Powerful trunk muscles pull on the slender caudal they may expel air. More derived teleosts exhibit the physoclis-
peduncle. Since the body does not bend, all of the thrust comes from tous (Gr., phys, bladder, clist, closed) condition in which the
beats of the stiff, sickle-shaped caudal fin. pneumatic duct is lost in adults. In physoclistous fishes, gas is
hic70049_ch24_514-542.indd 531 7/25/07 1:24:48 PM

charged steel gas cylinder. Yet the oxygen pressure in the fish’s
Swim bladder
blood cannot exceed 0.2 atmosphere—in equilibrium with the
oxygen pressure in the atmosphere at the sea surface.
Hearing and Weberian Ossicles

Fishes, like other vertebrates, detect sounds as vibrations in the
inner ear. Detecting these vibrations is difficult for aquatic ver-
tebrates because their bodies are nearly the same density as the
A surrounding water, letting sound waves pass through the fish’s
body nearly undetected.
A particularly elegant solution to this problem is found in the
Dorsal aorta
Ovale
ostariophysans, a group of teleosts that contains the minnows,
characins, suckers, and catfishes. The ostariophysans include
To about 7900 species and are usually the dominant fishes in diversity
heart and abundance in freshwater habitats. Their success may be partly
due to their Weberian ossicles, a set of small bones, which allow
Gas gland
them to hear faint sounds over a much broader range of frequency
B than other teleosts. Reception of sound begins at the swim blad-
Constrictor
Rete muscles
der, which can be easily vibrated because it is air-filled. Sound
mirabile vibrations are transmitted from the swim bladder to the inner ear
by the Weberian ossicles (Figure 24.28). This system has some
C similarity to the tympanum and middle ear bones of mammals
(pp. 744–745), but evolved independently. Adaptations to improve
Figure 24.27 hearing are not confined to ostariophysans. For example, herrings
A, Swim bladder of a teleost fish. The swim bladder lies in the coelom and anchovies have anterior expansions of the swim bladder that
just beneath the vertebral column. B, Gas is secreted into the swim directly contact the skull. The importance of the swim bladder in
bladder by the gas gland. Gas from the blood is moved into the
these fishes is demonstrated by experiments in which the swim
gas gland by the rete mirabile, a complex array of tightly-packed
capillaries that act as a countercurrent multiplier to increase oxygen bladder is artificially deflated, reducing sensitivity to sounds.
concentration. The arrangement of venous and arterial capillaries in
the rete is shown in C. To release gas during ascent, a muscular valve
opens, allowing gas to enter the ovale from which the gas is removed Respiration
by diffusion into the blood.
Fish gills are composed of thin filaments, each covered with
a thin epidermal membrane folded repeatedly into platelike
absorbed by blood from the ovale, a vascularized area of the lamellae (Figure 24.29). These are richly supplied with blood
swim bladder (Figure 24.27). Both types of fishes require gas to vessels. The gills are located inside the pharyngeal cavity and
be secreted into the swim bladder from the blood, although a are covered with a movable flap, the operculum. This arrange-
few shallow-water-inhabiting physostomes may gulp air to fill ment provides excellent protection to delicate gill filaments,
their swim bladder. streamlines the body, and makes possible a pumping system for
Physiologists who were at fi rst baffl ed by the secretion moving water through the mouth, across the gills, and out the
mechanism now understand how it operates. In brief, the gas operculum. Instead of opercular flaps as in bony fishes, elasmo-
gland secretes lactic acid, which enters the blood, causing a branchs have a series of gill slits (see Figure 24.8) out of which
localized high acidity in the rete mirabile that forces hemo- the water flows. In both elasmobranchs and bony fishes the
globin to release its load of oxygen. The capillaries in the rete branchial mechanism is arranged to pump water continuously
are arranged in parallel, creating a countercurrent multiplier and smoothly over the gills, although to an observer it appears
(p. 677), allowing oxygen to reach high concentrations in the that fish breathing is pulsatile. The flow of water is opposite the
gas gland and to diffuse into the swim bladder. The final gas direction of blood flow (countercurrent flow), the best arrange-
pressure attained in the swim bladder depends on the length ment for extracting the greatest possible amount of oxygen from
of the rete capillaries; they are relatively short in fishes living water. Some bony fishes can remove as much as 85% of the dis-
near the surface, but are extremely long in deep-sea fishes. solved oxygen from water passing over their gills. Very active
The amazing effectiveness of this device is exemplified by a fishes, such as herring and mackerel, can obtain sufficient water
fish living at a depth of 2400 m (8000 feet). To keep the bladder for their high oxygen demands only by swimming forward con-
inflated at that depth, the gas inside (mostly oxygen, but also tinuously to force water into their open mouth and across their
variable amounts of nitrogen, carbon dioxide, argon, and even gills. This process is called ram ventilation. Such fishes would be
some carbon monoxide) must have a pressure exceeding 240 asphyxiated if placed in an aquarium that restricts free-swimming
atmospheres, which is much greater than the pressure in a fully movements, even if the water is saturated with oxygen.
hic70049_ch24_514-542.indd 532 7/25/07 1:24:49 PM

