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No¡¡doubt¡¡many¡¡instincts¡¡of¡¡very¡¡difficult¡¡explanation¡¡could¡¡be¡¡opposed¡¡to¡¡the¡¡theory¡¡of¡¡natural¡¡selection£»¡¡cases£»¡¡in¡¡which¡¡we¡¡cannot¡¡see¡¡how¡¡an¡¡instinct¡¡could¡¡possibly¡¡have¡¡originated£»¡¡cases£»¡¡in¡¡which¡¡no¡¡intermediate¡¡gradations¡¡are¡¡known¡¡to¡¡exist£»¡¡cases¡¡of¡¡instinct¡¡of¡¡apparently¡¡such¡¡trifling¡¡importance£»¡¡that¡¡they¡¡could¡¡hardly¡¡have¡¡been¡¡acted¡¡on¡¡by¡¡natural¡¡selection£»¡¡cases¡¡of¡¡instincts¡¡almost¡¡identically¡¡the¡¡same¡¡in¡¡animals¡¡so¡¡remote¡¡in¡¡the¡¡scale¡¡of¡¡nature£»¡¡that¡¡we¡¡cannot¡¡account¡¡for¡¡their¡¡similarity¡¡by¡¡inheritance¡¡from¡¡a¡¡common¡¡parent£»¡¡and¡¡must¡¡therefore¡¡believe¡¡that¡¡they¡¡have¡¡been¡¡acquired¡¡by¡¡independent¡¡acts¡¡of¡¡natural¡¡selection¡£¡¡I¡¡will¡¡not¡¡here¡¡enter¡¡on¡¡these¡¡several¡¡cases£»¡¡but¡¡will¡¡confine¡¡myself¡¡to¡¡one¡¡special¡¡difficulty£»¡¡which¡¡at¡¡first¡¡appeared¡¡to¡¡me¡¡insuperable£»¡¡and¡¡actually¡¡fatal¡¡to¡¡my¡¡whole¡¡theory¡£¡¡I¡¡allude¡¡to¡¡the¡¡neuters¡¡or¡¡sterile¡¡females¡¡in¡¡insect¡communities£º¡¡for¡¡these¡¡neuters¡¡often¡¡differ¡¡widely¡¡in¡¡instinct¡¡and¡¡in¡¡structure¡¡from¡¡both¡¡the¡¡males¡¡and¡¡fertile¡¡females£»¡¡and¡¡yet£»¡¡from¡¡being¡¡sterile£»¡¡they¡¡cannot¡¡propagate¡¡their¡¡kind¡£¡¡
The¡¡subject¡¡well¡¡deserves¡¡to¡¡be¡¡discussed¡¡at¡¡great¡¡length£»¡¡but¡¡I¡¡will¡¡here¡¡take¡¡only¡¡a¡¡single¡¡case£»¡¡that¡¡of¡¡working¡¡or¡¡sterile¡¡ants¡£¡¡How¡¡the¡¡workers¡¡have¡¡been¡¡rendered¡¡sterile¡¡is¡¡a¡¡difficulty£»¡¡but¡¡not¡¡much¡¡greater¡¡than¡¡that¡¡of¡¡any¡¡other¡¡striking¡¡modification¡¡of¡¡structure£»¡¡for¡¡it¡¡can¡¡be¡¡shown¡¡that¡¡some¡¡insects¡¡and¡¡other¡¡articulate¡¡animals¡¡in¡¡a¡¡state¡¡of¡¡nature¡¡occasionally¡¡become¡¡sterile£»¡¡and¡¡if¡¡such¡¡insects¡¡had¡¡been¡¡social£»¡¡and¡¡it¡¡had¡¡been¡¡profitable¡¡to¡¡the¡¡community¡¡that¡¡a¡¡number¡¡should¡¡have¡¡been¡¡annually¡¡born¡¡capable¡¡of¡¡work£»¡¡but¡¡incapable¡¡of¡¡procreation£»¡¡I¡¡can¡¡see¡¡no¡¡very¡¡great¡¡difficulty¡¡in¡¡this¡¡being¡¡effected¡¡by¡¡natural