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ghter¡­coloured¡¡than¡¡those¡¡confined¡¡to¡¡cold¡¡and¡¡deeper¡¡seas¡£¡¡The¡¡birds¡¡which¡¡are¡¡confined¡¡to¡¡continents¡¡are£»¡¡according¡¡to¡¡Mr¡¡Gould£»¡¡brighter¡­coloured¡¡than¡¡those¡¡of¡¡islands¡£¡¡The¡¡insect¡­species¡¡confined¡¡to¡¡sea¡­coasts£»¡¡as¡¡every¡¡collector¡¡knows£»¡¡are¡¡often¡¡brassy¡¡or¡¡lurid¡£¡¡Plants¡¡which¡¡live¡¡exclusively¡¡on¡¡the¡¡sea¡­side¡¡are¡¡very¡¡apt¡¡to¡¡have¡¡fleshy¡¡leaves¡£¡¡He¡¡who¡¡believes¡¡in¡¡the¡¡creation¡¡of¡¡each¡¡species£»¡¡will¡¡have¡¡to¡¡say¡¡that¡¡this¡¡shell£»¡¡for¡¡instance£»¡¡was¡¡created¡¡with¡¡bright¡¡colours¡¡for¡¡a¡¡warm¡¡sea£»¡¡but¡¡that¡¡this¡¡other¡¡shell¡¡became¡¡bright¡­coloured¡¡by¡¡variation¡¡when¡¡it¡¡ranged¡¡into¡¡warmer¡¡or¡¡shallower¡¡waters¡£¡¡
When¡¡a¡¡variation¡¡is¡¡of¡¡the¡¡slightest¡¡use¡¡to¡¡a¡¡being£»¡¡we¡¡cannot¡¡tell¡¡how¡¡much¡¡of¡¡it¡¡to¡¡attribute¡¡to¡¡the¡¡accumulative¡¡action¡¡of¡¡natural¡¡selection£»¡¡and¡¡how¡¡much¡¡to¡¡the¡¡conditions¡¡of¡¡life¡£¡¡Thus£»¡¡it¡¡is¡¡well¡¡known¡¡to¡¡furriers¡¡that¡¡animals¡¡of¡¡the¡¡same¡¡species¡¡have¡¡thicker¡¡and¡¡better¡¡fur¡¡the¡¡more¡¡severe¡¡the¡¡climate¡¡is¡¡under¡¡which¡¡they¡¡have¡¡lived£»¡¡but¡¡who¡¡can¡¡tell¡¡how¡¡much¡¡of¡¡this¡¡difference¡¡may¡¡be¡¡due¡¡to¡¡the¡¡warmest¡­clad¡¡individuals¡¡having¡¡been¡¡favoured¡¡and¡¡preserved¡¡during¡¡many¡¡generations£»¡¡and¡¡how¡¡much¡¡to¡¡the¡¡direct¡¡action¡¡of¡¡the¡¡severe¡¡climate£¿¡¡for¡¡it¡¡would¡¡appear¡¡that¡¡climate¡¡has¡¡some¡¡direct¡¡action¡¡on¡¡the¡¡hair¡¡of¡¡our¡¡domestic¡¡quadrupeds¡£¡¡
Instances¡¡could¡¡be¡¡given¡¡of¡¡the¡¡same¡¡variety¡¡being¡¡produced¡¡under¡¡conditions¡¡of¡¡life¡¡as¡¡different¡¡as¡¡can¡¡well¡¡be¡¡conceived£»¡¡and£»¡¡on¡¡the¡¡other¡¡hand£»¡¡of¡¡different¡¡varieties¡¡being¡¡produced¡¡from¡¡the¡¡same¡¡species¡¡under¡¡the¡¡same¡¡conditions¡£¡¡Such¡¡facts¡¡show¡¡how¡¡indirectly¡¡the¡¡conditions¡¡of¡¡life¡¡must¡¡act¡£¡¡Again£»¡¡innumerable¡¡instances¡¡are¡¡known¡¡to¡¡every¡¡naturalist¡¡of¡¡species¡¡keeping¡¡true£»¡¡or¡¡not¡¡varying¡¡at¡¡all£»¡¡although¡¡living¡¡under¡¡the¡¡most¡¡opposite¡¡climates¡£¡¡Such¡¡considerations¡¡as¡¡these¡¡incline¡¡me¡¡to¡¡lay¡¡very¡¡little¡¡weight¡¡on¡¡the¡¡direct¡¡action¡¡of¡¡the¡¡conditions¡¡of¡¡life¡£¡¡Indirectly£»¡¡as¡¡already¡¡remarked£»¡¡they¡¡seem¡¡to¡¡play¡¡an¡¡important¡¡part¡¡in¡¡affecting¡¡the¡¡reproductive¡¡system£»¡¡and¡¡in¡¡thus¡¡inducing¡¡variability£»¡¡and¡¡natural¡¡selection¡¡will¡¡then¡¡accumulate¡¡all¡¡profitable¡¡variations£»¡¡however¡¡slight£»¡¡until¡¡they¡¡become¡¡plainly¡¡developed¡¡and¡¡appreciable¡¡by¡¡us¡£¡¡
Effects¡¡of¡¡Use¡¡and¡¡Disuse
From¡¡the¡¡facts¡¡alluded¡¡to¡¡in¡¡the¡¡first¡¡chapter£»¡¡I¡¡think¡¡there¡¡can¡¡be¡¡little¡¡doubt¡¡that¡¡use¡¡in¡¡our¡¡domestic¡¡animals¡¡strengthens¡¡and¡¡enlarges¡¡certain¡¡parts£»¡¡and¡¡disuse¡¡diminishes¡¡them£»¡¡and¡¡that¡¡such¡¡modifications¡¡are¡¡inherited¡£¡¡Under¡¡free¡¡nature£»¡¡we¡¡can¡¡have¡¡no¡¡standard¡¡of¡¡comparison£»¡¡by¡¡which¡¡to¡¡judge¡¡of¡¡the¡¡effects¡¡of¡¡long¡­continued¡¡use¡¡or¡¡disuse£»¡¡for¡¡we¡¡know¡¡not¡¡the¡¡parent¡­forms£»¡¡but¡¡many¡¡animals¡¡have¡¡structures¡¡which¡¡can¡¡be¡¡explained¡¡by¡¡the¡¡effects¡¡of¡¡disuse¡£¡¡As¡¡Professor¡¡Owen¡¡has¡¡remarked£»¡¡there¡¡is¡¡no¡¡greater¡¡anomaly¡¡in¡¡nature¡¡than¡¡a¡¡bird¡¡that¡¡cannot¡¡fly£»¡¡yet¡¡there¡¡are¡¡several¡¡in¡¡this¡¡state¡£¡¡The¡¡logger¡­headed¡¡duck¡¡of¡¡South¡¡America¡¡can¡¡only¡¡flap¡¡along¡¡the¡¡surface¡¡of¡¡the¡¡water£»¡¡and¡¡has¡¡its¡¡wings¡¡in¡¡nearly¡¡the¡¡same¡¡condition¡¡as¡¡the¡¡domestic¡¡Aylesbury¡¡duck¡£¡¡As¡¡the¡¡larger¡¡ground¡­feeding¡¡birds¡¡seldom¡¡take¡¡flight¡¡except¡¡to¡¡escape¡¡danger£»¡¡I¡¡believe¡¡that¡¡the¡¡nearly¡¡wingless¡¡condition¡¡of¡¡several¡¡birds£»¡¡which¡¡now¡¡inhabit¡¡or¡¡have¡¡lately¡¡inhabited¡¡several¡¡oceanic