i

Not all fishes are cold-blooded. In some fast-swimming species of tuna

and mackerel shark a wonderful net of arteries and veins conserves

the heat of metabolism to increase the power of the swimming muscles

v

by Francis G. Carey

ishes are regarded as cold-blooded

T h e o x y g e n in t h e w a t e r dowing cold. Indeed, for a fish to raise its body
animals, that is, animals whose through the & of a fish diffuses into temperature by metabolic processes
body temperature is the same as venous blood pumped to the gills by the alone would seem to be an impossibility.
the temperatuYe of their surroundings.
heart. The blood may reach the gills at
The fact is that not all hhes are cold- a somewhat elevated temperature as a
unas and mackerel sharks manage to
blooded. The first man to ~vriteabout a result of having accumulated metabolic L a y w1
< r m in apparent defiance of
warm fish was a British physician named heat. Heat, however, d&es
10 times nature because they possess a special
John Davy. He was voyaging in the faster than oxygen molecules do, so that structure in their circulatory system It
Tropics in 1835, and tlie ships company by the time the blood in the gills is satu- is the rete-mnabib, or wondeiful net,
was supplementing its rations by fishing
rated with oxygen its temperature has a tissue composed of ciose~yinterninDavy was intrigued by tlie copious blood
gled veins and arteries that was b-st
dropped to the temperature of the water.
The oxygenated blood now flows back noted in vertebrates by students of anatand mammal-like red flesh of one of the
fishes brought aboard, a species of tuna
through the fishs circuIatory system, and omy in the 19th century The Fiench
known as the skipjack. He took the temthe oxygen is utilized in the body tissues. naturalist Geoi ges Cuvier observed the
perature of several skipjack and discov- It is this process that generates the metapresence of the rete In the tuna in 1831,
ered that they were wanner than tlie bolic heat; the rise in temperature is the last year of his life Four years late!,
water they swam in by 10 degrees Celproportional to the amount of oxygen the same year that Davy measured the
sius (18 degrees Fahrenheit) In the extracted from the blood. In any given
temperature of tlie skipjack tuna, the
years since Davys observation most oth- volume of oxygenated blood the volume German anatomist Johannes M d e r forer species of tuna have proved to be
of extractable oxygen is less than 20 per- mally described the structures in the viswarm-blooded and so has one family of cent. Taking into account the usual ca- cera of the bluefin
The rete provides a thermal barrier
sharks: the . mackerel sharks. At the loric yield of a fishs foodstuffs, that limWoods Hole Oceanographic Institution ited supply of oxygen will support only against the loss of metabolic heat The
my colleagues and I have long been fas- enough metabolism to raise the tempera- mass of fine veins and arteries permits
cinated by these warm-blooded fishes.
ture of the blood one degree C. More- the free flow of blood for transport of
What is it that prevents most fishes over, even that slight temperature in- oxygen and other molecules, but at the
same time it short-circuits the flow of
f i om maintaining a body temperature
crease is lost to the water when the blood
that is more than negligibly higher than next passes through the gills, so that no heat from the body tissues to tlie
and shunts the accumulated heat back
tlie temperature of the surrounding wa- heat accumulates to keep the fish warm.
to the tissues. In the ret2 an artery, carter? It is not, as might be supposed, the er than the surrounding water.
loss of surface heat that comes from beChanges in the fkhs rate of metabo- rying cool, oxygen-] ich blood from the
ing immersed in a cool medium Rather lism do not affect the cycle, because any gills to the body tissues, branches to
it is the heat loss implicit in the fishes exercise 01 other activity that produces form a mass of small vessels The aiteiial
mode of respiration All land animals
more metabolic heat also demands more network runs parallel to and is interminand some aquatic ones obtain the oxygen oxygen. An increased demand of oxygen gled with a similar mass of &e veins
means a gieater loss of heat thhrough the carrying warm, oxygen-depleted blood
required to support their metabolism by
breathing air, which is rich in oxygen gills, so thal whether the fish is at rest or from the tissues back to the gills The
and has a low capacity for absorbing exercising violently it will always be system of closely associated arteries and
lieat. Water, from which fish must obtain their o.xygen, contains scarcely 2 5
percent of the oxygen found in an equal
RHTE !WIRdBILE, or wonderful net, of the bluefin tuna is seen in cross section. The rete
volume of air but has 3,000 times tlie
is a system of parallel smaH arteries and veins that supplies and drains the band of darkcapacity Of air
absorbing heat As colored muscle that is used for sustained swimming effort. The system constitutes a counthe Ieat lost in extracting Osygen tercurrent heat exchanzrr: venous blood warmed by metabolism gives up its heat to cold,
from water can be loo~ooo
greater newly oxygenated arterial blood fresh from the fishs gills. The effect is to increase the temthan the loss in elctractkg the Same perature and thus the power of the muscle. In thk bluefin rete sample the vessels have been
amount of oxcgen from air.
visualized by the injection of latex: red latex in the arteries and blue latex in the veins.

