the life history inbreedingcages. Dr.
Howard
thoughtthatMr.
Simpson's
method
of observationand
recordwas most
surelycal culated tofurnish the data forcorrect conclusions.The
final paperof theeveningwas
by Prof.Cook, and
entitled:
THE EARWIG'S FORCEPS AND THE PHYLOGENY OF INSECTS.
By O.
F.COOK.
The
earwigs areagroup
oftropical insectswith very few rep resentatives in temperateregions.But
in spite of their retiring habits theyhave received a considerableamount
ofpopular atten tion,becauseof the fearinspired by the rather formidable pair of forceps carried at the end of the body. Entomologistsknow,
however,that the creatures are quite harmless,and
that their forceps are not only free from any poison glands,but are not suf ficiently strong tomake
awound
or puncture. In fact, no ade quate explanation ofthefunction of the forcepsseems to exist,as evidencedby
the followingsummary
of the scientificknowledge
of the subjectby
Dr. Sharp, the eminent entomologist of the BritishMuseum.
" The
pair of forcepswithwhich
thebody
isarmed
at itsex tremity forms another character almost peculiar to the earwigs, butwhich
exists in the genusJapyx
of the Thysanura.These
forceps varymuch
in the different genera of the family; they sometimes attain a large sizeandassume
veryextraordinary and distorted shapes.They
are occasionally used by the insectsas ameans
of completing the process of packingup
the wings, butin
many
species itis not probable that they can be used for this purpose, because theirgreat size and peculiarly distorted forms renderthem
unsuitable forassisting in a delicate process of arrangement; theyare, too, always present in thewingless forms of the family. Their importancetothecreatureisatpresent quite obscure;
we
can onlycompare them
with the horns of lam-ellicorn Coleoptera,
which
have hithertoproved inexplicable, as far asutilityisconcerned.No
doubtthecalipers of the earwigs givethem
animposing
appearance,and
itmay
beofsome
little advantageon
this account; they are notknown
to be usedas offensive instruments forfighting, but they are occasionally broughtintoplayforpurposesof defence, the creaturesusing
them
for the infliction of nips, which, however, are
by
nomeans
of a formidable character."*For
at leastonemember
ofthisgroup
thisdeficiency ofknowl
edge can be supplied by the fact that an earwig supposed to be*The
CambridgeNatural History, V, p.208, 1895.WASHINGTON.
Labia minor
Scudder,common
in the vicinity ofWashington,
usesits forceps to spread its wings,and
is apparently unableto resort to flightwithout theassistance of itscaudal armature.It is well
known
to entomologists that the hindwings
of the earwigdiffer from those of all other insects, except those of the Staphylinid beetles, in being folded transversely to fitunderthe short anterior wings,which
serve merely as protective covers.The method
of folding is, however, entirely different in thetwo
groups, and the suggestion of the above quotation that the ear-wiguses
the forcepsto fold thewings seem
tobe quite erroneous.It
was
probablyborrowed
by analogy from the beetles,where
the flexibleabdomen
is used, as itwere, to tuckthewings
undertheircovers.
The wings
of the Staphylinidas are stillexpanded when
the insect alights, and are sometimes allowedto remain sowhen
neglected through fright or annoyance.With
the earwig, on the contrary, the closing of thewings
is instantaneous and apparently quite automatic;when
the insect alights itswings
arecompletely folded,and
it runsaway
withoutthe necessity ofany of the preliminarycontortions of the Staphy-
linidae.
