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About Fishes

Fishes are by far the largest group of vertebrate ani

mals. T heir numbers may approach
cies (Nelson,

28,000 extant spe

1984), well over half of all vertebrates,

and thousands of other fish species lived in past geo

ment. Some families are virtually restricted to fresh wa

ter (e.g., percids) while others with many freshwater
representatives may show some tolerance for brackish
environments or have a few members occurring pri

logic times. Over half the world's species of fishes live

marily in marine habitats (e.g, cyprinodontiform fami

in marine environments; these are most diverse in the

lies). Still other, primarily marine, groups have a few

seas of the tropical Indo-Pacific region and in the Carib

members which have invaded freshwater environments

bean. Freshwater fishes are most diverse in South

(e.g., gobies, atheriniform fishes). T hese latter are best

America, which may harbor

5,000 species, and in

represented in regions where strictly freshwater groups

southeastern Asia. North America, including Mexico,

are absent

has a moderately diverse native freshwater fauna of

Central America, and oceanic islands. Groups that are


1,000 species.


less common, such as Australia, lower

able to move freely between waters of varied salinity

However, it should be pointed out that "fishes" is a

collective term for actually three very different groups

are termed euryhaline. Examples are the cyprinodon

tiform fishes (topminnows, mollies, and related

of currently living vertebrates and thus is not a "natu

groups). Many fishes are migratory, especially in re

ral" group, as, for instance, the mammals (class Mam

sponse to reproduction, and some are diadromous,

malia) are presumed to be. Included in fishes are the

spending portions of their lives in both marine and

class (or superclass) Agnatha, the primitive jawless ver

freshwater environments. Fishes that live most of their

tebrates represented today by the lampreys and, under

lives in the ocean but spawn in rivers (e.g., salmon,

older classifications, hagfishes; the class

herrings) are termed anadromous; those that do the op

Chondrichthyes, which includes the sharks, chimaeras,

posite (e.g., freshwater eels) are catadromous fishes.

skates, and rays, characterized by a cartilagenous rather

than bony skeleton; and the class Osteichthyes, the
bony fishes. Osteichthyes alone contains over


Most fishes spend much of their life in one or two

basic habitat types, such as rivers or smaller streams
(lotic habitats), springs, lakes, and swamps (lentic), es

species, making it the largest vertebrate class. All tetra

tuararies, coral reef, oceanic, and so on. A few are so

pod vertebrate groups are believed to have evolved from

generalized in habitat that they may be encountered in

the bony fishes, though the exact fish lineage (Jung

several habitats within freshwater or marine realms.

fishes, lobefins, or other) from which they evolved has

been in dispute. To put this into perspective, "higher"

For purposes of general reference, roughly based on

dimensions, stream habitats include very large rivers a

vertebrates, including humans, are more closely related

half kilometer or more in width, such as the Mississippi

to bony fishes than these fishes are to some other "fish"

or lower Tennessee, large rivers averaging

groups, such as sharks.

in width (e.g., lower Cumberland, upper Tennessee),

All fish groups are characterized by gills, which evo

lutionarily gave rise to various structures, including the

medium rivers

200-400 m

50-200 m width, (e.g., lower Duck or

50 m width (e.g., Little,

Clinch), small rivers under

jaws of chondrichthyans and bony fishes and the ear

Buffalo), large and small creeks, and spring runs.

bones of tetrapods. Chondrichthyans and bony fishes

Creeks and small rivers may be montane (high gradient,

share paired fins, which gave rise to the limbs of tetra

fast-flowing, predominately rocky substrate), upland

pods. These adaptations and many others allow fishes to

(moderate gradient and flow, substrate variable but gen

respire, move, feed, excrete, reproduce, and otherwise

erally v-.ith extensive rocky areas), or lowland (low gra

function in their watery environment.

dient, sluggish current, predominantly depositional

substrates such as sand and silt) in character; larger
streams may be upland or lowland in character. All of

Habitats and Lifestyles

these basic categories and subcategories of stream types

have characteristic fish faunas associated with them

Over half of the species of fishes, including many

which are further modified in composition by prevailing

whole families, are restricted to the marine environ-

habitat types associated with regional geology (see gen-



eral descriptions in previous section, Waters and Geol

ogy of Tennessee). Because fishes have evolved such
close associations with these habitat types, many spe
cies show great fidelity to physiographic provinces.
This fidelity can promote fragmentation and isolation of
populations over time as the surface geology changes,
and it probably has resulted in the evolution of several
new species.
Natural lentic habitats consist of several kinds of
lakes (lacustrine habitats), swamps, and backwaters of
larger streams. Natural lakes are created in a variety of
ways, including glacial modification of the land surface
(as in much of northern North America), subsidence
due to tectonic activity (e.g., Reelfoot Lake) or dissolu
tion of underlying strata (e.g., karst lakes of Florida),
changing river courses (oxbow lakes), or natural dam
ming of streams by landslides or beavers. Humans have
added a large contingent of unnatural lentic environ
ments through the impoundment of streams in which
some lentic fish species have flourished while stream
fishes largely disappeared.
Swamps or marshes are characterized by expanses of
shallow, standing water which may fluctuate seasonally
with rainfall. Permanent swamp habitats usually have
submergent or emergent vegetation and may be forested
(e.g., cypress, tupelo). Waters may be clear, stained
brown by tannic-acid, or seasonally turbid (muddy). A
number of fish species are strictly or often associated
with swampy environments.
Caves constitute an additional unique habitat type. A
few species of fish lead a strictly subterranean (hypo
gean or trogloditic) existence. Other species
(troglophilic) are often associated with caves but may
lead an epigean (surface) existence part of the time. Hy
pogean fishes, which are typically blind and devoid of
pigment, have evolved several specializations to cope
with their lightless and low-nutrient environment, as
discussed in the section herein on amblyopsid fishes .
Beyond general habitat preferences, fishes demon
strate a variety of basic behaviors and physical adapta
tions. Some fishes are rather sedentary, spending much
of their time near a favored haunt or moving about a
small area. Such fishes tend to be solitary or occur in
small, loose aggregations, and some may be constantly
or seasonally territorial . Other fishes rove freely over
large areas and are constantly on the move. These tend
to be schooling fishes and may frequent the midwater
(pelagic) realm or cruise very near the surface or bot
tom. Many midwater species are filter feeders (e.g.,
shad, paddlefish), straining plankton from the water
with special gill rakers, while others actively feed on
midwater organisms or dash to the surface to take fallen