¡¡selection¡£¡¡But¡¡I¡¡must¡¡pass¡¡over¡¡this¡¡preliminary¡¡difficulty¡£¡¡The¡¡great¡¡difficulty¡¡lies¡¡in¡¡the¡¡working¡¡ants¡¡differing¡¡widely¡¡from¡¡both¡¡the¡¡males¡¡and¡¡the¡¡fertile¡¡females¡¡in¡¡structure£»¡¡as¡¡in¡¡the¡¡shape¡¡of¡¡the¡¡thorax¡¡and¡¡in¡¡being¡¡destitute¡¡of¡¡wings¡¡and¡¡sometimes¡¡of¡¡eyes£»¡¡and¡¡in¡¡instinct¡£¡¡As¡¡far¡¡as¡¡instinct¡¡alone¡¡is¡¡concerned£»¡¡the¡¡prodigious¡¡difference¡¡in¡¡this¡¡respect¡¡between¡¡the¡¡workers¡¡and¡¡the¡¡perfect¡¡females£»¡¡would¡¡have¡¡been¡¡far¡¡better¡¡exemplified¡¡by¡¡the¡¡hive¡bee¡£¡¡If¡¡a¡¡working¡¡ant¡¡or¡¡other¡¡neuter¡¡insect¡¡had¡¡been¡¡an¡¡animal¡¡in¡¡the¡¡ordinary¡¡state£»¡¡I¡¡should¡¡have¡¡unhesitatingly¡¡assumed¡¡that¡¡all¡¡its¡¡characters¡¡had¡¡been¡¡slowly¡¡acquired¡¡through¡¡natural¡¡selection£»¡¡namely£»¡¡by¡¡an¡¡individual¡¡having¡¡been¡¡born¡¡with¡¡some¡¡slight¡¡profitable¡¡modification¡¡of¡¡structure£»¡¡this¡¡being¡¡inherited¡¡by¡¡its¡¡offspring£»¡¡which¡¡again¡¡varied¡¡and¡¡were¡¡again¡¡selected£»¡¡and¡¡so¡¡onwards¡£¡¡But¡¡with¡¡the¡¡working¡¡ant¡¡we¡¡have¡¡an¡¡insect¡¡differing¡¡greatly¡¡from¡¡its¡¡parents£»¡¡yet¡¡absolutely¡¡sterile£»¡¡so¡¡that¡¡it¡¡could¡¡never¡¡have¡¡transmitted¡¡successively¡¡acquired¡¡modifications¡¡of¡¡structure¡¡or¡¡instinct¡¡to¡¡its¡¡progeny¡£¡¡It¡¡may¡¡well¡¡be¡¡asked¡¡how¡¡is¡¡it¡¡possible¡¡to¡¡reconcile¡¡this¡¡case¡¡with¡¡the¡¡theory¡¡of¡¡natural¡¡selection£¿¡¡
First£»¡¡let¡¡it¡¡be¡¡remembered¡¡that¡¡we¡¡have¡¡innumerable¡¡instances£»¡¡both¡¡in¡¡our¡¡domestic¡¡productions¡¡and¡¡in¡¡those¡¡in¡¡a¡¡state¡¡of¡¡nature£»¡¡of¡¡all¡¡sorts¡¡of¡¡differences¡¡of¡¡structure¡¡which¡¡have¡¡become¡¡correlated¡¡to¡¡certain¡¡ages£»¡¡and¡¡to¡¡either¡¡sex¡£¡¡We¡¡have¡¡differences¡¡correlated¡¡not¡¡only¡¡to¡¡one¡¡sex£»¡¡but¡¡to¡¡that¡¡short¡¡period¡¡alone¡¡when¡¡the¡¡reproductive¡¡system¡¡is¡¡active£»¡¡as¡¡in¡¡the¡¡nuptial¡¡plumage¡¡of¡¡many¡¡birds£»¡¡and¡¡in¡¡the¡¡hooked¡¡jaws¡¡of¡¡the¡¡male¡¡salmon¡£¡¡We¡¡have¡¡even¡¡slight¡¡differences¡¡in¡¡the¡¡horns¡¡of¡¡different¡¡breeds¡¡of¡¡cattle¡¡in¡¡relation¡¡to¡¡an¡¡artificially