¡¡islands£»¡¡tenanted¡¡by¡¡no¡¡beast¡¡of¡¡prey£»¡¡has¡¡been¡¡caused¡¡by¡¡disuse¡£¡¡The¡¡ostrich¡¡indeed¡¡inhabits¡¡continents¡¡and¡¡is¡¡exposed¡¡to¡¡danger¡¡from¡¡which¡¡it¡¡cannot¡¡escape¡¡by¡¡flight£»¡¡but¡¡by¡¡kicking¡¡it¡¡can¡¡defend¡¡itself¡¡from¡¡enemies£»¡¡as¡¡well¡¡as¡¡any¡¡of¡¡the¡¡smaller¡¡quadrupeds¡£¡¡We¡¡may¡¡imagine¡¡that¡¡the¡¡early¡¡progenitor¡¡of¡¡the¡¡ostrich¡¡had¡¡habits¡¡like¡¡those¡¡of¡¡a¡¡bustard£»¡¡and¡¡that¡¡as¡¡natural¡¡selection¡¡increased¡¡in¡¡successive¡¡generations¡¡the¡¡size¡¡and¡¡weight¡¡of¡¡its¡¡body£»¡¡its¡¡legs¡¡were¡¡used¡¡more£»¡¡and¡¡its¡¡wings¡¡less£»¡¡until¡¡they¡¡became¡¡incapable¡¡of¡¡flight¡£¡¡
Kirby¡¡has¡¡remarked¡¡£¨and¡¡I¡¡have¡¡observed¡¡the¡¡same¡¡fact£©¡¡that¡¡the¡¡anterior¡¡tarsi£»¡¡or¡¡feet£»¡¡of¡¡many¡¡male¡¡dung¡­feeding¡¡beetles¡¡are¡¡very¡¡often¡¡broken¡¡off£»¡¡he¡¡examined¡¡seventeen¡¡specimens¡¡in¡¡his¡¡own¡¡collection£»¡¡and¡¡not¡¡one¡¡had¡¡even¡¡a¡¡relic¡¡left¡£¡¡In¡¡the¡¡Onites¡¡apelles¡¡the¡¡tarsi¡¡are¡¡so¡¡habitually¡¡lost£»¡¡that¡¡the¡¡insect¡¡has¡¡been¡¡described¡¡as¡¡not¡¡having¡¡them¡£¡¡In¡¡some¡¡other¡¡genera¡¡they¡¡are¡¡present£»¡¡but¡¡in¡¡a¡¡rudimentary¡¡condition¡£¡¡In¡¡the¡¡Ateuchus¡¡or¡¡sacred¡¡beetle¡¡of¡¡the¡¡Egyptians£»¡¡they¡¡are¡¡totally¡¡deficient¡£¡¡There¡¡is¡¡not¡¡sufficient¡¡evidence¡¡to¡¡induce¡¡us¡¡to¡¡believe¡¡that¡¡mutilations¡¡are¡¡ever¡¡inherited£»¡¡and¡¡I¡¡should¡¡prefer¡¡explaining¡¡the¡¡entire¡¡absence¡¡of¡¡the¡¡anterior¡¡tarsi¡¡in¡¡Ateuchus£»¡¡and¡¡their¡¡rudimentary¡¡condition¡¡in¡¡some¡¡other¡¡genera£»¡¡by¡¡the¡¡long¡­continued¡¡effects¡¡of¡¡disuse¡¡in¡¡their¡¡progenitors£»¡¡for¡¡as¡¡the¡¡tarsi¡¡are¡¡almost¡¡always¡¡lost¡¡in¡¡many¡¡dung¡­feeding¡¡beetles£»¡¡they¡¡must¡¡be¡¡lost¡¡early¡¡in¡¡life£»¡¡and¡¡therefore¡¡cannot¡¡be¡¡much¡¡used¡¡by¡¡these¡¡insects¡£¡¡
