OFTHE
BIOLOGICAL SOCIETY OF WASHINGTON
NOTES ON THE COLORATION OF FISHES!
BY r.AKTOX
A.BEAN AND ALFRED
C.WEED.
[Published by permissionofthe Secretarj of theSmithsonian Institution.]
While
it is wellknown
that the color of fishes of thesame
species variesmore
or less itmay
not be amiss to call especial attention tosome
of themore marked
changes thatmay
benoticed.
The
color changes in living fishesmay
bedue
tovarious emotions as fright, curiosity, anger, or sexual excite- ment, or they
may
hedue
to environment, as the colorof the bottom, the color of the water, distance of the fishfrom
the surface orfrom
bottom, distance from objects attached to thebottom
or floating at the surface, presence in theimmediate
vicinity of grass, sticks, etc.Many
peoplewho
are familiar with the fact that the colorand
color patternmay change
in thesame
species havean
idea that this takes place slowly.We hope
toshow
that this does not necessarily followand
will givesome
illustrations of color changeswhich we
have noted in various species in aquariums, principally that of theBureau
of Fisheries atWashington,
D. ('.The
LargeMouth
Black Bass (Micrepterus salmoides)when
viewed asan
ordinarily preservedspecimen
usuallyshows more
or less traces of a black lateral band. In the large specimens this
may
be very faint but can usually be discerned. Similar specimensof theSmallMouth
Black Bass(Micropterusdolomieu)show no
tracesof this blackband
but are plain olivaceous in color.There
are several livingspecimensofeach speciesin theaquarium
at theBureau
ofFisherieswhere we
have spentsome
time watching the color changes.One
largespecimen which showed
verystrongly the typical black lateralband
of the Large18—Proc. Biol. Soc.Wash., Vol.XXIV,1911. (69)
70 Bean and Weed —Coloration of Fishes.
Mouth
Black Basschanged
in less thantwo
seconds, so that theband was
entirely invisibleand
the fishappeared to be aSmall Mouth
Black Bass. In preserved specimens ofboth species the color, exceptforthe black lateral band, is plain; thatis, there areno
mottlings or marmorations, butin theaquarium, where
thefish are soused to seeingvisitors that theypay no
attentiontotheir presence, the color
may
be seen tochange from
plain olivaceous without the black lateralband
to plain olivaceous with such a band,or toa mottlingof lightand
dark green with orwithout the black band.The
LargeMouth
Black Basswhen showing
the mottled color usuallyshows
the blackband
; speci-mens
of theSmall Mouth
Black Bass usually do notshow
the blackband and
themottlingshave
agreater tendencyto appear as dark cross bars. It is difficult to determine the exciting causes of these changes as fish hiding in crevicesof the rocks willshow
both colorationsand
fishesswimming show
both colorpatterns. It appears, however, that fish
swimming
freely or hidingin ratherdark
holeshave
agreater tendency to exhibit the plain coloration,whilethose restingon
thebottom
orswim- ming
close to the rocks at the side of theaquarium have
a greater tendencytoshow
themarbled
ormottled color pattern.Preserved specimensof
Tautog
(Tautoga onitis) are usually nearly plain black but withmore
or less distinctvertical lightand dark
bars of about equal width. In theaquarium
theyshow
remarkable variations ofthesetwo
color patterns.When swimming
freely the color isusually plain blackand when
rest- ingon
thebottom
or at the sides of theaquarium
the bright pattern isvisible.However,
these colors are not constantand swimming
fishmay
be seenshowing
the bars whilethose restingon
thebottom show
thedark
color.The
color changes are frequently very rapid.We have
seen a fishswimming
in avertical circle about
two
feet in diametershowing
the bright coloration only near thebottom
of theaquarium and
the black color in theupper
halfofthe circleat each turn.Inthe
New York Aquarium
at BatteryPark
noticesareposted in frontof certain tanks containingsome
of the tropical angel fish, calling attention of visitors to rapid changes in colorand
color pattern of this fish.These
changes are truly marvelous.Waves
of blue startingat thehead
pass to the tailand
disap-pear.