¡¡imperfect¡¡state¡¡of¡¡the¡¡male¡¡sex£»¡¡for¡¡oxen¡¡of¡¡certain¡¡breeds¡¡have¡¡longer¡¡horns¡¡than¡¡in¡¡other¡¡breeds£»¡¡in¡¡comparison¡¡with¡¡the¡¡horns¡¡of¡¡the¡¡bulls¡¡or¡¡cows¡¡of¡¡these¡¡same¡¡breeds¡£¡¡Hence¡¡I¡¡can¡¡see¡¡no¡¡real¡¡difficulty¡¡in¡¡any¡¡character¡¡having¡¡become¡¡correlated¡¡with¡¡the¡¡sterile¡¡condition¡¡of¡¡certain¡¡members¡¡of¡¡insect¡communities£º¡¡the¡¡difficulty¡¡lies¡¡in¡¡understanding¡¡how¡¡such¡¡correlated¡¡modifications¡¡of¡¡structure¡¡could¡¡have¡¡been¡¡slowly¡¡accumulated¡¡by¡¡natural¡¡selection¡£¡¡
This¡¡difficulty£»¡¡though¡¡appearing¡¡insuperable£»¡¡is¡¡lessened£»¡¡or£»¡¡as¡¡I¡¡believe£»¡¡disappears£»¡¡when¡¡it¡¡is¡¡remembered¡¡that¡¡selection¡¡may¡¡be¡¡applied¡¡to¡¡the¡¡family£»¡¡as¡¡well¡¡as¡¡to¡¡the¡¡individual£»¡¡and¡¡may¡¡thus¡¡gain¡¡the¡¡desired¡¡end¡£¡¡Thus£»¡¡a¡¡well¡flavoured¡¡vegetable¡¡is¡¡cooked£»¡¡and¡¡the¡¡individual¡¡is¡¡destroyed£»¡¡but¡¡the¡¡horticulturist¡¡sows¡¡seeds¡¡of¡¡the¡¡same¡¡stock£»¡¡and¡¡confidently¡¡expects¡¡to¡¡get¡¡nearly¡¡the¡¡same¡¡variety£»¡¡breeders¡¡of¡¡cattle¡¡wish¡¡the¡¡flesh¡¡and¡¡fat¡¡to¡¡be¡¡well¡¡marbled¡¡together£»¡¡the¡¡animal¡¡has¡¡been¡¡slaughtered£»¡¡but¡¡the¡¡breeder¡¡goes¡¡with¡¡confidence¡¡to¡¡the¡¡same¡¡family¡£¡¡I¡¡have¡¡such¡¡faith¡¡in¡¡the¡¡powers¡¡of¡¡selection£»¡¡that¡¡I¡¡do¡¡not¡¡doubt¡¡that¡¡a¡¡breed¡¡of¡¡cattle£»¡¡always¡¡yielding¡¡oxen¡¡with¡¡extraordinarily¡¡long¡¡horns£»¡¡could¡¡be¡¡slowly¡¡formed¡¡by¡¡carefully¡¡watching¡¡which¡¡individual¡¡bulls¡¡and¡¡cows£»¡¡when¡¡matched£»¡¡produced¡¡oxen¡¡with¡¡the¡¡longest¡¡horns£»¡¡and¡¡yet¡¡no¡¡one¡¡ox¡¡could¡¡ever¡¡have¡¡propagated¡¡its¡¡kind¡£¡¡Thus¡¡I¡¡believe¡¡it¡¡has¡¡been¡¡with¡¡social¡¡insects£º¡¡a¡¡slight¡¡modification¡¡of¡¡structure£»¡¡or¡¡instinct£»¡¡correlated¡¡with¡¡the¡¡sterile¡¡condition¡¡of¡¡certain¡¡members¡¡of¡¡the¡¡community£»¡¡has¡¡been¡¡advantageous¡¡to¡¡the¡¡community£º¡¡consequently¡¡the¡¡fertile¡¡males¡¡and¡¡females¡¡of¡¡the¡¡same¡¡community¡¡flourished£»¡¡and¡¡transmitted¡¡to¡¡their¡¡fertile¡¡offspring¡¡a¡¡tendency¡¡to¡¡produce¡¡sterile¡¡members¡¡having¡¡the¡¡same¡¡modification¡£¡¡And¡¡I¡¡believe¡¡that¡¡this¡¡process¡¡has¡¡been¡¡repeated£»¡¡until¡¡that¡¡prodigious¡¡amount¡¡of¡¡difference¡¡between¡¡the¡¡fertile¡¡and¡¡sterile¡¡females¡¡of¡¡the¡¡same¡¡species¡¡has¡¡been¡¡produced£»¡¡which¡¡we¡¡see¡¡in¡¡many¡¡social¡¡insects¡£¡¡
But¡¡we¡¡have¡¡not¡¡as¡¡yet¡¡touched¡¡on¡¡the¡¡climax¡¡of¡¡the¡¡difficulty£»¡¡namely£»¡¡the¡¡fact¡¡that¡¡the¡¡neuters¡¡of¡¡several¡¡ants¡¡differ£»¡¡not¡¡only¡¡from¡¡the¡¡fertile¡¡females¡¡and¡¡males£»¡¡but¡¡from¡¡each¡¡other£»¡¡sometimes¡¡to¡¡an¡¡almost¡¡incredible¡¡degree£»¡¡and¡¡are¡¡thus¡¡divided¡¡into¡¡two¡¡or¡¡even¡¡three¡¡castes¡£¡¡The¡¡castes£»¡¡moreover£»¡¡do¡¡not¡¡generally¡¡graduate¡¡into¡¡each¡¡other£»¡¡but¡¡are¡¡perfectly¡¡well¡¡defined£»¡¡being¡¡as¡¡distinct¡¡from¡¡each¡¡other£»¡¡as¡¡are¡¡any¡¡two¡¡species¡¡of¡¡the¡¡same¡¡genus£»¡¡or¡¡rather¡¡as¡¡any¡¡two¡¡genera¡¡of¡¡the¡¡same¡¡family¡£¡¡Thus¡¡in¡¡Eciton£»¡¡there¡¡are¡¡working¡¡and¡¡soldier¡¡neuters£»¡¡with¡¡jaws¡¡and¡¡instincts¡¡extraordinarily¡¡different£º¡¡in¡¡Cryptocerus£»¡¡the¡¡workers¡¡of¡¡one¡¡caste¡¡alone¡¡carry¡¡a¡¡wonderful¡¡sort¡¡of¡¡shield¡¡on¡¡their¡¡heads£»¡¡the¡¡use¡¡of¡¡which¡¡is¡¡quite¡¡unknown£º¡¡in¡¡the¡¡Mexican¡¡Myrmecocystus£»¡¡the¡¡workers¡¡of¡¡one¡¡caste¡¡never¡¡leave¡¡the¡¡nest£»¡¡they¡¡are¡¡fed¡¡by¡¡the¡¡workers¡¡of¡¡another¡¡caste£»¡¡and¡¡they¡¡have¡¡an¡¡enormously¡¡developed¡¡abdomen¡¡which¡¡secretes¡¡a¡¡sort¡¡of¡¡honey£»¡¡supplying¡¡the¡¡place¡¡of¡¡that¡¡excreted¡¡by¡¡the¡¡aphides£»¡¡or¡¡the¡¡domestic¡¡cattle¡¡as¡¡they¡¡may¡¡be¡¡called£»¡¡which¡¡our¡¡European¡¡ants¡¡guard¡¡or¡¡imprison¡£¡¡
It¡¡will¡¡indeed¡¡be¡¡thought¡¡that¡¡I¡¡have¡¡an¡¡overweening¡¡confidence¡¡in¡¡the¡¡principle¡¡of¡¡natural¡¡selection£»¡¡when¡¡I¡¡do¡¡not¡¡admit¡¡that¡¡such¡¡wonderful¡¡and¡¡well¡established¡¡facts¡¡at¡¡once¡¡annihilate¡¡my¡¡theory¡£¡¡In¡¡the¡¡simpler¡¡case¡¡of¡¡neuter¡¡insects¡¡all¡¡of¡¡one¡¡caste¡¡or¡¡of¡¡the¡¡same¡¡kind£»¡¡which¡¡have¡¡been¡¡rendered¡¡by¡¡natural¡¡selection£»¡¡as¡¡I¡¡believe¡¡to¡¡be¡¡quite¡¡possible£»¡¡different¡¡from¡¡the¡¡fertile¡¡males¡¡and