36

I
GILLS

HEART

POSTCARDINALVEIN

DORSAL AORTA

DORSAL AORTA

POSTCARDINAL VEIN
1
7
CUTANEOUS VEIN

CUTANEOUS ARTERY

V.4SCULAR SYSTEMS of cold-blooded (top) and warm ( b o t t o m )fishes are different. The
main vessels in most fishes, the cold-blooded ones, run along the backbone and radiate outward to the small vessels (not shown) that supply the muscle. In warm fi-hes the central
vessels are smaller; most of the blood flows through cutaneous vessels under the skin and
thence through countercurrent nets I not shown) that supply the muscle. This arrange.
tnent puts the cold end of the exchanger near the skin and the warm end in warm ti-sues.

veins acts as a Countercurrent heat eschanger. There are large areas of contact
between the two kinds of fine vessel, and
heat is readily transferred through the
vessels' thin walls. Therefore the walm
venous b l o o d i s c o o l e d a s i t flo~vs
through the rete, so that little or no heat
is lost when the venous blood at last
reaches the gills. At the same time the
cold arterial blood is warmed, so that
when it reaches the interior of the fish,
it has nearly reached the temperature of
the warm body tissues.
A rete of this type serves exclusively
as a countercurrent heat exchanger. Its
vessels, although small, are too large and
their walls are too thick to aIIow any significant difusion of oxygen molecules
from the arterial blood to the venous
blood. Other retia that do serve as gas
eschangers are found in many species of
fish [see '' 'The Wonderful Net,'
by
P. F. Scliolander; SCIENTIFIC ,IJIERICA
April, 19571. The tiny vessels that coniprise thein are capillaries, and so oxygen
molecules readily pass from one to another. Such retia are found in the swim
bladder and the eye.
"

-'

major adaptive advantage of an elevated body temperature is greatIy

enhanced muscle power. If the clifference in temperature between two otherwise equivalent muscles is 70 degrees C.,
the warmer muscle is able to contract
and relru three times more rapidly, wi t l i
no reduction in the force applied a t each
contraction; thus it can generate three
times as much power. This kind of increase in power is evidentlv essential in
flying animals such as birds and bats;
their body temperatures are characteristically high. Even some large 'insects
with a heavy wing loading elevate their
body temperature when they fly. For esample, cicadas and locusts are often unable to take off until they have warmed
up by shivering or by basking in the sun.
Like flying, high-speed swimming is a
demanding form of locomotion. Water is
a dense and viscous medium, and to
move through it rapidly requires not
only fine streanilining but also substantial muscle power. The high body temperature of tunas and mackerel sharks
apparently helps to provide the estra
power needed for high-speed swimming:
the most important of the vascular I y o t
es&angers are those that serve the darkcolored swimming muscles.
Vertebrate muscle is generally a mixBLOOD SUPPLY TO MUSCLE of the bluefin tuna is primarily through four pairs of
ture of dark-colored fibers and light-co!cutaneous vessels. I multitude of smaller vessels brJnch from them, forming the thick
orerj ones. The dark fibers contain high
vascular d.ib above and I d o u the dark muscle, the rete, from which :hc dark muscle ij
concentrations of the ory2en-tmnspor:supplied. The light muscle is supplied by bands of alternating arteries and veins that slant
iiig pigments and enzymes associated
through the muzcle from the segmental vessels, thus serving as two-dimensional heat exchangers. Other, non-heat-exchanging arteries and veins run out from the central resrel~. with oxiiintive metabolism; they are also