While one
iswatching
thefish the color willbecome
allblue or all brownish or half blue
and
half brown, etc.Indeed
the changes are toonumerous
to mention.The
Pigfish (Orthopristis rhn/sopterus)shows
very remarkable changes in colorand
color pattern.There
are three perfectly distinctcolor patternswhich
appear to be about equallycom- mon. One
isaperfectly plain light ordark
gray withoutspots or bars ofany
kind.Another
has thissame
grayground
with severalheavy
black cross bars,which sometimes
aremore
or less brokenup
intodark
mottlings.A
thirdshows
instead of the cross bars a black linefrom
theend
of the snout to the base of the dorsal finand
along each side of this fin.Below
this is a
narrow
light stripe with aheavy
blackband
belowit.A
second darkband
extendsfrom
the eye straightto thecaudal peduncle.The
lower edge of thisband
isjust abovetheupper
lip.
Between
thisand
thedark band
aboveit is a light stripe about thesame
width as either of thesedark
bands.The upper
darkband
crosses the front of thehead
aboutmidway
between the lowerband and
the front of the dorsal fin.There
arealso a few small black lines across the front of thehead which make
themarkings
appearmuch
like a bridle.One
of these lines passesfrom
theupper
edge of the opercle to theupper
edgeof orbitand
then acrossfrontof head.Sometimes
thetwo
setsof barsfadeout exceptat pointswhere
theycross, giving rise to a seriesof square black blotcheson
the side of the fish.Any
of these color patternsmay
appearinstantane- ously, followingany
other.Sometimes
thechange may
belikened to the fading of
one
pictureand
the appearance of anotherin a stereopticon.Specimens
of the Pinfish(Lagodon
rhomboides) in the aqua-rium show
mostly a color patternmade up
simply oflongitudi- nal blue stripes of greater orless intensityon
agrayish ground.However,
at times black cross bars appear.The
appearanceand
disappearance of these barsmay
be practically instantane- ous or itmay
be quite slow.The ground
color between the blue linesoftenbecomes
a brightshell pink.Many
ofourcommon
fresh-water sunfishesshow
remarkable changes in their color in life.The Common
Sunfish orTobacco Box
" (Lepomis gibbosus) does notshow
these changes so well assome
of the other species.The
'Green
Sunfish"
(Lepomis cyanellus), a widely distributed fish in the
72
Bean and Weed —Coloration of Fishes.
Mississippi Valley,
which
has been introduced intothePotomac
River,shows
these color changesmore
thanany
other sunfish withwhich we
are acquainted, in factthechanges of colorand
color patterns are almost asnumerous and
as rapid as in the angelfish. In addition to thenormal
colorchanges, anincrease or reduction in the intensity of the color, or achange from dark
to light color,due
to emotions orenvironment, there aremany
other changes forwhich we
can onlyguess atthe cause.We
shall probably neverknow why
one fishwhen
frightened will turn pale while another in thesame aquarium
willbecome
very strongly barred.The commonest
color, visible in preserved specimens, in dead fishand
in fish just takenfrom
the water, is aplaindark
olivaceous withmore
orlessdark red in the vertical finsand
withtwo
blue lines across the cheek.In the
aquarium
thismay
be seen tochange
to a rich goldenbrown
with or without vertical cross bars or to a plain light coloralmostsilvery.The
vertical cross bars aresometimes
aspronounced
as in the Yellow Perch (Percafiavescens), at other times the whole side of the fish is covered withminute
shining specks withoutany
apparent regular arrangement.Any
ofthese color patterns
may
appear practically instantaneously.The change from
vertical barsto the plain coloration withlight specksmay
occur apparently in a flash.Where
a few speci-mens
of approximately equal size are confined in a smallaquarium
these colorchangesmay
bemost
readily noted.The
intensification in color of the largest specimen
when
a smaller onehappens
to stray into forbidden territoryand
thesudden
paling of thelatter as he retreatsmay
lie seen very frequently, especiallywhen
the fish are beingfed.The Long Eared
Sunfish (Lepomis auritus),known
also asRed
BreastedBream,
iscommonly
amore
or lessyellowish fish with orange spotsand
mottlings over the entire body, thesesometimes showing
asmore
or lessindistinct cross bars.There
aremany
specimens of this species in theaquarium
at theBureau
ofFisherieswhere we have watched
their color changes.Most
ofthe specimensshowed
adark
olive colorwith theverti- cal finsand
the breastdark
red.Sometimes
faint cross bars show. This coloration is sosimilar to thatof Lepomiscyanellus that only the position ofsome
blue lineson
theoperculum and
theshapeofthe"
ear'' indicated thespecific difference.