Figure 24.28
Weberian ossicles are small bones that
Brain transmit sound vibrations received in the
swim bladder to the inner ear. Teleosts
with this apparatus can detect faint
sounds over a much broader range of
frequency than other fishes.
Semicircular
canals
Weberian
ossicles
Vertebrae
Ribs
Swim bladder
A surprising number of fishes can live out of water for vary- (Gr. ēlektron, something bright, phoros, to bear), have degener-
ing lengths of time by breathing air. Several devices are employed ate gills and must supplement gill respiration by gulping air through
by different fishes. We already have described the lungs of a vascular mouth cavity. One of the best air breathers of all is the
lungfishes, gars, and the extinct rhipidistians. Freshwater eels Indian climbing perch, Anabas (Gr. anabainō, to go up), which
often make overland excursions during rainy weather, using the spends most of its time on land near the water’s edge, breathing air
skin as a major respiratory surface. Electric eels, Electrophorus through special air chambers above much-reduced gills.
Gill filaments Gill raker

Filament with
Gill rakers B lamellae
A
Gill arch
Gill arch
Figure 24.29
Gills of fish. The bony, protective flap covering
the gills (operculum) has been removed, A, to
Blood C reveal branchial chamber containing the gills.
flow There are four gill arches on each side, each
bearing numerous filaments. A portion of a gill
arch (B) shows gill rakers that project forward
to strain out food and debris, and gill filaments
Lamella Water that project to the rear. A single gill filament
flow (C) is dissected to show the blood capillaries
within the platelike lamellae. Direction of
water flow (large arrows) is opposite the
Filament
direction of blood flow.
hic70049_ch24_514-542.indd 533 7/25/07 1:24:51 PM

Osmotic Regulation that a freshwater fish devotes only a small part of its total energy
maintaining itself in osmotic balance.
Freshwater is an extremely dilute medium with a salt concentra-
tion (0.001 to 0.005 gram moles per liter [M]) much below that
of the blood of freshwater fishes (0.2 to 0.3 M). Water therefore
tends to enter their bodies osmotically, and salt is lost by dif- Perhaps 90% of all bony fishes are restricted to either a freshwater
fusion outward. Although the scaled and mucus-covered body or a seawater habitat because they are incapable of osmotic regu-
surface is almost totally impermeable to water, water gain and lation in the “wrong” habitat. Most freshwater fishes quickly die
salt loss do occur across thin membranes of the gills. Freshwa- if placed in seawater, as will marine fishes placed in freshwater.
ter fishes are hyperosmotic regulators with several defenses However, some 10% of all teleosts can pass back and forth with
against these problems (Figure 24.30). First, excess water is ease between both habitats. These euryhaline fishes (Gr. eurys,
pumped out by the opisthonephric kidneys (p. 673), which broad, hals, salt) are of two types: those such as many flounders,
are capable of forming very dilute urine. Second, special salt- sculpins, and killifish that live in estuaries or certain intertidal areas
absorbing cells located in the gill epithelium actively move salt where the salinity fluctuates throughout the day; and those such
ions, principally sodium and chloride, from water to the blood. as salmon, shad, and eels, that spend part of their life cycle in
This absorption, together with salt present in the fish’s food, freshwater and part in seawater.
replaces diffusive salt loss. These mechanisms are so efficient
Large Active tubular

glomerulus reabsorption
of NaCl
FRESHW
FIS
Food,
freshwater
GILLS KIDNEYS
Active absorption of NaCl, Excretion
Intestinal of dilute urine
water enters osmotically
wastes Urine
Active tubular
Glomerulus secretion
Figure 24.30 of MgSO4
Osmotic regulation in freshwater and
marine bony fishes. A freshwater fish
maintains osmotic and ionic balance
in its dilute environment by actively
absorbing sodium chloride across the
gills (some salt is gained with food). To MARIN
flush out excess water that constantly FISH
enters the body, the glomerular kidney
produces a dilute urine by reabsorbing
Food,
sodium chloride. A marine fish must seawater
drink seawater to replace water lost
osmotically to its salty environment.
Sodium chloride and water are
absorbed from the stomach. Excess
sodium chloride is actively transported
outward by the gills. Divalent sea GILLS STOMACH INTESTINAL WASTES KIDNEYS
salts, mostly magnesium sulfate, are
Active secretion of Passive absorption MgSO4 voided Excretion of MgSO4,
eliminated with feces and secreted by
NaCl, water loss of NaCl and water with feces urea, little water
the tubular kidney.
hic70049_ch24_514-542.indd 534 7/25/07 1:24:53 PM