The Fishes

of Tennessee

prey (e.g., many minnows). Still others, such as striped

bass, are roving predators feeding on other midwater
fishes. Surface-oriented fishes (e.g., topminnows, some
silversides) cruise just beneath the surface, feeding on
fallen or emergent organisms; such fishes usually have
an upturned mouth and flattened dorsal profile to facili
tate this life-style. Many fishes are bottom-oriented
(benthic) in behavior, swimming just above the bottom
(epibenthic; e.g., suckers), resting upon it (e.g., darters
and scuJpins), or even burrowing into it. Some of these
species have a downwardly oriented mouth to facilitate
feeding. Those benthic species that frequent swifter wa
ters may have several hydrodynamic adaptations, in
cluding a depressed or streamlined body profile,
modified fins, and reduction of the swimbladder to
lessen buoyancy. Benthic and epibenthic species are
generally very closely associated with a specific sub
strate type (gravel, sand, small or large rocks, vegeta
tion, detritus) and current regime. Though none occur
in North America, a few fishes (e.g., mudskippers,
climbing perches) have remarkably evolved adaptations
permitting them to spend short periods of time on land!
Because they feed on a variety of organisms and, in
tum, are preyed upon by a diversity of predators, in
cluding terrestrial ones, fishes are vital links in nature's
food web, especially between the aquatic and terrestrial
environments. Fishes of different kinds or life-stages
feed on everything imaginable-including microscopic
organisms, such as diatoms and bacteria; plankton;
plants; macroinvertebrates (e.g., insects, crustaceans,
mollusks); other fishes of all sizes; and other verte
brates, both aquatic and terrestrial. In tum, they are fed
upon by everything from large aquatic insects to other
fishes and reptiles and a variety of birds and mammals,
including humans, for whom they have provided one of
the most important protein sources for thousands of
Like nearly every living organism, fishes are host to
a variety of parasites, both external and internal, includ
ing: fungi; protozoans; acanthocephalan, cestode, trem
atode, and nematode worms; and various arthropods,
such as mites, copepods, and isopods (see Hoffman,
1967). Perhaps among the most obvious to collectors of
freshwater species are the immature stages (metacer
cariae) of certain trematode worms which form con
spicuous black cysts on the skin and may be extremely
abundant on fishes in some populations. Also con
spicuous when present are such external parasites as
leeches, "anchor worms" (females of certain copepods
which attach to the skin or fins), and "fish lice"
(copepods). Copepods and large isopods may some
times be found living in the gill cavity of fishes. When

dissecting a fish , it is not uncommon to find worms

(trematodes , nematodes , cestodes) living in the gut or
freely in the body cavity, or to find immature stages en
cysted in the flesh . Perhaps one of the most interesting
parasites is the larval stage (glochidia) of freshwater
mussels (Unionidae) , which are discharged by the fe
male mussel on the gills or fins of passing fi shes where
they undergo early development; subsequently these
drop to the stream bottom to begin their sedentary life
style after having had their dispersal facilitated by the
fish host . Some mussels have actually evolved color
patterns to lure fishes in proximity to facilitate larval in
festation . Though sometimes unsightly, when in natural
equ il ibrium, fish parasites seldom have an impact on the
health of their hosts .

Anatomy and Function

In the following sections we attempt to acquaint readers

with the basic physical features of fishes and how they
function to facilitate their existence in the aquatic envi
ronment. Many terms will be introduced which will be
of use to those who wish to identify fishes through the

keys and descriptions in this book . Much of the infor

mation presented here , plus much additional detailed in
forn1ation , is inferable from the general works of Lagler
et al . (1977), Bond (1979) , and Moyle and Cech (1982,
1988) , and from speci al ized works such as those of Al
exander (1967) , Hoar and Randall (1969-1984), H arder
(1975) , and Caillet et al . (1986).

Head, Body, Fins, and Locomotion. The body form s

and fin configurations and positioning in fi shes consti
tute an almost endless array; some of the more basic
forms are illustrated in Figure 10 . In lateral profile , ba
sic body shapes range from hori zontally elongate , or
extremely attenuate , to deep-bodied (vertically exagger
ated) ; in cross section , elongate fishes may be terete or
fusiform (cylindrical) (e . g . , chub minnow s , gar, mack
erel ) , somewhat laterally compressed (parallel-sided)
(e . g . , striped bass), or dorsally depressed (fl attened
from above; e . g . , sculpins). Deep-bodied fishes are all
laterally compressed to varied degrees; dorsally de
pressed fishes range from relatively elongate (e . g . ,
sculpins) to extremely broad ( e . g . , skates and rays) in
dorsal view.