From
this olive coloration fishes
would
quicklychange
to a goldenbrown
with or without dark cross bars or with barsformed
of small spots slightly darker than theground
color.One
fishwhich we
watched forsome
timeand
which, hy its actions, appeared to be amale more
or lessapproaching
the breeding conditionwas
a brightyellow colormottled over thewhole
side with dark orangespotseachabout thesizeofascale.The
breastwas
a clear bright orange.The
pectoral finswere a clearlemon
yellow.There was
hardly a trace of cross bars although at times a few could be seen quite indistinctly. This fishwas
observed several timesand seemed
toshow no
color changes except,on
oneoccasion, a slight tendencyof the yellowground
colorto appeargrayish.The
Blue Sunfish (Lepomispallidas)which
isknown
alsoas Bluegill,Copper Nosed Bream,
BlueBream,
Black Sunfish,Roach (Sodus
Bay,New York),
etc.,shows
various color pat- terns.Sometimes
the fish will be a plain iron color withno markings
atall except the black spoton
the opercular Hap.At
othertimes itis a plain yellowish green withno markings
oritmay show
either of these colorswith faint or dark cross bars, or the lightground
colormay
he almost entirely obscured byheavy
dark bars withdark
mottlings betweenthem.
In Eftneacanthus gloriosus, a
common
sunfish of the vicinity ofWashington,
color changesmay
be notedwhich
arevery sim-ilar to those of Lepomiscyannellus.
The
changesfrom
vertical darkand
light barsto the colorationof bright specks is practi- callythesame
as intheGreen
Sunfish exceptthatinEnneacanthus theheavy
black stripe through the eye never,so far aswe know, becomes
entirely invisible.The War Mouth
(Chsenobryttusgulosus)shows
littlechange
in color except that the extent of the dark patches varies consid- erably so that thesame
fishmay
be grayish mottled with black or nearly all black with a few grayish streaks. Fish caughton hook and
line nearWashington
oftenshow much
red ordark orange in theground
color.Some
of ourcommon
fish,which
are frequently seen in aqua- riums, as theGar
Pike(Lepidosteus osseus)and
Strawberry Bass or Calico Bass (Pomoxis sparoidesand
Pomoxisannularis),appar- entlyshow no
changes exceptan
increase or reduction in the intensity ofthe coloration.74
Bean and Weed —Coloration of Fishes.
Preserved specimens of the Pickerel (Esox reticulatus)
show
very slight differences in color pattern,which
is usuallymore
orlessof a
network
ofdark
lineson
a lighter ground, butgreat differences in its visibility.These
differences are probably largelydue
to the condition of the fishwhen
takenfrom
the waterand
difference in manipulation in preserving the speci-men.
Fishesfrom
clear water usuallyhave
the color patternmore
pronounced, while thosefrom muddy
water, particularly water of a distinctly yellow color, frequentlyshow no
traces of the color patternwhen
first takenand
if killed in the preserva- tiveimmediately
willshow
none.However,
if these fish are placed for a fewminutes
in clear water the color patternbecomes
visible,and
itmay
appear in cold water even after the fish has died,showing
that the color cellsmay remain
activefora time
under
favorable conditions. Fishes of this speciesswimming
nearthe surface in theaquarium show
averystronglymarked
color pattern. Insome
ofthem
the reticulations extendentirely overthe back, while in others the backismore
or lessplain dark olivaceous.
When
these fish are restingon
abottom
of clear yellow sand thevertical bars of the reticula- tions tend to disappearand
portions ofthe horizontal bars alsobecome
fainter so that themarkings
resemblewave marks on
the bottom.The
colorof the backbecomes
plain greenishyel- low,exceptforthepresenceof scattered black specksresembling, in colorand
arrangement, black sand grains in the neighbor- hood.The
fish resting near the surface resembles a floating stick covered with spots of light refractedby
the small waves.In this species the color changes are not in the nature of a
change
in color pattern,but simply variations in its intensity.Inthe Yellow Perch (Percaflavescens) the color pattern does not change, but the intensity of the black cross bars is quite variable.