Marine bony fishes are hypoosmotic regulators that

encounter a completely different set of problems. Having a much
lower blood salt concentration (0.3 to 0.4 M) than the seawater
around them (about 1 M), they tend to lose water and gain salt.
A marine teleost fish quite literally risks drying out, much like
a desert mammal deprived of water. To compensate for water
loss, a marine teleost drinks seawater (Figure 24.30). Excess salt
accompanying the seawater is disposed in multiple ways. Major
sea salt ions (sodium, chloride, and potassium) are carried by
the blood to the gills where they are secreted outward by special
salt-secretory cells. The remaining sea salt ions, mostly mag-
nesium, sulfate, and calcium, are voided with feces or excreted
by the kidneys. Unlike a freshwater fish’s kidneys, which form
urine by the usual filtration-resorption sequence typical of most
vertebrate kidneys (pp. 673–677), a marine fish’s kidneys excrete
divalent ions by tubular secretion. Since very little if any filtrate is
formed, the glomeruli have lost their importance and disappeared
in some marine teleosts. Pipefishes and the frogfish shown in
Figure 24.32 are examples of “aglomerular” marine fishes. Figure 24.31
Wolf eel, Anarrhichthys ocellatus, feeding on a sea cucumber it has
captured and pulled to the opening of its den.
Feeding Behavior
For any fish, feeding is one of its main concerns in day-to-day
living. Although many a luckless angler would swear otherwise,
a fish devotes more time and energy to eating, or searching for
food to eat, than to anything else. Throughout the long evolution
of fishes, there has been unrelenting selective pressure for those
adaptations that enable a fish to win the eat-or-be-eaten contest.
Certainly the most far-reaching single event was the evolution
of jaws (p. 511). Jaws freed fishes from a largely passive filter-
feeding existence, enabling them to adopt a predatory mode of
life. Improved means of capturing larger prey demanded stron-
ger muscles, more agile movement, better balance, and improved
special senses. More than any other aspect of its life habit, feeding
behavior shapes the fish.
Most fishes are carnivores and prey on a myriad of animal
foods from zooplankton and insect larvae to large vertebrates.
Some deep-sea fishes are capable of eating victims nearly twice
their own size—an adaptation for life in a world where meals are
infrequent. Most advanced ray-finned fishes cannot masticate their
food as we can because doing so would block the current of water
across the gills. Some, however, such as wolf eels (Figure 24.31), Figure 24.32
have molarlike teeth in their jaws for crushing prey, which may Longlure frogfish, Antennarius multiocellatus, awaits its meal. Above
include hard-bodied crustaceans. Others that do grind their food its head swings a modified dorsal fin spine ending in a fleshy tentacle
use powerful pharyngeal teeth in their throat. Most carnivorous that contracts and expands in a convincing wormlike manner. When
fish swallow their prey whole, using sharp-pointed teeth in their a fish approaches the alluring bait, the huge mouth opens suddenly,
creating a strong inward current that sweeps the prey inside. The
jaws and on the roof of the mouth to seize prey. The incompress-
entire process takes only 4 milliseconds!
ibility of water assists many large-mouthed predators in captur-
ing prey. When the mouth is suddenly opened, water rushes in,
sweeping the victim inside (Figure 24.32).
A second group of fishes are herbivores that eat plants and Suspension feeders that crop the abundant microorgan-
macroalgae. Plant eaters are relatively uncommon among fishes, isms of the sea form a third and diverse group of fishes ranging
but are crucial intermediates in the food chain in some habitats. from fish larvae to basking sharks. However, the most charac-
Plant eaters are most common in coral reefs (parrotfishes, dam- teristic group of plankton feeders are herringlike fishes (menha-
selfishes, and surgeonfishes) and in tropical freshwater habitats den, herring, anchovies, capelin, pilchards, and others), mostly
(some minnows, characins, and catfishes). pelagic (open-sea dwellers) fishes that travel in large schools.
hic70049_ch24_514-542.indd 535 7/25/07 1:25:06 PM