Figure 10. Some basic body forms of fishes: top, terete (chub minnow); middle, laterally compressed (sunfish); bottom, dorsally
depressed (sculpin).



spinous dor sal fin




dorsal spines

gill cover
\o percleI

gill juncture

isthm us

anal spines



pelvic fin

anal fin

/th orac ic positionl

adipose fin


axillary process

pelvic fin /abdominal position\

Figure 11. Body regions and basic anatomical features of a fish: a) a spiny-rayed fish (perch) with anteriorly (thoracic) positioned
pelvic fins; b) a darter (small percid) showing details of urogenital area; c) a soft-rayed fish (trout) with abdominally positioned
pelvic fins and other fin features shown.

The body of a bony fish is divided into three main re

gions: the head, trunk, and caudal (Fig. l l a). The head

juncture of the gill flaps generally serves to demarcate

the rear of the head. The trunk, which contains the ab

includes that region from the tip of the snout to the pos

dominal cavity, lies between the head and caudal re

terior extremity of the gill cover. Dorsally, it is gener

gion. Dorsally, the region of the trunk ly ing between

ally delimited by the area where dorsal body muscu

the occiput and the dorsal fin is termed the nape. Ven

lature attaches to the rear of the skull (the occiput)

trally, the trunk subregions are the narrow isthmus (in

and is generally discemable as a transverse line an

terposed between the gill flaps), the breast (anterior to

terior to a hump in the dorsal profile. Ventrally, the

the pelvic fins), and belly. In most fishes the body is


The Fishes

of Tennessee

distinctl y constricted just behind the anal fin to form the

caudal peduncle. In some fishes the caudal region , the
beginning of which technically corresponds internally to
the first vertebra with a ventral spine (haemal spine)
rather than ribs (see Fig . 17) , simply tapers from some
where near midbody without a distinct peduncle being
formed ; this i s espec i ally true i n groups in which the
anal fin is reduced or lacking , as in eel s .
In lampreys (Fig . 12a) , the head is considered t o ex
tend posterior to and include the body segment bearing
the seventh (last) gill aperture . The trunk extends from

while more inferior mouths may be suctorial or mod

ified for scraping algae and associated organisms from
the bo ttom (e . g . , sturgeon , suckers). The jaws may be
protractile (extendable) or nonproctractil e ; in the l atter
case the premaxilla is either broadly or n arrowly con
nected to the s nout by a fleshy median frenum (Fi g . 14) .
In some fi s h groups (e . g . , gars , needlefishes) the jaw
bones may be greatly modified into beaklike or other
structures . In other groups (e . g . , catfishes , some min
nows), barbels, which are usually well endowed with
taste buds , may be present on the maxillary and mandi
bular areas (Fig . 15) .
The presence , locations , and types of teeth also vary
greatly among fishes . They may be present or absent on
any of the jaw or interior mouth bones (Fig . 13b), and a
few fi sh groups (e . g . , minnows) lack teeth in the mouth
region altogether. Various tooth types are shown in Fig
ure 16 .
In lampreys , jaws are lacking; instead , the mouth
consists of a b uccal funnel, a suction device , which has
variously developed toothlike structures (see termi nol
ogy in Fig . 1 2b). In parasitic species , the buccal funnel
and tooth structures function in attaching and holding to
slippery host fishes and to rasp a hole from which blood

behind the head to the muscle segment (myomere) bear

ing the anus or urogenital opening , and the remainder is
considered the c audal region .
The main external features of the head of bony fishes
and bones of the interior of the mouth are illustrated in
Figures 13a and b. Mouth shapes and positions vary
greatly among fi shes . In many fishes the mouth is ter
minal, located at the anterior extremity of the head , but
in others it may be slightly overhung by the snout (sub
terminal) , far beneath the snout (inferior), or upturned
(superior) for surface feeding. In general , terminally lo
cated mouths are utilized for biting and seizing (e . g . ,
sunfish), or water intake for filter feeding (e . g . , shad) ,


caudal region




anterior teeth


diphycercal caudal fin

urogenital papilla & opening

supraoral lamina

lateral teeth
ct==== circumoral tee

marginal tooth row ----tc

transverse lingual lamina

infra oral lamina

Figure 12. Lamprey anatomy: a) body regions and external features; b) features of oral region.









pectoral girdle






pre opercular








meta pterygoid





palat ines

Figure 13, Bones of the head region of a fish: a) jaw bones and superficial skull bones (redrawn and modified from Lagler et a!.,



b) bones of the roof of the mouth region (ventral view),

The Fishes

of Tennessee

The superficial facial bones and sensory canal system

of a bony fish are shown in Figure 13 and 26. The size
of the eye, or orbit, varies greatly among fishes and
varies with growth (allometry) in an individual, with
the eyes of larvae and juveniles usually being propor
tionately larger than those of adults . The infraorbital (or
suborbital) bones may be poorly developed or partially
absent in some fishes with corresponding alterations of
the canal system borne by those bone s . The sensory
canal system (cephalic latera lis system; see Fig . 26 )
varies in development among fishes, and some canal
portions may be elaborated, interrupted, or absent, or
even cavernous, as in fi shes of the drum family. The
bones (opercle, preopercle, and others) associated with
the gill flap region vary widely in shape and may or
may not bear spines or other features .
The body of a fish is composed of sequential muscle
segments termed myomeres (Fig . Ila) which, in small
scaled or scaleless species, may be vi sible through the
skin or as creases on preserved specimens . In lampreys
(Fig . 12a), they are usually quite evident, and their
numbers are important in identifi cation of species .
In many bony fish groups a lateral-line canal (see
Sense Organs section , below) is evident along the side
of the body near the midline (Fi g . II a) extending from
the cephalic system on the head near the top of the gill
opening to the base of, or onto, the caudal fi n . It may

up p e r lip (p r emaxillaries)