Sometimes
they are almost invisibleand
thefish ap- pearsto be aclear yellowcolor, whileat othertimes they are so broadand
intense that the fish appears almost black.The ground
colorchanges with thebands from
a light yellow to a dark yellowish gray.The
SeaRobin
(Prionotus evolans), the Puffer (Spheroidesmar
tdatus),and
the Filefish (Alutera scripta)show no change
in the color pattern, but considerable changesfrom
golden to grayish in theground
color.Specimens
of the SeaRobin
which had
buried themselves in the sand were light gray with black markings, while those that wereswimming
were various shadesofyellowand brown marked
with darker brown.Early in the
summer
of 1908 a small catfish (Amciurus sp.)was
taken in a dip net in theChesapeake and Ohio
Canal, atChain
Bridge, near Georgetown, D. C.When
first taken itwas
entirely black. Tins color persisted for several daysand was
then replaced by amost
unexpected pattern.The body was
black, butat the base of dorsal, adipose dorsal, analand
caudal finswas
anarrow
whiteband
so thateach fin appeared cutofffrom
the body,which seemed
to be a little pieceof black stick withsome
little pieces of rotted leaves (the black part of the fins) near it.The
caudal finwas
in reality truncate, but therewas
a large white patch in itwhich made
it appear that the upperand
lower rays were greatly produced. This color in turn, after a few weeks,was
replaced by the black color ofthe adult.Although many
notes on the coloration of fishes have been published they are scattered through the literature of every branch of ichthyology.We
offer the following partial listof themore
recent papers:1897
—
A. E. Verrill—
Nocturnal Protective Coloration of Fishes.Am.
Nat., Vol.
XXXI,
Feb., 1897, 99-103.Am.
Journ. Sci., Vol.Ill, 1897, 132-136. Abstractof paper read before
Am.
Morph.Soc, Dec. 30, 1896.
1905—D. S.
Jordan—
A Guideto theStudyof Fishes.New
York, 1905.Vol. I, pp. 226 to 236.
1907—Raveret-Wattel— " Le Pecheur" 18 anee No. 421, Sept. 30, 1907, p. 7(37. "Lacoloration des Poissons"
gives an account of the slow colorchanges in fishesunderthe influenceof great changes in color ofenvironment.
1908
—
FrancisWard —
Markings and colors in fish.How
they protect their wearers. Scientific American, Supplement No. 1714, p.297.
New
York, 1908.A
few notes on protective relation of color to surroundings in Stone Loach, Pike and Gudgeon.Experiments on Pike with head indark and bodyin light and
viceversa. This paperwas apparently published or offered for publicationin "
CountryLife," butwehave not founditinthat magazine.
76
Bean and Weed —
Coloration of Fishes.1909—C. Tate
Regan—
Proc. Zool. Soc. London, Feb. 16. 1909, p. 130.Gives notes on four species and criticises Dr. Evermann's note onspecies of Bodianus based on color in Fishes of Porto Rico, Bull.U. S. F.
C,
1900 (1902).1909—Jacob Reighard
— An
experimental studyof warning coloration in coral reef fishes. Carnegie Inst. Pub. 103, pp. 257-325 (1909).Experimentsinfeeding variously colored baits towildrisbes.
1909—0. H.
Townsend—
13th Ann. Rep. N. Y. Zool. Soc, Jan. 1909.Observationsoninstantaneous changes in color
among
tropicallislies.
1910—0. H.
Townsend— "The
Century Magazine," September, 1910.Chameleons of the Sea. Notes instantaneous color changes in
many
tropical risbes and slower ones in some northern ones.Practically a reprintof 13th Ann. Rept. N. Y. Zool. Soc,Jan., 1909, pp. 1-28.
1910
—
F. B. Sumner—
Adaptive colorchangesamong
hsbes. Bull.Zool.Soc. X. Y., No. 42,p. 699, 1910. (v>lorchanges of Pleuronecti- d.r. Fivephotographsofthe samehsh ondifferent bottoms.
1910—R. C.