Both phytoplankton and smaller zooplankton are strained from eel. They had been called leptocephali (Gr. leptos, slender,
the water with sievelike gill rakers (see Figure 32.1, p. 710). kephal¯e, head) by early naturalists, who never suspected
Because plankton feeders are the most abundant of all marine their true identity. In 1905 Johann Schmidt, supported by the
fishes, they are important food for numerous larger but less Danish government, began a systematic study of eel biology,
abundant carnivores. Many freshwater fishes also depend on which he continued until his death in 1933. With cooperation of
plankton for food. captains of commercial vessels plying the Atlantic, thousands of
Other groups of fishes include scavengers, such as hag- leptocephali were caught in different areas of the Atlantic with
fishes, that consume dead and dying animals, and detritivores, plankton nets Schmidt supplied. By noting where larvae in dif-
such as some suckers and minnows, that consume fine, particu- ferent stages of development were captured, Schmidt and his
late organic matter. Some fishes use a parasitic mode of feeding colleagues eventually reconstructed the spawning migrations.
in which they consume parts of other live fishes. Examples of When adult eels leave the coastal rivers of Europe and
these include lampreys (p. 518) and the candiru, Vandellia, a North America, they swim steadily and apparently at great depth
tiny elongate catfish that feeds on the gill epithelia of host fishes. for 1 to 2 months until they reach the Sargasso Sea, a vast area
Finally it should be noted that most fishes, though specialized of warm oceanic water southeast of Bermuda (Figure 24.33).
for a narrow diet, may use other foods when available. Here, at depths of 300 m or more, the eels spawn and die.
Digestion in most fishes follows the vertebrate plan. Except Minute larvae then begin an incredible journey back to the
in several fishes that lack distinct stomachs, food proceeds from streams of Europe and North America. Since the Sargasso Sea
stomach to tubular intestine, which tends to be short in carni- is much closer to the American coastline, American eel larvae
vores (see Figure 24.15) but may be extremely long and coiled in make their journey in only about eight months, compared to
herbivorous and detritivorous forms. In herbivorous grass carps, three years for European eel larvae. Males typically remain in
for example, the intestine may be nine times the body length, an brackish water of coastal rivers, while females migrate as far
adaptation for the lengthy digestion required for plant carbohy- as several hundred kilometers upstream. After 8 to 15 years of
drates. In carnivores, some protein digestion may be initiated in growth, females, now 1 m long, return to the ocean to join the
the acid medium of the stomach, but the principal function of smaller males in the journey back to the spawning grounds in
the stomach is to store often large and infrequent meals while the Sargasso Sea.
awaiting their reception by the intestine.
Digestion and absorption proceed simultaneously in
the intestine. A curious feature of ray-finned fishes, espe- Recent enzyme electrophoretic analysis of eel larvae confirmed not
cially teleosts, is the presence of numerous pyloric ceca (see only the existence of separate European and American species but
Figure 24.15) found in no other vertebrate group. Their primary also Schmidt’s belief that the European and American eels spawn in
function appears to be fat absorption, although all classes of partially overlapping areas of the Sargasso Sea.
digestive enzymes (protein-, carbohydrate-, and fat-splitting) are
secreted there.
Homing Salmon
Migration The life history of salmon is nearly as remarkable as that of
freshwater eels and certainly has received far more popular
Freshwater Eels attention. Salmon are anadromous (Gr. anadromos, run-
For centuries naturalists had been puzzled about the life his- ning upward); they spend their adult lives at sea but return to
tory of freshwater eels, Anguilla (an-gwilla) (L. eel), a com- freshwater to spawn. Atlantic salmon, Salmo salar (L. salmo,
mon and commercially important species of coastal streams of salmon, sal, salt), and Pacifi c salmon (six species in the
the North Atlantic. Eels are catadromous (Gr. kata, down, genus Oncorhynchus [on-ko-rinkus] [Gr. onkos, hook, rhyn-
dromos, running), meaning that they spend most of their lives chos, snout]) have this practice, but there are important dif-
in freshwater but migrate to the sea to spawn. Each fall, large ferences among the seven species. Atlantic salmon may make
numbers of eels were seen swimming down rivers toward the repeated upstream spawning runs. The six Pacific salmon spe-
sea, but no adults ever returned. Each spring countless num- cies (sockeye, coho, pink, Chinook, chum, and Japanese masu)
bers of young eels, called “elvers” (Figure 24.33), each about each make a single spawning run (Figure 24.34), after which
the size of a wooden matchstick, appeared in coastal rivers they die.
and began swimming upstream. Beyond the assumption that The virtually infallible homing instinct of the Pacific spe-
eels must spawn somewhere at sea, location of their breeding cies is legendary: after migrating downstream as a smolt,
grounds was completely unknown. (a juvenile stage) a sockeye salmon ranges many hundreds of
The first clue was provided by two Italian scientists, Grassi miles over the Pacific for nearly 4 years, grows to 2 to 5 kg
and Calandruccio, who in 1896 reported that elvers were not in weight, and then returns almost unerringly to spawn in the
larval eels but rather were relatively advanced juveniles. True headwaters of its parent stream. Some straying does occur and
larval eels, they discovered, were tiny, leaf-shaped, com- is an important means of increasing gene flow and populating
pletely transparent creatures that bore little resemblance to an new streams.
hic70049_ch24_514-542.indd 536 7/25/07 1:25:09 PM

