OF WASHINGTON, VOLUME
XI, 19(19. 151NUPTIAL COLORS IN THE CHRYSOMELID^E.
[Coleoptera.]
BY
FREDERICK KNAB.Among
certain forms of Chrysomelidse theimago
takeson
an entirely different appearance with the attainment of sexual maturity. Eithernew
colors areadded
to the existing pattern or there is a complete change in coloration.These
colors are not acquired as the result of themore
or less gradual chiti- nizationand
pigmentationwhich
follows eclosion ("Aussfarb- ungsprozess" of theGermans)
hutmake
their appearance only with thedevelopment
of the sexual powers.This
matur- ing of the sexualpowers may
be a matter ofmonths and
oftenit only
comes
about with the advent ofanother season, formost
Chrysomelidae are single-broodedand
do notbecome
sex- uallymature
until after they have hibernated or aestivated.Perhaps it
would
bemore
exact to say that the physiological changeswhich
determine sexual activity take place during the resting period (hibernation or aestivation).One
of themost
strikingexamples
of color-transformation of this character has been studiedby
Caulfield in a northern species of Cassidini; its significance, however, appears to have been wholly overlooked.The
beetle in question isPhysonota
helianthi Randall.The
differencebetween
the parent beetlesand
thenew
generationproduced from them
is so great that Caulfield very naturally concluded the insect to be double-brooded and
dimorphic.* l>ut further observationdemon-
strated that this insect is single-broodedand
in the course of time completely changes its appearance.fThe
beetleswhich
appear in the spring or earlysummer
arebrilliant golden greenand
so theyremain
untilthey copulate, lay their eggs,and
final- ly die.The
offspring of these golden green parents are wholly different in appearance. Afteremergence
thepronotum and
elytra are shining
creamy
white with translucent marginal portionsand
with five black spotsupon
thepronotum
; soon, with the progressing chitinization, the elytra change to a dull blackish color with irregular ivory spotsand
blotchesupon
them. In this sober garb the beetlesmay
befound upon
their food-plant during the later part of thesummer and
in theautumn;
they feed but littleand show no
sexual desires,and
it is in this state that they go into hibernation.
When
the beetles reappear in the following spring they haveassumed
the*Can. Ent., vol. 16, p.227 (1884).
fCan. Ent., vol. 19, p. 73 (1887).
152
PROCEEDINGS ENTOMOLOGICAL
SOCIETYbrilliant green coloration of theirparents
and
theyare then act- iveand
intentupon
reproduction. Dr.Dimmock and
the writer have corroborated Caulfield's observationsby
rearing the beetlesfrom
the eggsand
carryingthem
through to the following season.Transformations of a similar character have been observed
in certain species of
European
Cassidini, but their correlation with the physiological changes of the reproductive period ap- pears never tohave
been suspected.A much less obvious case
is that of another
American
cassidid,Chelymorpha
argus Lich- tenstein. This beetle at first,shows
an ochreous yellow ground- color above,and
it is not until it issuesfrom
hibernationand
is ready to reproduce that it
shows
the characteristic dull red ground-color. Incidentally itmay
bementioned
that atWinnipeg,
Manitoba, the writerfound
a race ofChelymorpha
arguswhich
is ochreous yellowwhen
sexually mature.Similar changes occur in certain Chrysomelini
when
sexual maturity is acquired.A
striking case is that ofZygogramma
signatipennis Stal,
which
the writerfound abundant upon
a large composite plant at Cordoba, Mexico.During
the wintermonths
the beetles could befound
but sparinglyupon
their food-plantand
appeared tobe very inactive.At
that time the ground-color of the elytrawas
a dullcreamy
white.With
the advent ofwarm
weatherand
the rains the beetlesbecame abundant
; theywere now
activeand many
pairswere
observed in copulation.The
ground-color of the elytrahad now changed
to bright green-gold. In certain of our native species of Calligrapha the ground-color of the elytra is at first
creamy
white,and
thistakeson
a silvery lusterwhen
thebeetle"becomes sexually mature. In Calligrapharhoda Knab
the contrast isenhanced by
the addition of red to the elytral pattern with sex- ual maturity; this change takes place in a stillmore
strikingmanner
in Calligrapha roivenaKnab. As
these beetleshave
been described in a previous article itwould
be superfluous togo
into the details of coloration.Tower
in hisbook
on evolution in the genus Leptinotarsa,*proposes
names
for certainMexican forms
of this genuswhich
he claims to be new.He
states that these speciesshow
colors in life
which
disappear after the death of the insectand
that these colors are of specific value. Doubtless in these cases also the colors in question are correlated with the sexual functions, forin all the cases previouslycited,the nuptial colors*An
Investigation of Evolution in Chrysomclid Beetles of the Genus Leptinotarsa. (Pub. No. 48, Carnegie Institution of Washington, 1906.)OF WASHINGTON, VOLUME
XI, 1909. 153disappear with death, as they are said to
do
in these species of Leptinotarsa. UnfortunatelyTower
has not specifiedwhat
these colors areand how
they are distributed.Should
these colors prove to be ofthe physiological character indicated, per- sons studying these beetles at different seasonswould
naturally conclude that the differences in colorationwere
of specific value; or even thesame
individual, in ignorance of their trm- nature,would
be led into error.Cases of nuptial colors occur in certain species of Halticini, although they appear to be
much
less frequent in thisgroup
than in the Cassidini or Chrysomelini.A
conspicuousexample
is
Disonycha
quinque-vittata Say, acommon
speciesupon
wil- lows inmany
parts ofNorth
America.These
beetles,when found
in the latesummer
have pale yellow elytra withnarrow
black longitudinal stripes.When
the beetlesemerge from
hi- bernation the following spring the elytrashow
in addition a largeamount
of crimson, disposed in broad stripes in the intervalsbetween
the black lines.A Mexican halticid, Homo-
phccta abdoniiiialis Chevrolat,
which
the writerfound common
at Cordoba, develops nuptialcolors.
During
the wintermonths
these beetleswere
frequentlyfound upon
their food-plant.They
thenhad
dull bluish black elytra, while thepronotum and most
of the legswere
a pale ivory-yellow.When
the season of sexual activity arrived, with the rainsand
thewarm
weather, the
pronotum and
the legschanged
to a delicate rose-red, while the elytra
showed
a peculiar blue sparkle.No
indications of nuptial colors have beenfound
in thetwo
great groups ofCamptosomata and Eupoda. While
it is true that ourknowledge
of the biology of these groups is very in- complete, these groups,and
particularly the one firstmentioned, stand apart inmany ways and
one can hardly expect close correspondence in themore
specialized traits.What
doescause surprise isthat the Hispini,
which
are so intimately linked with the Cassidini,show none
of the remarkable color-changes so frequent in the latter group.But
thismay
be due wholly to our incomplete knowledge,and
perhaps,when
the tropicalforms
have been investigated, thisphenomenon
will befound among
the Hispini also.These
notes demonstratehow meager
is ourknowledge
of the biologyof this importantgroup
of beetles.Our
literature ofeconomic entomology abounds
withmore
or less extended notices of the injurious species, yet it is rare to find an original observation recorded.The
accounts are mostlyborrowed from
earlier authors,and
such additions as aremade, from
scattered recordsand
without critical knowledge, are,more
often than not, inaccurate