Moreover,
unlike the beetle, the earwig does notopen
its
wings
readily orwhen
running.The
operation is obviously a special effortwhich
requires it to stand stilland
exert its undi vided attention for averyappreciable interval.The wing
coversand wings
are soon raised from the body, but the wings do not unfold until, by repeated quickupward movements
of the re curvedabdomen
and forcepstheyare, as it were,combed
outand
spread forflight. Occasionally an earwig seems tolosethepower
ofkeepingthe
wings
open, and repeatedly fallsdown
after short flights ofan inch or two, though apparentlymaking
efforts atlonger journeys. Instead ofdirect flights, they oftenrise
and
con tinue to gyrate in a spiral abouttwo
inches in diameter.It is, of course, possible that this observation
would
not applyto all the
winged
earwigs, and,tojudge from thepast,much
time will be needed for its verification in the different families and generaof thegroup.That
the present use of the forcepsremained
so long unnoticed is probably to be explained by the factthatthe earwigs, like the termites, are nocturnal or twilight insects,
and when
disturbed in the daytime never attempt to use theirwings, but run for shelterand
concealment inthe dark.The
insectswhich
wereseen toflyhad
beenattracted to a light in the even ing, and therealso flightseem-ed tobe undertaken onlywhen
the creatures feltthemselves atleisure.When annoyedor frightened
theyattempted only to run away
the faster. Several genera of
earwigs, including Apachya,
collected under similar circum
stances in Liberia, were
also seen to alight with theirwings
alreadyfolded,and
tobend
theirabdomens
whileresuming
flight,
though the natureand object of the
movements were
notthen ap-predated or noted in detail.
The
complexity of thewingfold is such, however, that the needof an accessory organ like the for ceps appears by nomeans
improbable, and in the absence ofany
other suggestion of general pertinence, itseems not unwarranted to proceed on the assumption that the primary use of the forceps of theearwig is the unfolding of the wings.It is, of course, tobe expected thatsuch organs asthe forceps
would
beutilized inother ways, though the failureofentomolo gists to discover such secondaryfunctionsmay
be taken as anindication that no very extensive adaptation has taken place.
The most
that can be said at present is that both in theearwigand
inJapyx
the long, slender hairs withwhich
the forceps are sparingly clothed arean indicationthatthetactilesensibility resid ing in the styletsofmany
insectsisat least partially retained.. Insome
genera the forceps havebecome
enlargedand
thickened to anextent strongly suggestive ofadefensiveuse,and
itisof further interest to note that such forms are often wingless, and that the broadeningand
thickening oftheabdomen
tends to diminish the flexibilitywhich
is retainedby themore
slender formand
laxer skeletal structure of thewinged
genera.The
unusually broadabdomen
of thewinged
African genusApachya
is an exception to this rule, but here flexibility is provided forby the extreme thinness ofthis part of thebody, whilsthe forceps areso peculiar as to suggest the existence ofsome
unique adaptation.There
also exist
winged
species with robust bodies and strong forceps, but, likemany
beetles, thesemay make
no use of their wings.Indeed, it iseasy to understand theevident tendency toward the
abandonment
of so specialized and difficult, and at thesame
time so relatively unnecessary an activity as flying seemsto beamong
the earwigs.