American European
MIGRATION PATTERNS LARVAL STAGES
Eel Eel
Just 1 1 Just
hatched Leptocephalus hatched
2 months 2 2 2 months
5
5 4
4 5 months 3 3 8 months
Europe
North
3 3
America 10 months 4 4 18 months
6 6
2
1 2
1
4
Africa
1 year 5 5 3 years
Elver
6-10 6 6 8-15
years years
Adult eel
Figure 24.33
Life histories of the European eel, Anguilla anguilla, and American eel, Anguilla rostrata. Migration patterns of European species are shown in red.
Migration patterns of American species are shown in blue. Boxed numbers refer to stages of development. Note that the American eel completes
its larval metamorphosis and sea journey in one year. It requires nearly three years for the European eel to complete its much longer journey.
Salmon runs in the Pacific Northwest have been devastated by a

lethal combination of spawning stream degradation by logging, pol-
lution and, especially, by more than 50 hydroelectric dams, which
obstruct upstream migration of adult salmon and kill downstream
migrants as they pass through the dams’ power-generating turbines.
In addition, the chain of reservoirs behind the dams, which has
converted the Columbia and Snake Rivers into a series of lakes,
increases mortality of young salmon migrating downstream by
slowing their passage to the sea. The result is that the annual run of
wild salmon is today only about 3% of the 10 to 16 million fish that
ascended the rivers 150 years ago. While recovery plans have been
delayed by the power industry, environmental groups argue that
in the long run losing the salmon will be more expensive to the
regional economy than making changes now that will allow salmon
stocks to recover.
Experiments by A. D. Hasler and others have shown that

homing salmon are guided upstream by the characteristic odor
of their parent stream. When salmon finally reach the spawning
beds of their parents (where they themselves were hatched),
they spawn and die. The following spring, newly hatched Figure 24.34
fry transform into smolts before and during the downstream Migrating Pacific sockeye salmon (Oncorhynchus nerka).
hic70049_ch24_514-542.indd 537 7/25/07 1:25:10 PM

migration. At this time they are imprinted (p. 791) with the single spawning. Less than one in a million eggs will survive the
distinctive odor of the stream, which is apparently a mosaic of numerous perils of the ocean to reach reproductive maturity.
compounds released by the characteristic vegetation and soil in Unlike the minute, buoyant, transparent eggs of pelagic
the watershed of the parent stream. They also seem to imprint marine teleosts, those of many near-shore bottom-dwelling
on odors of other streams they pass while migrating downriver (benthic) species are larger, typically yolky, nonbuoyant, and
and use these odors in reverse sequence as a map during the adhesive. Some bury their eggs, many attach them to vegeta-
upriver migration as returning adults. tion, some deposit them in nests, and some even incubate them
How do salmon find their way to the mouth of a coastal in their mouths (Figure 24.36). Many benthic spawners guard
river from the trackless miles of open ocean? Salmon move hun- their eggs. Intruders expecting an easy meal of eggs may be met
dreds of miles away from the coast, much too far to be able with a vivid and often belligerent display by the guard, which is
to detect the odor of their parent stream. Experiments suggest almost always male.
that some migrating fish, like birds, can navigate by orienting to Freshwater fishes almost invariably produce nonbuoyant
the position of the sun. However, migrant salmon can navigate eggs. Those, such as perch, that provide no parental care simply
on cloudy days and at night, indicating that sun navigation, if scatter their myriads of eggs among weeds or along the sedi-
used at all, cannot be a salmon’s only navigational cue. Fish ment. Freshwater fishes that do provide egg care, such as bull-
also (again, like birds) appear able to detect and to navigate to head catfishes and some darters, produce fewer, larger eggs that
the earth’s magnetic field. Finally, fishery biologists concede that enjoy a better chance for survival.
salmon may not require precise navigational abilities at all, but Elaborate preliminaries to mating are the rule for freshwater
instead may use ocean currents, temperature gradients, and food fishes. A female Pacific salmon, for example, performs a ritu-
availability to reach the general coastal area where “their” river is alized mating “dance” with her breeding partner after arriving
located. From this point, they would navigate by their imprinted at the spawning bed in a fast-flowing, gravel-bottomed stream
odor map, making correct turns at each stream junction until (Figure 24.37). She then turns on her side and scoops a nest hole
they reach their natal stream. with her tail. As eggs are laid by the female, they are fertilized
by a male (Figure 24.37). After the female covers the eggs with
gravel, the exhausted fish dies.
Reproduction and Growth Soon after an egg of an oviparous species is laid and fertil-
In a group as diverse as fishes, it is no surprise to find extraordi- ized, it absorbs water and the outer layer hardens. Cleavage fol-
nary variations on the basic theme of sexual reproduction. Most lows, and a blastoderm forms, astride a relatively enormous yolk
fishes favor a simple theme: they are dioecious, with exter- mass. Soon the yolk mass is enclosed by the developing blasto-
nal fertilization and external development of their eggs and derm, which then begins to assume a fishlike shape. Many fish
embryos (oviparity). However, as tropical fish enthusiasts are hatch as larvae, carrying a semitransparent sac of yolk, which
well aware, the ever-popular ovoviviparous guppies and mollies provides their food supply until the mouth and digestive tract
of home aquaria bear their young alive after development in the have developed. The larvae then begin searching for their own
ovarian cavity of the mother (Figure 24.35). As described earlier food. After a period of growth a larva undergoes a metamor-
in this chapter (p. 523), some viviparous sharks develop a kind phosis, especially dramatic in many marine species, including
of placental attachment through which the young are nourished eels (see Figure 24.33). Body shape is refashioned, fin and color
during gestation. patterns change, and the animal becomes a juvenile bearing the
Let us return to the much more common oviparous mode of unmistakable definitive body form of its species.
reproduction. Many marine fishes are extraordinarily profligate
egg producers. Males and females come together in great schools
and release vast numbers of gametes into the water to drift with
currents. Large female cod may release 4 to 6 million eggs at a
Figure 24.36
Male banded jawfish,
Opistognathus
macrognathus, orally
Figure 24.35 brooding its eggs.
Rainbow surfperch, The male retrieves
Hypsurus caryi, the female’s eggs and
giving birth. All incubates the eggs
of the West Coast until they hatch. During
surfperches (family brief periods when the
Embiotocidae) are jawfish is feeding, the
ovoviviparous. eggs are left in the
burrow.
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Asia Alaska
Oceanic distribution
Figure 24.37
Spawning Pacific salmon, Oncorhynchus, and development of eggs and young.
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Classification of Living Fishes