?&!y suqp1;ed w i t h blood. T h e lightc8lorsd Gben usudy have a less well:,e!rJg,ed blood supply. They can h c 4L1 a-:&out oxygen during activity by
metabo:king foodstuff through fermen;:don, recovering later when they are at
rest. Sluscles that are continuously active, for example the heart, are made up
largely or entirely of dark fibers. Muscles
that alternate between periods of rest
and short intervals of activity contain
few, if any, dark fibers.
As anyone who has seen a fish steak
knows, the two kinds of inuscle fiber are
sharply segregated in the axid musculature of fishes, For example, in many
t u n s a large inass of very dark muscle,
well supplied with blood, lies in a broad
band between the backbone and the skin
along the midplane of the body. This
dark muscle can operate continuously;
its contractions propel the &sh as i t

swims at ordinary cruising speeds The

blood that supplies the muscle reaches
the muscle fibers after passing through
a large heat-exchanging rete. As a result the muscle is wdrmer than any other
part of the fish's body.
In order to providc for this large heat
exchanger the arrangement of the tuna's
circulatory systeiv has been drastically
altered. Most fishes have a central distribution system for the blood. The main
blood supply to the muscle flows outward through a Iarge artery, the dorsal
aorta, and returns through the postcardinal vein; both are deep inside the
6sh just below the backbone. From these
central blood vessels segmental arteries
and veins radiate out to the periphery.
In the tuna the normal pattern of circulation to the muscle has been alniost
entirely reversed The major hlood vessels are no longer the central artery and

FISHES with warm bodies have streamlined

bodies a n d heavily muscled tails with crescent-shaped caudal fins.

FX%!'-SVIXGING

vein. Instead four artery-vein pairs just

under the skin, two on each side of the
fish, provide the main blood supply [see
top illustration on opposite page]. Froin
these cutaneous vessels Iarge numbers of
tiny blood vessels, each only a tenth of a
millimeter in diameter, branch off. The
tiny vessels intermingle to form slabs of
vascdar tissue, the retia, that lie close to
the upper and lower surfaces of the dark
muscle [see bottom illustration on opposite page 1. These retia are the heat exchangers that ensure the warmth of the
dark muscle. The blood to the light muscle also comes from the large cutaneous
vessels by way of pairs of s e p e n t a I
-arteries and veins that r u n up and down
over the surface of the muscle and send
n u m e r o u s branches into it. These
branches are in the form of vascular
bands, ribbons of alternating arteries
and veins that act as heat exchangers.

The ones illustrated here are three tunas: the bluefin (a),the SEpjack (b) and the wahoo (c), and a mackerel shark, the
(di.

39

The area of thermal contact between

arteries and veins in these two-dimenT:onal structures is smaller than it is in
&,e massive heat exchangers serving the
dark muscle, but apparently it IS adequate for the typically low rate of blood
&W to the light muscle and ensures that
much of this tissue too is warm.
One resdt of the inside-out arrangement of the main blood vessels in a warm
fish is that the large arteries and veins
that give rise to the retia are just under
the skin. This puts the cool end of the
heat exchanger near the surface of the
fish and the warm end deep in the interior, an arrangement that minimizes
surface heat loss.

By measuring the temperature of various parts of freshly caught bluefin tuna

we have learned that the warmest regions are within the dark muscle [see
illustration 011 page 4.21. The hot spots
are not, a s one might expect, in the
deepest part of the muscle. The reason
is that the deep interior receives some of
its blood supply from the dorsal aorta
through a system that has no specialized
heat exchangers. The highest muscle
temperature is found near the center of
the dark-muscle band on each side of the
fish, but substantial areas of muscle tissue, both dark and light, are considerably warmer than the temperature of the
water from ~ h c hthe fish are taken.

The bigeye tuna and the albacore

have circulatory systems that closely resemble the bhefin's. Other tuna species,
particularly the skipjack and to a lesser
extent the yellowfin, eIevate the ternperature of the muscle in a different wav
Although the pairs of cutaneous arteries
and veins are still present, the heat eschangers that arise from them may contain no more than a single layer of fine
blood vessels. In these fishes the pincipal heat exchanger is connected to the
central artery-vein pair below the backbone. This centra1 rete, a rodlike vascular mass that ma17 be larger in diameter
than the backbone itself, extends along
the vertebrae in the region above the