With
the earwigs, as with the termites, the wings probably serve thesingleimportantpurposeofcross-fertilization, interbreeding, or panmixia,which
conduces atonce to organic vigorand
to evolutionary progress.But owing
to theirmore
activehabits
and
theirfreedom from socialorganization and caste specialization, thepower
offlightisofmuch
less vital importance to the earwigs than to the termites, and although the latter use theirwings
forbuta singleflightallsexual individuals arewinged, whilemany
genera of earwigs long sinceabandoned
flight alto gether.The
existence ofsomany
wingless earwigs is not, however, anargument
against the use of the forcepswith thewings, noragainst theadequacy
of such an explanation of the evolutionary origin and universal presence of the forceps in thisgroup
of insects.Such
an objection could be maintained only on the theory that the ancestral earwigwas
wingless, and thatwings
have been in dependently developed by differentgenera of earwigs, a positionwhich nobody
is likely to maintain. Itis appreciated, however,thatthe failure ofany of thewingless earwigs to lose the forceps
is not in accord with
commonly
accepted evolutionary theories.Instead of dropping the forceps with the wings, the caudal ap pendages have, in
some
cases, apparently increased in size,and
have certainly continued to differentiate in form, attaining, for example, amarked asymmetry
in the winglessgenus Anisolabis, a conditionwhich
could not possibly have a functional signifi cance in connection with the wings,and
in all probability has none in any other relation.The
continued presence of the for ceps in the wingless earwigs is, moreover, paralleledasanevolu tionaryphenomenon by
the equally useless multiplicity of formwhich
appears in the forceps of thewinged
genera.The
func tion here ascribedto theorgans in question renders it highlyim
probable that there is theslightest use in the differencesof form,size,
and
armature of the forceps,and
the great variabilityof these characters also forbids the supposition thatany
definitely specialized usesremainunknown. Both
sexes ofLabia minor
use theirforceps inthe
same manner,
though theform
of theap
paratus isvery different,and
it is nearlytwice aslonginthemale
as in thefemale. This is probably oneof the endlessly
numerous
secondary sexual differences having no direct use, but perhaps serving an important purpose in contributing to the diversitywhich many
organisms maintain inside specific lines.The
general maintenance of a direct proportion
between
the length of theabdomen and
the length of the forceps* supports theview
that natural selection hastended merely tokeep the forceps long
enough
toreachback
to the wings.The
field of biology abounds, however, in similarphenomena which
appearto beanomalous and
mysteriouswhen viewed
from exclusively selectionalor static theories of evolution,butwhich
itseemspreferable to interpretas
examples
of a generallaw
of bio logicalchange
for itsown
sake, as it were,and
independent of natural selection.!As
accessories of the organsofflight the forcepsare,of course, to be lookedupon
asan adaptation, butfromwhat? Presumably
from thejointed stylets to be found in somany
groups, butmore
particularly from such as those of the peculiarinsect described
by Westwood,
underthename Dyscritina^
but subsequentlyre ported to be the larva of an earwig. Dyscritina,which
the writerhas observedand
collected in Liberia,may
besaid tocom-
*Inafewcases whereslender species haveshortforceps, the abdomen seemstobeunusuallyflexible,butingeneral the longforcepsgo with the longbodies.
tA. Kinetic Theory of Evolution, Science, N. S., XIII. No. 338, pp.969-978,June21, 1901.
%Trans. Ent. Soc. London, 1881,p. 601. PI. xxii, Figs.l-li.
bine the habits of the earwigs with those of the true Thysanura.
It lives
among
vegetable debris invery moist places, is extremely quick andagile in itsmovements,
is verysoftand delicate in tex ture,and
is provided witli a pair of long, many-jointed stylets instead of the forceps of the adult earwig. Curiously enough, these are exactly the differenceswhich
obtainbetween
Japyx,the only other insectwith forceps like the earwig, and the smallerand more
thysanuroidanalogueofCampodea
alsofoundinLiberiaand
described before this Society in 1899, under thename Pro japyx
* RecentlyProjapyx has been found to benotuncommon
in Porto Rico,
and
seems always tooccur in thesame
localities as the PortoRican
species of Japyx, thatis, in all sorts ofsitua tions from moistvalleys to the tops ofdry limestone hills.But
if Projapyx is really the larva of Japyx, students of the temperate species have failed of their full duty, orelsewe
have found repeated in this orderf the strange conditions of the ear wigswhere some
species havealarval stageand
ametamorphosis which
have been suppressed in the others.Itis realized that this reasoning reverses, for the present case, atleast, theopinion held
by Lubbock and
others that the larval stages of insects are derivative and adaptative, not ancestral and primitive.No
reason is, however, apparentwhy
one ofthese opinions should exclude the other,
and
in the present in stance it seems obvious that from the standpoint ofhexapod
structure Dyscritinaismuch
less specialized than the adultearwig, or than the larva of theHymenoptera
or Lepidoptera.If, instead of holding that the Dyscritina stage of the
earwig
is an adaptation,
we
interpret it as amore
primitive condition,it willbe necessary to apply the
same
reasoning to the orthop- terous groups towhich
the earwigs have been thought to be closely allied.Most
of the cockroaches,like mostof the earwigs, have nopronounced
metamorphosis, but in several genera thereisa transformation in both sexes, while in others onlythe males reach the
winged
condition.Metamorphosis
ismore
accentuatedamong
the cockroaches thanamong
the remaining orders of Orthoptera, so that the current opinion that these insects are primitive because they have nometamorphoses
is self-contra dictory.Having
thusemancipated ourselves from the notion that the cockroachesrepresent the primitive insect type, it will beeasier toappreciate at its proper value the long-obvious probability*Proc. Ent. Soc. Washington, iv.222, 1899.