The following Linnaean classification of major fish taxa follows that Subclass Holocephali (holo-sefa-li) (Gr. holos, entire,
of Nelson (2006). The probable relationships of these traditional kephalē, head): chimaeras, ratfishes. Scales absent;
groupings together with the major extinct groups of fishes are four gill slits covered by operculum; jaws with tooth plates;
shown in a cladogram in Figure 24.2. Other schemes of classifica- accessory clasping organ (tentaculum) in males; upper jaw
tion have been proposed. Because of the difficulty of determining fused to cranium. Examples: Chimaera, Hydrolagus; about
relationships among the numerous living and fossil species, we can 33 species, marine.
appreciate why fish classification has undergone, and will continue Class Actinopterygii (akti-nop-te-rijee-i) (Gr. aktis, ray,
to undergo, continuous revision. pteryx, fin, wing): ray-finned fishes. Skeleton ossified;
Phylum Chordata single gill opening covered by operculum; paired fins supported
Subphylum Vertebrata (Craniata) primarily by dermal rays; limb musculature within body; swim
Superclass Myxinomorphi bladder mainly a hydrostatic organ, if present; atrium and
Class Myxini (mik-siny) (Gr. myxa, slime): hagfishes. ventricle not divided; teeth with enameloid covering.
No jaws or paired fins; mouth with four pairs of tentacles; Subclass Cladistia (clə-distē-a) (Gr. cladi, branch): bichirs.
buccal funnel absent; 1 to 16 pairs of external gill openings; Rhombic ganoid scales; lungs; spiracle present; dorsal fin
vertebrae absent; slime glands present. Examples: Myxine, consisting of 5 to 18 finlets. Examples: Polypterus; about 16
Epaptretus; about 70 species, marine. species, freshwater.
Superclass Petromyzontomorphi Subclass Chondrostei (kon-drost¯e-¯i) (Gr. chondros,
Class Petromyzontida (pettr¯o-m¯i-zonti-də) (Gr. petros, cartilage, osteon, bone): paddlefishes, sturgeons.
stone, myzon, sucking): lampreys. No jaws or paired Skeleton primarily cartilage; caudal fin heterocercal; large
fins; mouth surrounded by keratinized teeth but no scutes or tiny ganoid scales present; spiracle usually present;
barbels, buccal funnel present; seven pairs of external gill more fin rays that ray supports. Examples: Polyodon,
openings; vertebrae present only as neural arches. Examples: Acipenser ; 29 species, freshwater and anadromous.
Petromyzon, Ichthyomyzon, Lampetra; 38 species, freshwater Subclass Neopterygii (neeop-te-rijee-i) (Gr. neo, new,
and anadromous. pteryx, fin, wing): gars, bowfin, teleosts. Skeleton
Superclass Gnathostomata (natho-sto-ma-ta) (Gr. gnathos, primarily bone; caudal fin usually homocercal; scales cycloid,
jaw, stoma, mouth). Jaws present; paired appendages present ctenoid, absent, or rarely, ganoid. Fin ray number equal
(secondarily lost in a few forms); three pairs of semicircular canals; to their supports in dorsal and anal fins. Examples: Amia,
notochord partly or completely replaced by centra. Lepisosteus, Anguilla, Oncorhynchus, Perca; about 27,000
Class Chondrichthyes (kon-drikthee-eez) (Gr. chondros, species, nearly all aquatic habitats.
cartilage ichthys, fish): cartilaginous fishes. Cartilaginous Class Sarcopterygii (sar-cop-te-rijee-i) (Gr. sarkos, flesh,
skeleton; teeth not fused to jaws and usually replaced; no swim pteryx, fin, wing): lobe-finned fishes. Skeleton ossified;
bladder; intestine with spiral valve; claspers present in males. single gill opening covered by operculum; paired fins with
Subclass Elasmobranchii (e-lazmo-brankeei) (Gr. sturdy internal skeleton and musculature within appendage;
elasmos, plated, branchia, gills): sharks, skates, and diphycercal tail; intestine with spiral valve; usually with
rays. Placoid scales or derivatives (scutes and spines) lungs; atrium and ventricle at least partly divided; teeth
usually present; five to seven gill arches and gill slits with enamel covering. Examples: Latimeria (coelacanths);
in separate clefts along pharynx; upper jaw not fused Neoceratodus, Lepidosiren, Protopterus (lungfishes); 8 species,
to cranium. Examples: Squalus, Raja, Charcarodon, marine and freshwater. Not monophyletic unless tetrapods
Sphyrna; about 937 species, mostly marine. are included.
Figure 24.38
Growth is temperature dependent. Consequently, fish Scale growth. Fish scales
disclose seasonal changes
living in temperate regions grow rapidly in summer when
in growth rate. Growth
temperatures are high and food is abundant but nearly stop is interrupted during
growing in winter. Annual rings in the scales, otoliths, and other winter, producing year
bony parts reflect this seasonal growth (Figure 24.38), a dis- marks (annuli). Each year’s
tinctive record of convenience to fishery biologists who wish increment in scale growth
is a ratio to the annual
to determine a fish’s age. Unlike birds and mammals, which
increase in body length.
stop growing after reaching adult size, most fishes after attain- Otoliths (ear stones) and
ing reproductive maturity continue to grow for as long as certain bones can also be
they live. This may be a selective advantage, since the larger used in some species to
the fish, the more gametes it produces and the greater its determine age and growth
rate.
contribution to future generations.
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SUMMARY
Fishes are poikilothermic, gill-breathing aquatic vertebrates with fins by undulatory contractions of the body muscles, which generate