ARK MUSCLE
DORSAL AORTA
POSTCARDINAL
VEIN

.RnE
MlRAElLE

40

j. Body-temperature measurements
;!c:F,jack attest to the efficiency of the
;;:--haped heat exchanger. these Iittle
fi& are often 10 degrees C. warmer than
the water and their warmest muscle is
adjacent to the backbone.
'la9p

t o

of

interesting coincidence that the

It: acankere1
sharks, a
that is evogroup

lutionarily far removed from higher bony

fishes such as the tunas, should have deveIoped a heat-exchange system that is
so much like the bhefh's. The deep interior blood vessels along a mackerel
shark's backbone are small. The major
blood supply circulates through a single
artery-vein pair running just below the
skin along each side of the fish. The
paired cutaneous blood vessels give rise
to a single massive rete that warms the
shark's dark muscle. In some mackerel
sharks, such as the mako shark, the heat
exchanger is a solid slab of blood-vessel
tissue resembling the bluefin's (see illustration on opposite page]. In others,
such as the porbeagle shark and the
white shark, the rete is diffuse, with
many bundles of fine blood vessels extending into the dark muscle. In mackerel sharks the body-temperature distribution closely resembles the distribution in the bluefin, most of the sharks
muscle is warmer than the water they
swim in, and the warmest parts of all
are twin regions within the dark mtiscle.
Tunas and mackerel sharks possess
heat exchangers that are not associated
with muscle tissue but serve the organs
of the body cavity In the albacore, bigeye and bluefin tunas dense bundles of
intermingied fine arteries and veins are
found on the surface of the liver; in a
large bluefin the bundles may be five
centimeters in diameter The arrangement in the mackerel sharks is M e r e n t
because the artery that in other fishes
carries the main blood supply to the viscera is either very s d or nonexistent.
Instead the visceral blood supply is provided by arteries that are i n s i g d c a n t in
most other fishes. These arteries penetrate into a much enlarged veiious space
knon7n as the hepatic sinus Venous
blood flows through this space on its way
from the viscera back to the heart The
arteries branch and branch again within
the hepatic sinus until tlie space IS vkr
tually filled with a spongelike mass ot
tin. vessels; the cool blood within them
is warmed by tlie bath of venous blood
Visceral retia do not keep the organs
of the body cavity warm continuously,
the temperature can vary from a level
equal to the warmest muscle to one only
slightly above the water temperature.

10\
5

I
10

,
I
30
3

c
15
20
WATER TEMPERATURE ( D E G R E E S CELSIUS)

MAXIiMI.Ti1 XUSCLE TEMPERATURES are plotted against water temperature for bluefin
(color), skipjack (solid black) and penowfin (open black). The bluefin r e p l a t e s its temperatnre almost independently of water temperature. The skipjack is warmer than the yell o w h but both tend to maintain a fixed temperature difference above water temperature.

The variations may be correiated wkth

the mackerel sharks' and tunas' digestive
activity. The tunas in particular have remarkably small body cavities, and the
mass of their visceral organs is limited.
Raising the temperature of the viscera
may speed digestion. Conversely, during
times of digestive inactivity the viscera
may be allowed to cool.
Since tunas are warm, we wondered
whether they could control their temperature at a constant level and thus be
independent of water temperature or
whether they warmed and cooled as the
water temperature changed. Other investigators had noted that yellowfin tuna
and skipjack tend to maintain a temperature a &xed number of degrees
above that of the water-something that
might be accomplished by an unregulated heat exchanger. These two species
are found only within a rather narrow
range of water temperature, however,
and so n7e turned our attention to the
bluefin tuna, which is equally at home
m tropical and in subpolar waters. We
measured the maximum temperature in
the muscle of bluefin taken along the
coast from the Bahamas to Newfoundland Comparison of their muscle tem-

perature with the temperature of the

water in which they were captured suggested that they control their temperature quite well Over a %-degree C
range of watei temperature the average
muscle temperature changed only siu
degrees, bluefin from the 30degree
waters of tlie Bahamas were only a few
degrees warmer than the water, whereas those from seven-degree northern
waters were 20 degrees above the water
temperature
There are tw7o ways an animal might
achieve such control One is by means
of the swift thermoregdatury adjustment to environmental temperature
changes that is characteristic of birds
and mammals, which can vary their rate
of heat loss in several ways and also generate exfxa heat by increasing their metabolism The other is the much slower
process of acclimatization, by which
cold-blooded animals adjust their activities to changing temperature and
which involves cellular and e v e
changes over an extended period of
time. In order to discover which of these
mechanisms the bluefin were using we
decided to try a field e q e r k e n t i n
which we would monitor the body tem41

into cold water. If the bluefin w7ere

capabk of rapid thermore,pIation of the
kind shown by mammals, their temperature should remain constant during a
sudden change in water temperature.