t
A
separate order Dicellurawas established for Japyx in 1896, forthe reason that it seemed more remote from the true Thysanura than thelatter are from the Orthoptera. See Brandtia, p.49.July30, 1896; also Proc. Ent. Soc. Washington, iv, 222, 1899.
OF 89
that the ancestral distinctionbelongstothe so-calledneuropterous orders with aquatic larva?
and
completemetamorphoses. Dys-
critina asthe larva of an earwig,
and
Projapyx as the larva of Japyx, bring the orthopterous series closer to the aquatic larvae of the Neuroptera, with their many-jointed stvlets,and
thus per mit us to think of the archetypal insect as a creature withmeta
morphosisand
withwings
instead ofbeginning with a thysanuran ancestor and being compelledto imaginethewings
as appearingkt independently at several points" as maintained
bv
Professor Smith.*For
the correctnessof this view, that the neuropterous orders with aquatic larvae are themore
primitive,! a largeamount
of evidencemight
be broughttogether, but perhaps themost
con spicuous advantageof this standpointlies in the factthat it per mits the suggestion of an originand method
ofdevelopment
for the insect wing,which
can scarcely be accounted forby any
rational evolutionary theory beginning with the assumption that the first insects
were
land animals.The wings
of the birds, pterodactyls, bats, fishes,and
other flyinganimals, areknown
to be modifications oforgans usedfor locomotion in water or on land, butwe
have been contented toassume
that the wings of insectswere made,
so to speak,from whole
cloth,and
have failedtoassociatethem
asthe homological equivalents and derivativesof older structures used for purposes otherthan flight.Kinetic evolution views as
normal
the progressivechange
of any particular part, butwould
not prearrangeand
carry forward thecomplex and
delicate adjustmentsof structureand
function necessary tothe perfection ofsuch organsaswings, since,except for flying,wings
areabout as useless structures for terrestrial in sects as could well be imagined,and some
representatives of nearly all theorders haveabandoned
them.But
ifthe discussion be transferred to the water,we
have, soto speak,
much
clearer sailing. Finstoosmall for flyingarestillvery useful to fish, anda gradual
and
natural increase of size ofswimming
organs to the pointwhere
theycan be used forflight is illustratedby
theanalogy of the flying fishes.J*
An
Essay on the Classification of Insects, Science, N. S., v, p'. 671, April 30. 1897.tThe
copulatorj apparatus of the Odonata, locatedonthesecond seg mentof theabdomen, is paralleled only in the Diplopoda.The
paired genital openings of the Ephemerida and the moulting of the insect in adultform are,if possible, evenmoreprimitivefeatures.I
Some
of the so-called flyingfish merelysoar for short distances on theirexpanded wings, but othersarecapable oftrueflight,notbyflapping theirwingslikebirds,but bv keepingthemin a state of very rapid vibraThe
lackofmeans
ofaerial existenceandoflocomotionon land has kept the fishfrom
terrestrial conquests, but aquatic insects are notthus restricted,and
thousands ofspecies, includingmem
bers of
many
different orders, arestill able tomake
the ancestral substitution of a terrestrial for an aquatic habitat.Many
alsoremain in the water asadults,
and
use theirwings
forswimming.