¡¡females£»¡¡in¡¡this¡¡case£»¡¡we¡¡may¡¡safely¡¡conclude¡¡from¡¡the¡¡analogy¡¡of¡¡ordinary¡¡variations£»¡¡that¡¡each¡¡successive£»¡¡slight£»¡¡profitable¡¡modification¡¡did¡¡not¡¡probably¡¡at¡¡first¡¡appear¡¡in¡¡all¡¡the¡¡individual¡¡neuters¡¡in¡¡the¡¡same¡¡nest£»¡¡but¡¡in¡¡a¡¡few¡¡alone£»¡¡and¡¡that¡¡by¡¡the¡¡long¡continued¡¡selection¡¡of¡¡the¡¡fertile¡¡parents¡¡which¡¡produced¡¡most¡¡neuters¡¡with¡¡the¡¡profitable¡¡modification£»¡¡all¡¡the¡¡neuters¡¡ultimately¡¡came¡¡to¡¡have¡¡the¡¡desired¡¡character¡£¡¡On¡¡this¡¡view¡¡we¡¡ought¡¡occasionally¡¡to¡¡find¡¡neuter¡insects¡¡of¡¡the¡¡same¡¡species£»¡¡in¡¡the¡¡same¡¡nest£»¡¡presenting¡¡gradations¡¡of¡¡structure£»¡¡and¡¡this¡¡we¡¡do¡¡find£»¡¡even¡¡often£»¡¡considering¡¡how¡¡few¡¡neuter¡insects¡¡out¡¡of¡¡Europe¡¡have¡¡been¡¡carefully¡¡examined¡£¡¡Mr¡¡F¡£¡¡Smith¡¡has¡¡shown¡¡how¡¡surprisingly¡¡the¡¡neuters¡¡of¡¡several¡¡British¡¡ants¡¡differ¡¡from¡¡each¡¡other¡¡in¡¡size¡¡and¡¡sometimes¡¡in¡¡colour£»¡¡and¡¡that¡¡the¡¡extreme¡¡forms¡¡can¡¡sometimes¡¡be¡¡perfectly¡¡linked¡¡together¡¡by¡¡individuals¡¡taken¡¡out¡¡of¡¡the¡¡same¡¡nest£º¡¡I¡¡have¡¡myself¡¡compared¡¡perfect¡¡gradations¡¡of¡¡this¡¡kind¡£¡¡It¡¡often¡¡happens¡¡that¡¡the¡¡larger¡¡or¡¡the¡¡smaller¡¡sized¡¡workers¡¡are¡¡the¡¡most¡¡numerous£»¡¡or¡¡that¡¡both¡¡large¡¡and¡¡small¡¡are¡¡numerous£»¡¡with¡¡those¡¡of¡¡an¡¡intermediate¡¡size¡¡scanty¡¡in¡¡numbers¡£¡¡Formica¡¡flava¡¡has¡¡larger¡¡and¡¡smaller¡¡workers£»¡¡with¡¡some¡¡of¡¡intermediate¡¡size£»¡¡and£»¡¡in¡¡this¡¡species£»¡¡as¡¡Mr¡¡F¡£¡¡Smith¡¡has¡¡observed£»¡¡the¡¡larger¡¡workers¡¡have¡¡simple¡¡eyes¡¡£¨ocelli£©£»¡¡which¡¡though¡¡small¡¡can¡¡be¡¡plainly¡¡distinguished£»¡¡whereas¡¡the¡¡smaller¡¡workers¡¡have¡¡their¡¡ocelli¡¡rudimentary¡£¡¡Having¡¡carefully¡¡dissected¡¡several¡¡specimens¡¡of¡¡these¡¡workers£»¡¡I¡¡can¡¡affirm¡¡that¡¡the¡¡eyes¡¡are¡¡far¡¡more¡¡rudimentary¡¡in¡¡the¡¡smaller¡¡workers¡¡than¡¡can¡¡be¡¡accounted¡¡for¡¡merely¡¡by¡¡their¡¡proportionally¡¡lesser¡¡size£»¡¡and¡¡I¡¡fully¡¡believe£»¡¡though¡¡I¡¡dare¡¡not¡¡assert¡¡so¡¡positively£»¡¡that¡¡the¡¡workers¡¡of¡