In¡¡some¡¡cases¡¡we¡¡might¡¡easily¡¡put¡¡down¡¡to¡¡disuse¡¡modifications¡¡of¡¡structure¡¡which¡¡are¡¡wholly£»¡¡or¡¡mainly£»¡¡due¡¡to¡¡natural¡¡selection¡£¡¡Mr¡£¡¡Wollaston¡¡has¡¡discovered¡¡the¡¡remarkable¡¡fact¡¡that¡¡200¡¡beetles£»¡¡out¡¡of¡¡the¡¡550¡¡species¡¡inhabiting¡¡Madeira£»¡¡are¡¡so¡¡far¡¡deficient¡¡in¡¡wings¡¡that¡¡they¡¡cannot¡¡fly£»¡¡and¡¡that¡¡of¡¡the¡¡twenty¡­nine¡¡endemic¡¡genera£»¡¡no¡¡less¡¡than¡¡twenty¡­three¡¡genera¡¡have¡¡all¡¡their¡¡species¡¡in¡¡this¡¡condition£¡¡¡Several¡¡facts£»¡¡namely£»¡¡that¡¡beetles¡¡in¡¡many¡¡parts¡¡of¡¡the¡¡world¡¡are¡¡very¡¡frequently¡¡blown¡¡to¡¡sea¡¡and¡¡perish£»¡¡that¡¡the¡¡beetles¡¡in¡¡Madeira£»¡¡as¡¡observed¡¡by¡¡Mr¡¡Wollaston£»¡¡lie¡¡much¡¡concealed£»¡¡until¡¡the¡¡wind¡¡lulls¡¡and¡¡the¡¡sun¡¡shines£»¡¡that¡¡the¡¡proportion¡¡of¡¡wingless¡¡beetles¡¡is¡¡larger¡¡on¡¡the¡¡exposed¡¡Dezertas¡¡than¡¡in¡¡Madeira¡¡itself£»¡¡and¡¡especially¡¡the¡¡extraordinary¡¡fact£»¡¡so¡¡strongly¡¡insisted¡¡on¡¡by¡¡Mr¡£¡¡Wollaston£»¡¡of¡¡the¡¡almost¡¡entire¡¡absence¡¡of¡¡certain¡¡large¡¡groups¡¡of¡¡beetles£»¡¡elsewhere¡¡excessively¡¡numerous£»¡¡and¡¡which¡¡groups¡¡have¡¡habits¡¡of¡¡life¡¡almost¡¡necessitating¡¡frequent¡¡flight£»¡¡these¡¡several¡¡considerations¡¡have¡¡made¡¡me¡¡believe¡¡that¡¡the¡¡wingless¡¡condition¡¡of¡¡so¡¡many¡¡Madeira¡¡beetles¡¡is¡¡mainly¡¡due¡¡to¡¡the¡¡action¡¡of¡¡natural¡¡selection£»¡¡but¡¡combined¡¡probably¡¡with¡¡disuse¡£¡¡For¡¡during¡¡thousands¡¡of¡¡successive¡¡generations¡¡each¡¡individual¡¡beetle¡¡which¡¡flew¡¡least£»¡¡either¡¡from¡¡its¡¡wings¡¡having¡¡been¡¡ever¡¡so¡¡little¡¡less¡¡perfectly¡¡developed¡¡or¡¡from¡¡indolent¡¡habit£»¡¡will¡¡have¡¡had¡¡the¡¡best¡¡chance¡¡of¡¡surviving¡¡from¡¡not¡¡being¡¡blown¡¡out¡¡to¡¡sea£»¡¡and£»¡¡on¡¡the¡¡other¡¡hand£»¡¡those¡¡beetles¡¡which¡¡most¡¡readily¡¡took¡¡to¡¡flight¡¡will¡¡oftenest¡¡have¡¡been¡¡blown¡¡to¡¡sea¡¡and¡¡thus¡¡have¡¡been¡¡destroyed¡£¡¡