for appendages. They include the oldest vertebrate groups, having thrust (propulsive force) and lateral force. Eel-like fishes oscillate the
originated from an unknown chordate ancestor in the Cambrian whole body, but in more rapid swimmers undulations are limited to
period or possibly earlier. Five classes of living fishes are recog- the caudal region or caudal fin alone.
nized. The jawless hagfishes (class Myxini) and lampreys (class Most pelagic bony fishes achieve neutral buoyancy in water
Petromyzontida), have an eel-like body form without paired fins, a using a gas-filled swim bladder, the most effective gas-secreting
cartilaginous skeleton, a notochord that persists throughout life, and device known in the animal kingdom. Sensitivity to sounds may
a disclike mouth adapted for sucking or biting. All other vertebrates be enhanced by Weberian ossicles that transmit sounds from the
have jaws, a major development in vertebrate evolution. Members swim bladder to the inner ear. Gills of fi shes, having effi cient
of class Chondrichthyes (sharks, rays, skates, and chimaeras) have countercurrent flow between water and blood, facilitate high rates
a cartilaginous skeleton (a degenerative feature), paired fins, excel- of oxygen exchange. All fishes show well-developed osmotic and
lent sensory organs, and an active, characteristically predaceous ionic regulation, achieved principally by the kidneys and gills.
habit. Bony fishes may be divided into two classes. Lobe-finned With the exception of agnathans, all fishes have jaws that are
fishes of class Sarcopterygii, represented today by lungfishes and variously modified for carnivorous, herbivorous, planktivorous, and
coelacanths, form a paraphyletic group if tetrapods are excluded, as detritivorous feeding modes.
done in traditional classification. Terrestrial vertebrates arose from Many fishes are migratory, and some, such as catadromous
one lineage within this group. The second is ray-finned fishes (class freshwater eels and anadromous salmon, make remarkable migra-
Actinopterygii), a huge and diverse modern assemblage containing tions of great length and precision. Fishes reveal an extraordinary
nearly all familiar freshwater and marine fishes. Modifications of the range of sexual reproductive strategies. Most fishes are oviparous,
skeletal and muscular systems in this group increased locomotion but ovoviviparous and viviparous fishes are not uncommon. Repro-
and feeding efficiency. ductive investment may be in large numbers of eggs with low sur-
Modern bony fishes (teleosts) have radiated to form approxi- vival (many marine fishes) or in fewer eggs with greater parental
mately 27,000 species that reveal an enormous diversity of adapta- care for better survival (freshwater fishes).
tions, body form, behavior, and habitat preference. Most fishes swim
REVIEW QUESTIONS
1. Provide a brief description of the fishes citing characteristics following synapomorphies to the diagram: claspers, cranium,
that would distinguish them from all other animals. endochondral bone, fleshy fins, jaws, vertebrae.
2. What characteristics distinguish hagfishes and lampreys from 10. List four characteristics of teleosts that contributed to their
all other fishes? How do they differ in morphology from each incredible diversity and abundance.
other? 11. Only eight species of lobe-finned fishes are alive today,
3. Describe feeding behavior in hagfishes and lampreys. How do remnants of a group that flourished in the Devonian period of
they differ? the Paleozoic. What morphological characteristics distinguish
4. Describe the life cycle of sea lampreys, Petromyzon marinus, lobe-finned fishes? What is the literal meaning of Sarcopterygii,
and the history of their invasion of the Great Lakes. the class to which lobe-finned fishes belong?
5. In what ways are sharks well equipped for a predatory life 12. Give the geographical locations of the three surviving genera
habit? of lungfishes and explain how they differ in their ability to
6. What function does the lateral-line system serve? Where are survive out of water. Which of the three is the most similar to
receptors located? the earliest, fossil lungfishes?
7. Explain how bony fishes differ from sharks and rays in the 13. Describe discovery of living coelacanths. What is the
following systems or features: skeleton, scales, buoyancy, evolutionary significance of the group to which they belong?
respiration, and reproduction. 14. Compare the swimming movements of eels with those of
8. Match ray-finned fishes in the right column with the group to trout, and explain why the latter are more efficient for rapid
which each belongs in the left column: locomotion.
_____ Chondrosteans a. Perch 15. Sharks and bony fishes approach or achieve neutral buoyancy
_____ Nonteleost b. Sturgeon in different ways. Describe the methods evolved in each
neopterygians c. Gar group. Why must a teleost fish adjust the gas volume in its
_____ Teleosts d. Salmon swim bladder when it swims upward or downward? How is
e. Paddlefish gas volume adjusted?
f. Bowfin 16. What is meant by “countercurrent flow” as it applies to fish
9. Make a cladogram that includes the following groups gills?
of fishes: chondrosteans, elasmobranchs, hagfishes, 17. How do Weberian ossicles increase a fish’s sensitivity to
holocephalans, lampreys, lungfishes, teleosts. Add the sounds?