Some
of the May-flies descend into thewater to lay their eggs,and
inthegenusPteronarcys
the adultwinged
insecthas external gills.The most
conspicuous suggestion for the formation ofwings
from gills is, perhaps,tobe foundin thelarvaeof theMay
flies,
where
the gills havebecome
subdorsal and the tracheae which, in other groups,hang
in brush-like clusters, are spread out instead asveins ofdelicate, leaf-likemembranes, and
even ar ranged inamanner
stronglysuggestive of the patterns of thewings
ofsome
adult insects of other groups.The
utilization of the anterior pairs ofsuchlamillargillsasswimming
organs,andtheir subsequent further specializationaswings, is thus a supposition requiringno abruptorimprobablechange
of structure or function,and
affordsa rational explanation oforgans otherwiseas mysteri ous morphologicallyas thewings
of angels.So much
for theargument
affordedby
thewinged
and wingless earwigs, and the similarity of the jointed stylets of the earwig larvastothose ofProjapyxand Campodea.
Shortlyafterwriting this sketch of phylogenetic possibilities, I received from Dr.Filippo Silvestri,* of
Bevagna,
Italy, a paper inwhich my meagre
accountof theanatomy
of theAfrican Projapyx isgreatly extended by observationson
a SouthAmerican
species. Dr.Silvestri not onlyagrees with
me
that Projapyx isthemost prim
itive of insects,but heholds inadditionthat it provesthe descent of theinsects
from
the diplopods,because hefindsthat the jointed styletsare spinning organshomologous
with those of Scolopen- drcllaand
with those of the diplopod orders Crelochetatand Monocheta. But
ifProjapyx is the larva ofJapyx
instead of a mature insect, Dr. Silvestri's reasoningmust
be reversed, andwe
should prepare ourselves to believe that theSymphyla,
Dip- lopodaand Pauropoda
do not represent the ancestors of the hex-tionthrough a small arc, like the insects.
The
distances traversed are toogreat,and therateofspeed tooslowandtoo uniform tobe explained bythemomentum
withwhichthefishleaves the water. Thisconclusionistheresultof
many
excellent opportunities of observation within thelast tenyears,principallyinthe Cape Verderegion of theAtlantic. The ob jection ofsome ichthyologists that theflyingfish isnotsoconstructedas tobe abletovibrate its fins in theairwouldalsorender theseorgansuse less in the water.* Boll. Mus. Zool. Anat.Comp. Univ. Torino,No. 399, Sept. 12, 1901.
t Brandtia,p. 41, 1896.
apods, butare, as it were, larviform off-shoots
from
the insectphylum.
In other words,we may compare
the diplopods with caterpillarsand
other larvae,and may
seriously undertake the study necessaryto determine the realityofwhat
have been sup posed to be merely superficial similarities, such as the form of the cephalic sclerites, the barbed hairsand
the repugnatorial pores.Polyxenus
looksenough
like a caterpillar,and
its large fossil relativePalceocampa would
have been evenmore
strongly suggestive of such an affinity.*Moreover,
ahexapod
origin for theDiplopoda would
explain thefactthat thediplopod larvas are hatched with the six anterior legs, the remaining pairs being attached to rings intercalated behind the genital segment.The anamorphous
Chilopodaarehatched with seven pairs oflegs, but the others areaddedby
intercalation in front of the genital seg ment,which
thus appears near the posteriorend
of thebody
in theonegroup and
near the anterior in theother.In this
way
itis possible to bridgethechasm which seemed
to so profoundly separate the Progoneata (Diplopoda,Symphyla, and Pauropoda)
from the Opisthogoneata(Hexapoda
and Chilo-*Thebarbedhairsof thelarvaaoftheMerocheta, andthebristlesof the Coelocheta, Monochetaand Colobognatha support theviewthat thesoft- bodied, hairyPolyxenus isthemostprimitive of existingdiplopod types.