The¡¡insects¡¡in¡¡Madeira¡¡which¡¡are¡¡not¡¡ground¡­feeders£»¡¡and¡¡which£»¡¡as¡¡the¡¡flower¡­feeding¡¡coleoptera¡¡and¡¡lepidoptera£»¡¡must¡¡habitually¡¡use¡¡their¡¡wings¡¡to¡¡gain¡¡their¡¡subsistence£»¡¡have£»¡¡as¡¡Mr¡£¡¡Wollaston¡¡suspects£»¡¡their¡¡wings¡¡not¡¡at¡¡all¡¡reduced£»¡¡but¡¡even¡¡enlarged¡£¡¡This¡¡is¡¡quite¡¡compatible¡¡with¡¡the¡¡action¡¡of¡¡natural¡¡selection¡£¡¡For¡¡when¡¡a¡¡new¡¡insect¡¡first¡¡arrived¡¡on¡¡the¡¡island£»¡¡the¡¡tendency¡¡of¡¡natural¡¡selection¡¡to¡¡enlarge¡¡or¡¡to¡¡reduce¡¡the¡¡wings£»¡¡would¡¡depend¡¡on¡¡whether¡¡a¡¡greater¡¡number¡¡of¡¡individuals¡¡were¡¡saved¡¡by¡¡successfully¡¡battling¡¡with¡¡the¡¡winds£»¡¡or¡¡by¡¡giving¡¡up¡¡the¡¡attempt¡¡and¡¡rarely¡¡or¡¡never¡¡flying¡£¡¡As¡¡with¡¡mariners¡¡ship¡­wrecked¡¡near¡¡a¡¡coast£»¡¡it¡¡would¡¡have¡¡been¡¡better¡¡for¡¡the¡¡good¡¡swimmers¡¡if¡¡they¡¡had¡¡been¡¡able¡¡to¡¡swim¡¡still¡¡further£»¡¡whereas¡¡it¡¡would¡¡have¡¡been¡¡better¡¡for¡¡the¡¡bad¡¡swimmers¡¡if¡¡they¡¡had¡¡not¡¡been¡¡able¡¡to¡¡swim¡¡at¡¡all¡¡and¡¡had¡¡stuck¡¡to¡¡the¡¡wreck¡£¡¡
The¡¡eyes¡¡of¡¡moles¡¡and¡¡of¡¡some¡¡burrowing¡¡rodents¡¡are¡¡rudimentary¡¡in¡¡size£»¡¡and¡¡in¡¡some¡¡cases¡¡are¡¡quite¡¡covered¡¡up¡¡by¡¡skin¡¡and¡¡fur¡£¡¡This¡¡state¡¡of¡¡the¡¡eyes¡¡is¡¡probably¡¡due¡¡to¡¡gradual¡¡reduction¡¡from¡¡disuse£»¡¡but¡¡aided¡¡perhaps¡¡by¡¡natural¡¡selection¡£¡¡In¡¡South¡¡America£»¡¡a¡¡burrowing¡¡rodent£»¡¡the¡¡tuco¡­tuco£»¡¡or¡¡Ctenomys£»¡¡is¡¡even¡¡more¡¡subterranean¡¡in¡¡its¡¡habits¡¡than¡¡the¡¡mole£»¡¡and¡¡I¡¡was¡¡assured¡¡by¡¡a¡¡Spaniard£»¡¡who¡¡had¡¡often¡¡caught¡¡them£»¡¡that¡¡they¡¡were¡¡frequently¡¡blind£»¡¡one¡¡which¡¡I¡¡kept¡¡alive¡¡was¡¡certainly¡¡in¡¡this¡¡condition£»¡¡the¡¡cause£»¡¡as¡¡appeared¡¡on¡¡dissection£»¡¡having¡¡been¡¡inflammation¡¡of¡¡the¡¡nictitating¡¡membrane¡£¡¡As¡¡frequent¡¡inflammation¡¡of¡¡the¡¡eyes¡¡must¡¡be¡¡injurious¡¡to¡¡any¡¡animal£»¡¡and¡¡as¡¡eyes¡¡are¡¡certainly¡¡not¡¡indispensable¡¡to¡¡animals¡¡with¡¡subterranean¡¡habits£»¡¡a¡¡reduction¡¡in¡¡their¡¡size¡¡with¡¡the¡¡adhesion¡¡of¡¡the¡¡eyelids¡¡and¡¡growth¡¡of¡¡fur¡¡over¡¡them£»¡¡might¡¡in¡¡such¡¡case¡¡be¡¡an¡¡advantage£»¡¡and¡¡if¡¡so£»¡¡natural¡¡selection¡¡would¡¡constantly¡¡aid¡¡the¡¡effects¡¡of¡¡disuse¡£¡¡