Fishes

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C H A P T E R

Placoderms Chondrichthyes Holocephalans

Cambrian Ordovician Silurian Devonian Carboniferous Permian

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Elasmobranchii Actinopterygii Sarcopterygii

Acanthodii† Limbs used for

Part of second visceral arch modified as

Mineralized tissue in dermis

2 or more pairs semicircular canals, vertebrae, extrinsic eye muscles

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Nostril Olfactory Brain Spinal Notochord Pharynx

C Teeth on tongue Tongue D

Class Petromyzontida: Lampreys

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There are 20 species of lampreys in North America. About half of

Migration Parasitic stage in lakes

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Characteristics of Class Petromyzontida Pharynx

Tongue protruded for

Subclass Elasmobranchii: Sharks,

J Tiger shark K Thresher shark

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Lateral line Pelvic fin

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Lateral line Ampullae

Opening to surface Lateral-line canal

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Figure 24.11 Celiac Kidney First

Stingrays have a slender and whiplike tail armed with one or

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term as a valid taxon. Rather, it is used as a term of convenience

First dorsal fin

Vertebra Lateral line

The major clade of neopterygians are teleosts (Gr. teleos,

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Basal plate Pulp cavity

Ganoid scales Cycloid scales Ctenoid scales

contractions on the tail, permitting greater speed. The dorsal fin

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Class Sarcopterygii: Lobe-Finned Fishes

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Australian lungfish African lungfishes South American lungfish

specimens was successful off the coast of the Comoro Islands.

waves of contraction moving backward along its body by alter-

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able to move through water while creating almost

Neutral Buoyancy and the

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Hearing and Weberian Ossicles

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Gill filaments Gill raker

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Large Active tubular

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Marine bony fishes are hypoosmotic regulators that

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Salmon runs in the Pacific Northwest have been devastated by a

Experiments by A. D. Hasler and others have shown that

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