WATERTEM P ERATU RE
19 DEGREES C.

WATER TEMPERATURE
21 DEGREES C

TEMPERATURES were recorded at various points in bluefin tuna (a! and mako sharks
( b ) .Isotherms show that the warmest parts of the fishes are within the dark muscle on each
side of the body. Much of the rest of the muscle is also considerably warmer than the water.

42

/ e found that the coast of Nova ScoaitJ

was 7n ideal setting for our project The Coolen brothers of Fox Point
operate pound nets in St Margarets
Bay, and they frequently catch bluefin
tuna in their nets Moreover, the coastal
waters of Nova Scotia are characterized
by a marked thermoche (a sharp diop
in temperature as the depth increases),
so that a free-swimming fish might encounter a wide range of water temperatures
We used a small harpoon equipped
with two thermistors in our first attempts
to record bluefin temperatures One
thermistor sensed water tempelature
and the other muscle temperature, they
were connected to an indicatoi by a
1,000-yard length of field-telephone wii e
that could unreel as the harpooned fish
swam away We failed on almost every
attempt the fish died or the harpoon w x
pulled out when the wire was snagged
by kelp or by lobster-pot lines It was at
this point that my colleague John Kanwisher suggested that we telemeter the
information from the fish.
The telemeterlng devices we selected
were battery-powered acoustical transmitters Their service life was from one
daj7 to three days and their range was as
much as Eve miles. The instrument package was mounted on a small harpoon
that could be driven through the fishs
thick skin with a minimum of injury to
the muscle. The thermistor that sensed
the muscle temperature was in the tip of
the harpoon and the one that sensed the
water temperature was attached. to the
transmitter, outside the fishs body on
the harpoon shaft For other experiments, in which we measured the stomach temperature, an instrument package
was pushed down the bluefins thioat
and into its stomach, the thermistol that
measured the water temperature was at
the end of a wire we led out thiough
the tunas gill slit The transmitteis
broadcast the tempeiature of the fish and
the temperature of the water during alternate one-minute periocis
Our tracking vessel was equipped
with a directional hydrophone that enabled us to follow the tuna We rotated
the hydiophone until the telemetei ed
signal was at a maximum and ;:?en
steered the vessel on that heading At
hourly intervals we lowered a bat??>thermograph to measure the tercpera-

signals.
~ , placed
v ~ transmitters on or in 14
bluefin tuna. The longest we tracked a
fish was 54 hours; by then the bluefin
had reached a position 130 miles offshore. We soon found that, as commercial fishermen had told us, tuna
avoid changes in water temperature if
they can. Most of our specimens remained near the surface or at least on
the warm side of the steep thermocline
that separated the upper waters from the
cold depths. Some of the bluefin would
dive through the tlermoche, but they
spent only a few minutes in the colder
water before returning to the warm side
Near the end of our efforts ~ 7 ewere
lucky enough to get a most satisfactory
res& The specimen was a 600-pound
bluefin with a transmitter in its stomach
The fish had been handled quite roughly
wkile the instrument was being installed
Perhaps for this reason as soon as it was
released from the pound at about 9:00
A.M. it swam down through the thermodine into water 14 degrees C colder
than the s q f a c e water in the pound.
When .the fish was released, its stomach
temperature registered 21 degrees C.
During its four-hour stay in the fivedegree water the temperature of its
stomach gradually fell to about 19 degrees. Early in the afternoon the fish l e turned to the warm side of tlie thermocline and remained in water that registered between 13 and 14 degrees for the
rest of the day. In spite of a change in
water temperature of nearly 30 degrees
C. the temperature of the fishs stomach
remained around 18 degrees The fact
that the deep-body temperature of the
fish remained nearly constant over extended periods in both cold water and
warm indicates that the bluefir was indeed thermoregulating. Just how the fish
do this we do not know. Presumably it is
by somehow varying the efficiency of the
heat-exchanging retia.
\Ve were not fortunate enough in our
muscle-temperature e.xperiments to have
a bluefin stay on the cold side of the
thermocline for any substantial lene@ of
time. One of the muscle-tagged fish,
however, did swim for hours in water
khat was gradually decreasing in temperature The water temperature dropped
four degrees C. in one 90-minute period.
During this interval the muscle temperature of the bluefin sIowIy rose from a

TIME

(HOURS)

TELEMETRY RECORD compares the temperature i n the muscle of a bluefin (color) with
that of the water !b&zck)through which it swam for three hours. The muscle temperature
was held constant as the water temperature declined gradually from about 1 4 t o 10 degrees.