Figure 14. Dorsal view of head of fish with non-protractile

upper jaw showing connection of frenum.

can be draw n . In nonparasitic lampreys , this organ fa

cilitates attachment to the substrate. It should be noted
that , unl ike bony fishes , the nostril in lampreys is a sin
gle median opening before the eye s . On the sides of the
head are seven circular openings which s erve as the gill
apertures . The lateral line pores of a lamprey's head are
mainly microscopic and may not be visible as illus
trated .

Figure 15. Barbels of fishes: left, a minnow (carp); center, a catfish; right, a sturgeon.

Figure 16. Some basic types of dentition in fishes: a) canifonn; b) villifonn or cardifonn (ventral view of upper jaw of catfish); c)
anterior incisifonn and posterior molarifonn teeth of lower jaw of a marine porgy (Sparidae).



be borne in a row of scales, or in the dermis of scaleless

spec ies, and may be incomplete , interrupted, or absent
in some groups and is even variable within spec ies in
some groups . In some fishes (e . g ., cavefi shes) the lat
eral line has vertical elaboration s .
Also externally visible on the body are the urogenital
opening and anus (Fig . lib). In most groups these
openings are just before the anal fin, but in two families
(pirateperch, cavefishes) these m igrate forward during
development to a jugular position just behind the is
thmus . The urogenital opening is posterior to the anus
and serves both the kidneys and gon ads; it is well sepa
rated from the anus or even produced into a tube c alled
a genital papilla. The term genital papilla is used when
the opening is on an elevated mound or tube .
The most prominent external features on the body of
most fishes are the fins . Individual and co llective termi
nology for fin s and their support structures are sho wn in
Figures lla,c, and 17. Fins vary greatly in shape and
size among and within fishes, and some groups lack
certain fin s . Dorsal fin s are present in most fishes; soft
rayed fishes (e . g ., trout, minnows) typically have a sin
gle dorsal fin (two or three in cods and relatives) while
spiny-rayed fishes (e . g ., perch) may have two separate
fins or a single continuous fin. An al fin s are single in
most fishes. The dorsal and anal fins are supported prin
cipally by the bladelike pterygiophores, inserted medi
ally between the lateral body muscul ature, and the fin
rays . Soft rays consist of bilaterally paired, segmented
structures which, except for the anterior one or few, are

usually branched one or more times tow ard the extremi

ties . "True" spines, such as those of perc iform fi s hes,
are solid, unpaired, and un segmented structures . The
"spines" of catfi shes, some m innows (e . g . , carp), and a
few other " lower" bony fish groups, which are gener
ally single, differ in that they develop from hyperostosis
(profuse bone tissue deposition) and fusion of seg
mented rays .
The caudal fin, or tail fin, is well developed in the
majority of fish groups but may be reduced or absent in
some (e . g ., some eels). The three basic types of caudal
fin are s hown in Figures 17 and 18. Lampreys have a
diphycercal caudal fin in which the rays radiate from
and surround the tip of the notochord . Certain primitive
groups, including the sturgeons, paddlefishes, gars,
bowfin, and sharks, have a heterocercal caudal fin in
which the upper lobe emanates from the upturned termi
nal vertebrae and the lower from the ventral aspects of
those bones . Typical of most bony fi s hes is the homo
cereal, u su ally relatively symetrical, confi guration ema
nating entirely from bladelike modifications of the
upturned terminal vertebra termed the hypural plate
complex (Fig . 17 ) . Homocercal caudal fins may have a
variety of shapes, including rounded, truncate, forked,
crescentic, and others .
Numerous families of lower teleosts (e . g ., smelt,
trout, catfish) have an additional median fin, the ad
ipose fi n. It is a fleshy fin lacking rays or spines and i s
typically located between t h e dorsal and caudal fins .

vertebral col umn

predorsal bones

homoce rcal ca udal fin

pectoral girdle
hypural complex
pelvic girdle

haemal spines
ple ural r i b s

Figure 17. Axial skeleton, fin support structures, and caudal fin type of a bony fish (centrarchid sunfish).


The Fishes of Tennessee

ascending, and descending. Some species with very

long-based dorsal fins, such as bowfins, achieve some
propulsion by wavelike undulations of this fin.