It¡¡is¡¡well¡¡known¡¡that¡¡several¡¡animals£»¡¡belonging¡¡to¡¡the¡¡most¡¡different¡¡classes£»¡¡which¡¡inhabit¡¡the¡¡caves¡¡of¡¡Styria¡¡and¡¡of¡¡Kentucky£»¡¡are¡¡blind¡£¡¡In¡¡some¡¡of¡¡the¡¡crabs¡¡the¡¡foot¡­stalk¡¡for¡¡the¡¡eye¡¡remains£»¡¡though¡¡the¡¡eye¡¡is¡¡gone£»¡¡the¡¡stand¡¡for¡¡the¡¡telescope¡¡is¡¡there£»¡¡though¡¡the¡¡telescope¡¡with¡¡its¡¡glasses¡¡has¡¡been¡¡lost¡£¡¡As¡¡it¡¡is¡¡difficult¡¡to¡¡imagine¡¡that¡¡eyes£»¡¡though¡¡useless£»¡¡could¡¡be¡¡in¡¡any¡¡way¡¡injurious¡¡to¡¡animals¡¡living¡¡in¡¡darkness£»¡¡I¡¡attribute¡¡their¡¡loss¡¡wholly¡¡to¡¡disuse¡£¡¡In¡¡one¡¡of¡¡the¡¡blind¡¡animals£»¡¡namely£»¡¡the¡¡cave¡­rat£»¡¡the¡¡eyes¡¡are¡¡of¡¡immense¡¡size£»¡¡and¡¡Professor¡¡Silliman¡¡thought¡¡that¡¡it¡¡regained£»¡¡after¡¡living¡¡some¡¡days¡¡in¡¡the¡¡light£»¡¡some¡¡slight¡¡power¡¡of¡¡vision¡£¡¡In¡¡the¡¡same¡¡manner¡¡as¡¡in¡¡Madeira¡¡the¡¡wings¡¡of¡¡some¡¡of¡¡the¡¡insects¡¡have¡¡been¡¡enlarged£»¡¡and¡¡the¡¡wings¡¡of¡¡others¡¡have¡¡been¡¡reduced¡¡by¡¡natural¡¡selection¡¡aided¡¡by¡¡use¡¡and¡¡disuse£»¡¡so¡¡in¡¡the¡¡case¡¡of¡¡the¡¡cave¡­rat¡¡natural¡¡selection¡¡seems¡¡to¡¡have¡¡struggled¡¡with¡¡the¡¡loss¡¡of¡¡light¡¡and¡¡to¡¡have¡¡increased¡¡the¡¡size¡¡of¡¡the¡¡eyes£»¡¡whereas¡¡with¡¡all¡¡the¡¡other¡¡inhabitants¡¡of¡¡the¡¡caves£»¡¡disuse¡¡by¡¡itself¡¡seems¡¡to¡¡have¡¡done¡¡its¡¡work¡£¡¡
It¡¡is¡¡difficult¡¡to¡¡imagine¡¡conditions¡¡of¡¡life¡¡more¡¡similar¡¡than¡¡deep¡¡limestone¡¡caverns¡¡under¡¡a¡¡nearly¡¡similar¡¡climate£»¡¡so¡¡that¡¡on¡¡the¡¡common¡¡view¡¡of¡¡the¡¡blind¡¡animals¡¡having¡¡been¡¡separately¡¡created¡¡for¡¡the¡¡American¡¡and¡¡European¡¡caver

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