I
i

10

I
3

TIME (HOURS)

RECORD O F TEMPERATURES from the muscle of a dusky shark (color)and the water it
swam in (black? shows a Ziderent relation. The dusky shark is a cold-blooded fish and its
muscle temperature stayed close to that of the water, dipping during even short dive:.

43

4
TIME (HOURS)

RECORD FROM TUNAS STOMACH shows the effect of the visceral, as opposed to the muscular, temperature-control system. The
temperature was telemetered from thermistors in the tunas stom-

little more than 23 degrees C. to above

24 degrees.
Our experiments showed that the
bluefin does not maintain its body temperature at a constant level as, for example, human beings do. The temperatures of fish from the same school can
fluctuate over a range of five degrees or
so. The greatest fluctuation we recorded
was provided by a second tuna tliat carried a stomach temperature transmitter.
At the time of release the temperature of
the fishs stomach was 79 degrees C.
During the b s t day of tracking its stomach temperature slowly began to rise. By
the. second day, although the water temperature had remained constant, the
fishs stomach was registering 26 degrees
C. Perhaps the fish was trying to digest
t 11 e t r a n s ni i t t e r 1
In other readings we found that muscle temperature tended to cycle over a
narrow range that is unrelated to the
temperature of the water. All these variations tell us that whereas tlie bluefins
various organs are being kept much
warmer than the surrounding water,
precise temperature control is either not
necessary or not possible In effect, the
fish has a sloppy thermostat. Nonetlieless, these fish can be said to thermoregulate in tlie same sense tliat birds and
mammals do.
How did the tunas and mackerel
sharks come to develop these remarkably efficient heat exchangers? Althougli
44

ach (color) and outside the fish in the water (black). M7hen the
tuna was released, it swam down into cold water and stayed there
for four hours, but its stomach temperature decreased only a little.

the rete is certainly a complex vascular

array, its mode of evolution was probably quite simple. For one thing, arteries and veins generally follow the
same path as they travel between organs
and within tissues. Every such pairing of
an artery and a vein is a rudimentary
heat exchanger; if at one end of the
system there is.either heating or cooling,
then tliere will be some heat transfer between the paired blood vessels. This
rudimentary exchange is probably what
accounts for our observation that fishes
tliat ought not to be able to warm themselves at all by metabolic means nevertheless may have muscle temperatures
from one to two degrees C higher than
the water temperature.

Suppose

that such a slight rise in tissue

temperature offers a genuine selective advantage to an evolving species. In
such an event there already exists (as
part of tlie normal embryonic development of the circulatory system) a basis
for the development of more advanced
heat exchangers. The circulatory system
of the developing embryo Erst takes the
form not of discrete blood vessels but of
beds of interconnecting spaces and channels. Later, as certain channels become
important, these routes become larger
and other channels atrophy and disappear. liloreover, there is a tendency
toward the deveiopment of multiple
channels. Several such channels may

forrn within a single bed, with the result

that the vascular system of the adult will
have some duplicate components Therefore in order to form the mass of parallel
blood vessels that is an efficient heat exchanger, only a mod8cation of tlie normal embryonic pattern is required, not a
radical genetic change
A fast-swimming predatory fish finds
a n abundance of food awaiting it in tlie
form of fast-swimming squid, herring
and mackerel that slower predators cannot capture. In terms both of streamlining and of muscle power the tunas die
probably the swiftest predatoi s of tlie
open ocean Yellowfin tuna and wahoo
tuna are noted fol their speed they have
been observed to reach speeds in escess
of 40 miles per hour during 10- to 20second sprints Mackerel sharks (at least
the mako and porbeag!e) probably also
enjoy tlie benefits of fast swiniming
Such a shark, with its bulky, musculai
body, it3 streamlining and its narrow,
crescent-shaped tail, looks remaikably
like a tuna That the two unrelated
groups of fishes independently evolved
the same means of raising their body
temperature must surely be connected
to tlie adaptive advantage of increased
swimrmng speed, the extra power ava~lable from warm muscle must have been
decisive in achieving that speed It 15 a
classic example of parallel adaptations
that evidently gave both groups access
to an underexploited source of food