Figure 18. Primitive caudal fin types: a) diphycercal; b)

The paired fins (Fig. 1 1 a), the pectorals and the pel
vics, are bilateral appendages which vary considerably
in placement among fish groups. Pectoral fins are al
ways just behind the head but may be inserted relatively
low (e.g., suckers) or high (e.g., silversides) on the
sides of the body. The pelvic fins (also called ventrals)
may be situated near midbody (abdominal; e.g., trout,
minnows), which is thought to be the evolutionarily
more primitive positioning, or they may be situated an
teriorly (thoracic or jugular) beneath or in advance of
the pectoral fins (e.g., perch, sunfish), as is characteris
tic of "higher" fish groups. The pectoral and pelvic fins
are supported internally (Fig. 17) by complexes of large
bones (girdles) which are not directly associated with
the vertebral column. In most groups only soft ray s are
present in the pectoral fin, but in some (e.g., catfishes)
the first rays may be fused into a spine. In many spiny
rayed fishes the pelvic fin has a single spine. The axil
lary processes shown in Figure 1 1 are characteristic of
only a few fish groups, such as salmonids and herrings,
and a small axillary process occurs in many cyprinids.
Fishes move through the water by using both body
movements and the fins. Primarily, forward motion is
attained by side-to-side flexure of the body, caused by
alternating contractions of the myomeres, culminating
in thrust from the caudal fin. Fins are used primarily for
short bursts, stabilization, and stopping or reversing.
Median fins serve as stabilizers and, when curled to the
side, as brakes. Paired fins are for short maneuvering,

Gills and Respiration. In bony fishes the gills consist of

five arches, each composed of several bones, suspended
posteriorly beneath the skull (Fig. 19). T hese pharyn
geal arches and their associated structures are collec
tively sometimes called the branchial basket. Each arch
bears numerous filaments posteriorly that are richly sup
plied with arterial blood. Anteriorly, the major bones
(limbs) of the arch usually have inwardly directed gill
rakers of varied numbers and shapes which serve to
strain food or other matter passing over the gills. In
many fish, especially perciforms, teeth are present on
various pharyngeal bones; these may be patches of
small teeth for grasping or large molariform (rounded,
molarlike) teeth for grinding prey, such as mollusks. In
minnows and suckers, in place of the gill rakers, the
fifth arch has developed large pharyngeal teeth (Fig.
20) which li just ahead of the esophagus and serve to
tear or grind food, and which vary in shape from sharp
pointed to hooked to molariform.
The heart of a bony fish is located just posterior to
the gills in the ventral portion of the body cavity (see
Fig. 23), and blood is pumped directly into the gill
arches. Respiration is achieved by pumping water over
the gills with alternating expansions of the mouth cavity
and opening of the gill flaps or by passage of water
through the gill chamber while swimming. Gaseous ex
change (oxygen uptake, carbon dioxide release) occurs
in the gill filaments, and oxy genated blood is pumped
to the rest of the body. Gill filaments are also the site of
chloride cells which serve to maintain salt balance in
fishes. In many fishes, additional filamentous organs,
the pseudobranchs, are located on the lining of the gill
chamber in the preopercle area (Fig. 19). These may
provide oxygen to the eyes and possibly augment gas
production for the swimbladder.
In lampreys, the gills consist simply of seven paired
pouches on either side of the alimentary tract which
open medially to this tract and have small external vents
visible along the sides of the head (Fig. 12a). These
pouches are lined with filaments through which gas ex
change takes place when water is pumped over them by
contractions and expansions of the pouches. Gills of
sharks, rays, and their relatives are more similar to
those of bony fishes, consisting of arches, but the
chamber has multiple slits opening to the outside rather
than a single gill flap, and a spiracle (reduced anterior
gill slit) may be present dorsally for intake of water.



rt. pharyngeal arches 1

pseudobra nch (rt. side)

1 st left arch
gill ra kers

lo w er limb

mod ified 5th arc h w ith tooth plates

Figure 1 9 . G ill chamber region o f a f i s h showing gill structures and pseudobranch location.

Skin and Scales. The bodies of fishes are covered with

a somewhat permeable skin , and most have bony scales
or derivitive structures on much of the skin surface , but
a few groups (e . g . , freshwater sculpins and several cat
fish families) lack scale s . The skin is endowed with mu
cous cells which produce the characteristic slime of
fi shes ; this secretion may function as a sealant agai nst
infections and to reduce friction in swimming. Also
present in the skin are the organs of color, the chroma
tophores and iridocytes. The former are capable of con
traction and expansion to change coloration and impart
true color through the possession of different pigments
(e . g . , melanophores have black pigment, erythrophores
red , and so on). Iridocytes reflect external l ight sources
and result in silvery reflections and iridescent effects .


The Fishes of Tennessee

The basic scale types (Fig . 2 1a-d) are placoid

(sharks and relatives ) , ganoid (gars , sturgeons), and
cycloid, the more familiar type found on most fishes .
The first two types are thick and enamel-like and lack
the circular bony ridges characteristic of cycloid scales .
The parts of a cycloid scale are shown in Figure 2 1c .
The concentric growth rings and annual growth checks
(annuli) are important in age studies of fishes . In many
fish groups (primarily spiny-rayed) cycloid scales are
modified (Fig . 2 I d), having tiny spines on the posterior
surface either as an integral part of the scale (few ma
rine groups) or attached at the base by platelets
(ctenoi d ) , giving the fishes a rough texture (e . g . , perch ,
sunfish) . Some specialized derivatives of scales are the
external armorl ike pl atings of seahorses and pipefishe s ,



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. ..
.. . ' . . .. . . .. . . ".- - .

' -"

-.- - -







Figure 2 1 . Basic fish scale types: a) placoid ; b) ganoid; c)
cycloid; d) ctenoid (drawn, in part, by P. Yarrington).