It
may
alsobesaidthattheskeletons of thedifferentorders ofDiplopodaaretoo diverseto.be rationallyexplained bydescent from a single hard- bodied type. In the Merocheta the segmental rings aresolid and
com
plete,without eventracesof sutures torepresent pleural or ventralplates.
In the Coelocheta, Monocheta, and Colobognatha the ventral plates are free; in the Diplocheta,Zygocheta, and Anocheta they are adnate, but are distinctbysutures.
The
pleurae are free in the Oniscomorphaand Limacomorphaand inthe Siphonotidoe; adnate in the Polyzonidse and remaining Colobognatha, andin theMonochetaand Anocheta; notraces of pleural elements have been reported in the Diplocheta,Zygocheta, Coelochetaand Merocheta. Finally, in the order Anocheta, the dorsal partof thesegmental ringis composedof threetransverse bands,acon dition perhaps paralleledonly in the larvae of the saw-flies.From
the entomological standpoint thesedifferenceswould bethoughtvery grave;indeed, they
maybe
said tobe altogether too graveforexplanationby evo lutionarychangesin partsalready hardened. If,however,wethink ofthemasindependentacquisitions offirmarmorbysoft-skinned animalswehave ampleanalogiesinother groups. Thisinterpretation doesnot,of course, decrease the actual diversity, butitenables us to credit the evidence of the otherwise great similarity of structure and function
among
the Diplopoda. Itpermits Polyxenustobemoreclosely associated with the other Diplopoda, and brings the Diplopoda,as a group, closer to the Symphylaand tothe Hexapoda.poda).
That
there could be any derivative relationshipbetween
a
group
ofanimals with the reproductive system opening in the anterior part of thebody
(Progoneata)and
others withthe reverse arrangement ^Opisthogoneata)seemed
impossible, unless thecommon
originwere
traced back toworms
with unspecialized reproductive segments.Should
the present suggestion prove to have a foundation in fact,we
shall havefirmerground
forbeliev ing that the insectsand
the four classescommonly grouped
as44
Myriapods"
do in reality constitute a natural assemblage.*Nor do
the possibilitiesof integration end here, sincethe asso ciation of the Progoneata with the insects as derivatives of an originallyaquaticgroup
reopensthewhole
question oftheiraffini ties with the Crustaceaand
Arachnida, andmay
result in therehabilitation of the
Arthropoda
as a naturalphylum
or primary division of the animalkingdom.
The
paperwas
discussed brieflyby
Messrs.Simpson,
Gill and Stiles.Mr. Simpson
said that Prof.Comstock
had given seriousconsideration to the theorythat the primitiveand
originalinsects
were
aquaticand
winged, buthad
finallyabandoned
itas untenable.Ke had noted that the tracheal gills of certain May
flies areof almost the
same
pattern as thewing
veins. Thissimilarity, however, he
had
foundtobeaccidental. Dr.Gill said he thoughtthattherewas
noground
forbelievingthat insectswere
derivedfrom myriapods.There was
a possibility, however,that they hadsprung
from an entomostracantype, though thiswas
merely anhypothesis. Insectsmay
haveexisted inpaheozoictimes, but therewas
no evidence of it.He
maintained that flying-fish do not have atrue flight, but that theirenormously enlarged pec toral finsserve merely as parachutes for their sustentation inthe air until the initialmomentum
of the fish leaving the water isexhausted.
Prof.
Cook,
however, maintainedthat certain of theflying-fish really progress through the air by a rapid vibration of the wings.He
spoke of the resemblance ofsome
geologic Crus tacea to theIarva3 of certain species of cockroaches as of inter estand
suggestive.*For this the name Labrata was proposed in 18^6 (Brandtia, p.30).
heLabrata weredeemedco-ordinate with tl
Crustacea)and theMalacopoda (Peripatus).