¡intermediate¡¡size¡¡have¡¡their¡¡ocelli¡¡in¡¡an¡¡exactly¡¡intermediate¡¡condition¡£¡¡So¡¡that¡¡we¡¡here¡¡have¡¡two¡¡bodies¡¡of¡¡sterile¡¡workers¡¡in¡¡the¡¡same¡¡nest£»¡¡differing¡¡not¡¡only¡¡in¡¡size£»¡¡but¡¡in¡¡their¡¡organs¡¡of¡¡vision£»¡¡yet¡¡connected¡¡by¡¡some¡¡few¡¡members¡¡in¡¡an¡¡intermediate¡¡condition¡£¡¡I¡¡may¡¡digress¡¡by¡¡adding£»¡¡that¡¡if¡¡the¡¡smaller¡¡workers¡¡had¡¡been¡¡the¡¡most¡¡useful¡¡to¡¡the¡¡community£»¡¡and¡¡those¡¡males¡¡and¡¡females¡¡had¡¡been¡¡continually¡¡selected£»¡¡which¡¡produced¡¡more¡¡and¡¡more¡¡of¡¡the¡¡smaller¡¡workers£»¡¡until¡¡all¡¡the¡¡workers¡¡had¡¡come¡¡to¡¡be¡¡in¡¡this¡¡condition£»¡¡we¡¡should¡¡then¡¡have¡¡had¡¡a¡¡species¡¡of¡¡ant¡¡with¡¡neuters¡¡very¡¡nearly¡¡in¡¡the¡¡same¡¡condition¡¡with¡¡those¡¡of¡¡Myrmica¡£¡¡For¡¡the¡¡workers¡¡of¡¡Myrmica¡¡have¡¡not¡¡even¡¡rudiments¡¡of¡¡ocelli£»¡¡though¡¡the¡¡male¡¡and¡¡female¡¡ants¡¡of¡¡this¡¡genus¡¡have¡¡well¡developed¡¡ocelli¡£¡¡
I¡¡may¡¡give¡¡one¡¡other¡¡case£º¡¡so¡¡confidently¡¡did¡¡I¡¡expect¡¡to¡¡find¡¡gradations¡¡in¡¡important¡¡points¡¡of¡¡structure¡¡between¡¡the¡¡different¡¡castes¡¡of¡¡neuters¡¡in¡¡the¡¡same¡¡species£»¡¡that¡¡I¡¡gladly¡¡availed¡¡myself¡¡of¡¡Mr¡¡F¡£¡¡Smith's¡¡offer¡¡of¡¡numerous¡¡specimens¡¡from¡¡the¡¡same¡¡nest¡¡of¡¡the¡¡driver¡¡ant¡¡£¨Anomma£©¡¡of¡¡West¡¡Africa¡£¡¡The¡¡reader¡¡will¡¡perhaps¡¡best¡¡appreciate¡¡the¡¡amount¡¡of¡¡difference¡¡in¡¡these¡¡workers£»¡¡by¡¡my¡¡giving¡¡not¡¡the¡¡actual¡¡measurements£»¡¡but¡¡a¡¡strictly¡¡accurate¡¡illustration£º¡¡the¡¡difference¡¡was¡¡the¡¡same¡¡as¡¡if¡¡we¡¡were¡¡to¡¡see¡¡a¡¡set¡¡of¡¡workmen¡¡building¡¡a¡¡house¡¡of¡¡whom¡¡many¡¡were¡¡five¡¡feet¡¡four¡¡inches¡¡high£»¡¡and¡¡many¡¡sixteen¡¡feet¡¡high£»¡¡but¡¡we¡¡must¡¡suppose¡¡that¡¡the¡¡larger¡¡workmen¡¡had¡¡heads¡¡four¡¡instead¡¡of¡¡three¡¡times¡¡as¡¡big¡¡as¡¡those¡¡of¡¡the¡¡smaller¡¡men£»¡¡and¡¡jaws¡¡nearly¡¡five¡¡times¡¡as¡¡big¡£¡¡The¡¡jaws£»¡¡moreover£»¡¡of¡¡the¡¡working¡¡ants¡¡of¡¡the¡¡several¡¡sizes¡¡differed¡¡wonderfully¡¡in¡¡shape£»¡¡and¡¡in¡¡the¡¡form¡¡and¡¡number¡¡of¡¡the¡¡teeth¡£¡¡But¡¡the¡¡important¡¡fact¡¡fo