Figure 2 0 . Modified fifth pharyngeal arches of fishes: a) with

partly hooked teeth for ripping and tearing (minnow); b) with
comb-like teeth (sucker); c) wi th molarifonn teeth (redear
sunfish, a snail-eater); d) heavily modified with molariform
teeth for crushing mollusks (freshwater drum).



nous photophores of deep-sea fishes which may func

tion in species recognition and illumination of prey.
In several fish groups , breeding tuber cles, or "pearl
organs , " develop on the head , body, or fins (Fig .


during the spawning season (see Wiley and Collette ,

1 970). T hese are horny protuberances which are se

creted in response to seasonal hormonal changes , pri
marily in males , and presumably function in enhanced
contact during spawning . T hese tubercles slough off af
ter the spawning season .

Figure 2 2 . Examples of fish breeding tubercles: a) head and

body tubercles of a stoneroller m i nnow (genus Campostoma);
b) tuberc les on pectoral fin rays of a cyprinid in uniserial (one
per fin-ray segment) configuration.

Sense Or gans and Senses. T he sensory systems of

fishes are necessarily quite different from those of ter
restrial vertebrates . For instance , in bony fishes, the
cephalic lateralis system is instrumental in detection of
sound or vibrations . T he canals of the head and lateral

belly scutes of herring , and the scalpel-like spine on the

line are equipped with cells called neuromasts which

caudal peduncle of the marine surgeonfishes .

have hairlike cilia oriented in such a way as to detect di

Scales vary tremendously in size between and within

fish groups and may be firmly attached or easily lost

rectionality of vibrations . T his information is transmit

ted directly to the brain via nerves which lie in

(deciduous). The coverage of scales (squamation) also

proximity to the canals . T hus the "ears" of fishes are in

varies greatly among fishes . When present , scales are

part very different from those of terrestrial vertebrates .

usually distributed on much of the sides and back of the

T he function of this system and its relationships to be

body but may be present or absent on the belly, breast ,

havior were treated extensively by Disler

and nape (dorsal surface from occiput to dorsal fin ori

( 1 960) .

T here is also an inner ear, more analagous to those of

gin). On the head , they may be completely absent , or

other vertebrates , but which , instead of containing ear

present or absent on the opercles , cheek area , and top

bones or ossicles, has three calcareous "ear stones" or

of the head . Only a few groups have the head almost

otoliths. One of these (the lapillus) functions in main

completely scaled .

taining body orientation and the remaining two (as

A variety of other features may be found on the skin

ter icus, sagitta) in sound reception . In several fish

of fishes . Many species have external taste buds , usu

groups , features have evolved to enhance transmission

ally visible only microscopically as tiny bumps , located

of vibrations to the inner ear area . In minnows , suckers ,

on the skin surface , particularly on the ventral portion

catfishes , and relatives (ostar iophysan or otophysan

of the head , and some even on the body. Small fleshy

fishes) , the anterior vertebrae are modified to form a

appendages , such as lappets or papillae , are present on

chain of bones , the Weber ian ossicles , interlinking the

the skin of some fishes (e . g . , sculpins) . Perhaps some

swimbladder, which is very sensitive to vibrations , to

of the most remarkable organs of the skin are the lumi-

the inner ear area . In a few other groups the swimblad-


The Fishes

of Tennessee

der has forward extensions to the auditory region .

Lampreys lack an elaborate inner ear sy stem, having
only a small auditory capsule . Thanks to the aid of sen
sitive hydrophonic equipment , many fishes , including
even small minnows , are now known , to produce
sounds (see Tavolga , 197 1 ). Sound communications
may be much more important in fishes than was pre
viously thought.
Chemoreceptors are well developed in fishes . Taste
buds , or gustatory organs , may occur externally on the
skin of fish (e . g . , some catfishes) and on barbels about
the mouth ; these organs are variously distributed on the
fins , lips , and in the mouth cavity and throat area as
wel l . Intimately related with taste in the location of
food is the sense of smell , which also is important in
orientation , communication , and perhaps predator de
tection. The olfactory senses are extremely well devel
oped in fishes , and some species of fishes are believed
capable of detecting substances at a few parts per tril
lion . For example , salmon homing to their natal streams
to spawn after years at sea are believed to locate these
waters over hundreds of miles through smell which was
imprinted soon after hatching. Odiferous chemicals ,
called pheromones , are emitted by some fishes (e. g . ,
the "fright" substance of minnows) for communication
through the water to signal danger, locate one another,
or perhaps facilitate courtship. The organs of smell , the
nasal rosettes, are located in the capsule served by the
incurrent and excurrent nares (Fig. 1 3a) through which
water is funneled. The single nasal opening of lampreys
(Fig . 12a) , through which water is alternately taken in
and expelled , leads to a sac lined with olfactory tissue.
Vision in fishes is of varied utility among groups.
Many fishes are sight feeders (e.g. , minnows , trout ,
sunfishes) , and many depend on well-developed vision
for not only feeding but species recognition and locat
ing cover. Other species , which inhabit turbid or very
deep waters , or subterranean waters , may have the eyes
and visual capacity much reduced or even nonfunctional
with concommitant increased development of other
senses , such as sound detection (e . g . , cave fishes) or
taste (e . g . , catfishes). In other fishes (e . g . , walleye ,
some marine groups) which are nocturnally active , the
ey es have evolved a brilliant reflective layer (tapetum
lucidum) on the inner surface of the eyeball to maxi
mize light gathering and , in some of these groups , the
eyes are very large. Several species of fishes are known
to be capable of color recognition and , based on the
bright color patterns , which probably facilitate recogni
tion in many species , this capability may be pervasive
among at least species inhabiting well-lighted environ
ments .

The eye of a fish , like that of a terrestrial vertebrate ,

is a ball-shaped organ with a transparent cornea and cir
cular opaque iris ; however, unlike the eye of terrestrial
vertebrates , the iris of the fish eye is scarcely dilative
and the lens is essentially spherical rather than elliptical
to compensate for the refractive properties of water.
The eyeball bulges somewhat from the head , and the
lens protrudes from within it , to give the fish virtually a
circular field of vision ; within the field , the lens focuses
at some distance to the sides but at close range to the
front , presumably to facilitate feeding . In some groups
(e . g . , herrings) vertically oriented , fixed eyelids
(adipose eyelids) have evolved (Fig. 33) .
As for other senses , fishes are capable of temperature
perception , and many species remain closely associated
with certain temperature regimes. The sense of touch is
thought to be somewhat limited in fishes , but some spe
cies (e . g. , some catfishes) are known to be sensitive to
tactile stimuli . Finally, one sensory system that may be
very important in fishes is that of electroreception . The
extent of development of these capabilities to sense ex
tremely weak currents is not well known for many
fishes , but it is known to be developed to varied degrees
in a number of families , reaching its highest state in
such groups as the South American electricfishes (Gy m
notiformes) which are also capable of emitting weak
charges . (The ability to emit stunning voltage levels has
developed in the notorious electric eel genus , Elec
trophorous . ) Catfishes have microscopic pit organs in
the skin for reception of minute electrical currents. This
system may be very important in location of objects or
prey as well as in communication , orientation , and nav
igation in some groups.
Skeleton and Internal Organs . We have already
touched on various skeletal components associated with
the head and fins and will only briefly further discuss
the fish skeleton . In higher bony fishes (Osteichthyes)
the body is supported internally by a more or less com
pletely ossified (bony) skeleton (Fig . 17) . However, in
chondrostean fishes (sturgeons , paddlefishes) much of
the skeleton is cartilagenous . In lampreys , the skeleton
is even less developed , consisting of a cartilagenous
cranial structure and gill complex and a spinal cord
around which vertebrae do not completely form ; thus
the embryonic notochord is retained through adulthood.
In bony fishes , skeletal features vary considerably
among groups . The presence and number of predorsal
bones varies , as does the number of epipleural ribs , and
epineurals , also known as "intramuscular" bones ,
which are common to such groups as herrings , pikes ,
trout , suckers , and minnows but not to "higher" groups ,



such as perches and sunfishe s ; these are often responsi

ble for complaints about some of these fishes being "too
bony to eat . "
When one dissects a fish , o r guts one while cleaning ,
several internal organs are readily apparent (Fig . 23).
The organs are contained within the thin layer lining of
the body cavity, the peritoneum, which is usually silver
or whitish but may be speckled with black pigment or
v irtually black , in which case it is usually visible as
such through the body wal l . Dark peritonea are typical
of some detritivorous and herbivorou s species . The
more variable organs to be discussed are the alimentary
trac t , gas bladder, and gonad s .
T h e alimentary tracts of many fi shes have a well
developed stomach ( e . g . , trout , perch , sunfishes) while
those of others are not well defined , appearing as part
of the tubular intestine . In such species as detritivorous
and herbivorous minnows , the gut may be extremely
long with many loops or coils . In species with well
defined stomachs , pyloric caeca often append from the
union between the stomach and gut; these vary in size
and number among groups .

In many fishes the gas bladder, or "swimbladder, " is

one of the l argest internal organs , located dorsally in
the body cavity and having the appearance of a white
balloon . This organ functions in regulating buoyancy
and is thus reduced or even absent i n fishes that have
more bottom-oriented life-styles . There are two basic
types of gas bladder. In many "lower" bony fishes , such
as minnows and trouts , a physostomous gas bladder ex
ists , which maintains a primitive connection to the ali
mentary tract , and which is presumably initially filled
during early development by gUlping air; the bladder is
regulated in later life by gas secretions from the blood .
The tubular connection between the esophagus and
swimbladder (pneumatic duct) allows some of these
fishes ( e . g . , gars , bowfins) to use the swimbladder as a
lung and breathe by gulping air at the water surface .
The primitive connection has been completely lost i n
higher fish group s (physoclistous condition). G a s blad
ders may be single chambered but are multichambered
in many species . The gas bladder also serves as a reso
nating chamber for the reception and production of
sound . Connections from thi s organ to the inner ear

swi m bladder
u ri n a ry b l adder

i ntest i n e
Figure 2 3 . Major internal organs of a fish (trout). Liver i s distended ventrally to reveal underlying organs; kidney i s generally
bound in tis sue along ventral aspect of vertebral column.


The Fishes of Tennessee

area are discussed above under Sense Organs and

Senses . For sound production , in some groups , spe
cialized muscles have evolved externally or internally
about the swimbladder to generate vibrations (e . g . , in
the drum family) .
In most fishes the gonads are tubular organs lying on
each side of the body cavity and terminating at the
urogenital opening , though they have different shapes in
a few fish groups . In some primitive groups (e .g . ,
bowfin , paddlefish) and salmonids and relatives , they
do not connect directly to the oviducts , with the eggs
first being shed into the body cavity and funneled into
the ducts . The gonad of lampreys is a single tubular
organ which is also remote from the urogenital opening .
Near the spawning season , the gonads of fishes are ob
vious and , in the case of females , may fill a substantial
portion of the body cavity; out of the spawning season
the gonads may be flaccid and difficult to discern . Ova
ries are recognizable by their typically yellowish color
and the granular appearance caused by the spherical egg
bodies within them , while testes are usually white and
contain thick fluid (milt) .

Reproduction and Early Development

Fishes exhibit diverse modes of reproduction; for a sys

tematic treatment of these , see Breder and Rosen
( 1966). The majority of fishes spawn by having the
female extrude eggs , which are externally fertilized by
sperm-bearing milt from the male (oviparous reproduc
tion) . These fishes may simply broadcast buoyant eggs
in midwater (e .g . , shad) or sinking (demersal) eggs on
the bottom (e .g . , walleye) , in which case the number
extruded is generally extremely high . Other species uti
lize crevices in rocks or submerged logs while others
construct gravel nests in which to deposit a lesser num
ber of eggs and abandon (e . g . , salmonids) or guard
them (e . g . , catfish , some darters) ; this latter task is of
ten accomplished by the male. A few fishes (e . g . , am
blyopsid cavefishes) incubate eggs in the mouth or gill
cavity until hatching . Some groups (e . g . , the "live
bearers ," Poeciliidae) have internal fertilization accom
plished by a male intromittant organ , the gonopodium ,
formed from a modified anal fin , with eggs incubated
internally in the female and born fully developed (viv
iparity) . This process is described more fully under the
family Poeciliidae .
In a few species (e . g . , some poeciliids , cyprinids) re
production by gynogenesis or hybridogenesis has
evolved in which all-female clonal populations have re
sulted from certain hybrid crosses (Dawley and Bogart ,

1989; Allendorf and Ferguson , 1990) . In gynogenesis ,

females of the hybrid "species" develop diploid eggs
such that , while sperm from the males of one of the
original parental species stimulates initial development
of the eggs , none of their genetic material is incorpo
rated into the zygote , thus resulting in exact clones of
the females . In hybridogenesis , diploid females of
hybrid origin produce haploid eggs containing the un
altered genome of only one of the original parents .
These are fertilized by normal males to produce "hemi
clonal" progeny that are all female and will repeat the
process to produce additional , different hemiclones .
Some fishes are hermaphroditic , possessing both
male and female gonadal tissues , and functioning both
as males and females in successive spawning acts (syn
chronous hermaphroditism); this condition is common
in some marine groups and is known occasionally in
such groups as topminnows (Fundulidae) and striped
bass (Moronidae) . Other species (mostly marine) un
dergo complete sex reversal , functioning as one sex
when younger and the other as an older adult (consecu
tive hermaphroditism).
With a few exceptions , most temperate fishes spawn
sometime between late winter and midsummer when
waters are warming . Before spawning , fishes generally
engage in some form of courtship, which may range
from simple following behavior to elaborate displays
especially by the males-which can consist of swim
ming patterns , physical contact , fin-flaring , and colora
tion changes . Before and after spawning , males of
nesting species often vigorously defend territories about
their nests against intruders of the same or different spe
cies . Some nest-building species (e . g . , sunfishes) en
gage in "cuckoldry," in which younger males posing as
females enter the nests and fertilize some of the eggs .
During the actual spawning act , in midwater
spawners , the spawning pair usually just swim upward
in close proximity extruding eggs and sperm . In benthic
spawners , it is usual for the male to press the female to
the substrate in some fashion with the urogenital open
ings in close proximity. Fishes are generally observed to
vibrate briefly at the time of releasing gonadal products .
In species that bury eggs , the female or both sexes may
vigorously vibrate the anal fin to entrench the eggs.
The eggs of mid water spawners may drift for several
days and undergo early development while in transit.
Demersal (heavier than water) eggs , which may be ad
hesive , generally develop while on the substrate . Newly
hatched fishes (Fig . 24a ,b) are termed larvae or "fry. "
These have little resemblance to adult fishes and con
tinue to undergo development (ontogeny) after emer
gence from the egg . Earliest stages usually survive for



pectoral fin bud

b ra i n l o be s

y o lk s ac

ot o l i t h s


oil globule

inte stine

g a s b l ad d er

pelvic f i n bud

incipient fin ray s

Figure 24. Early life h istory stages of a fish: a) yolk-sac l arva.; b) pos tlarva; c) early j uvenile.

several days on nutrients gained from absorption of the

peratures . These early l i fe"stages of a fish lead a very

yolk sac . This stage i s followed by a transition to an ac

tenuous existence , and a large percentage of fishes fall

tively feeding stage (usually on microinvertebrates)

victim to predation or other perils during this period and

termed a postlarva . Eventually, postlarvae transform

thus never reach adulthood . References of particular in

into small fishes , more or less resembling adults ,

terest on the early development of fishes are Fish

postlarval stage ranges from a few days to several

( 1 9 32), Hogue et al . ( 1 976), S nyder ( 1 983), Moser et

( 1 984), and the series of volumes in preparation by
Wallus et al . (e . g . , 1 990) on Ohio River drainage larval

weeks and , in general , is inversely related to water tem-

fishes .

termed "j uveniles . " The time of early development

from spawning through hatching and tran sition to the


Th e Fishes

of Tennessee

al .