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In¡¡the¡¡bird£»¡¡the¡¡fibula¡¡is¡¡small¡¡and¡¡its¡¡lower¡¡end¡¡diminishes¡¡to

a¡¡point¡£¡¡The¡¡tibia¡¡has¡¡a¡¡strong¡¡crest¡¡at¡¡its¡¡upper¡¡end¡¡and¡¡its

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to¡¡be¡¡no¡¡tarsal¡¡bones£»¡¡and¡¡only¡¡one¡¡bone£»¡¡divided¡¡at¡¡the¡¡end

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appears¡¡in¡¡the¡¡place¡¡of¡¡the¡¡metatarsus¡£



In¡¡the¡¡young¡¡bird£»¡¡however£»¡¡the¡¡pulley¡­shaped¡¡apparent¡¡end¡¡of

the¡¡tibia¡¡is¡¡a¡¡distinct¡¡bone£»¡¡which¡¡represents¡¡the¡¡bones¡¡marked

As¡££»¡¡Ca¡££»¡¡in¡¡the¡¡crocodile£»¡¡while¡¡the¡¡apparently¡¡single

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one¡¡another¡¡and¡¡with¡¡an¡¡additional¡¡bone£»¡¡which¡¡represents¡¡the

lower¡¡row¡¡of¡¡bones¡¡in¡¡the¡¡tarsus¡¡of¡¡the¡¡crocodile¡£



In¡¡other¡¡words£»¡¡it¡¡can¡¡be¡¡shown¡¡by¡¡the¡¡study¡¡of¡¡development¡¡that

the¡¡bird's¡¡pelvis¡¡and¡¡hind¡¡limb¡¡are¡¡simply¡¡extreme¡¡modifications

of¡¡the¡¡same¡¡fundamental¡¡plan¡¡as¡¡that¡¡upon¡¡which¡¡these¡¡parts¡¡are

modelled¡¡in¡¡reptiles¡£



On¡¡comparing¡¡the¡¡pelvis¡¡and¡¡hind¡¡limb¡¡of¡¡the¡¡ornithoscelidan

with¡¡that¡¡of¡¡the¡¡crocodile£»¡¡on¡¡the¡¡one¡¡side£»¡¡and¡¡that¡¡of¡¡the

bird£»¡¡on¡¡the¡¡other¡¡£¨Fig¡£¡¡6£©£»¡¡it¡¡is¡¡obvious¡¡that¡¡it¡¡represents¡¡a

middle¡¡term¡¡between¡¡the¡¡two¡£¡¡The¡¡pelvic¡¡bones¡¡approach¡¡the¡¡form

of¡¡those¡¡of¡¡the¡¡birds£»¡¡and¡¡the¡¡direction¡¡of¡¡the¡¡pubis¡¡and

ischium¡¡is¡¡nearly¡¡that¡¡which¡¡is¡¡characteristic¡¡of¡¡birds£»

the¡¡thigh¡¡bone£»¡¡from¡¡the¡¡direction¡¡of¡¡its¡¡head£»¡¡must¡¡have¡¡lain

close¡¡to¡¡the¡¡body£»¡¡the¡¡tibia¡¡has¡¡a¡¡great¡¡crest£»¡¡and£»¡¡immovably

fitted¡¡on¡¡to¡¡its¡¡lower¡¡end£»¡¡there¡¡is¡¡a¡¡pulley¡­shaped¡¡bone£»¡¡like

that¡¡of¡¡the¡¡bird£»¡¡but¡¡remaining¡¡distinct¡£¡¡The¡¡lower¡¡end¡¡of¡¡the

fibula¡¡is¡¡much¡¡more¡¡slender£»¡¡proportionally£»¡¡than¡¡in¡¡the

crocodile¡£¡¡The¡¡metatarsal¡¡bones¡¡have¡¡such¡¡a¡¡form¡¡that¡¡they¡¡fit

together¡¡immovably£»¡¡though¡¡they¡¡do¡¡not¡¡enter¡¡into¡¡bony¡¡union£»

the¡¡third¡¡toe¡¡is£»¡¡as¡¡in¡¡the¡¡bird£»¡¡longest¡¡and¡¡strongest¡£

In¡¡fact£»¡¡the¡¡ornithoscelidan¡¡limb¡¡is¡¡comparable¡¡to¡¡that¡¡of¡¡an

unhatched¡¡chick¡£





Fig¡£¡¡6¡£Bird¡£¡¡Ornithoscelidan¡£¡¡Crocodile¡£

The¡¡letters¡¡have¡¡the¡¡same¡¡signification¡¡in¡¡all¡¡the¡¡figures¡£

Il¡££»¡¡Ilium£»¡¡a¡£¡¡anterior¡¡end£»¡¡b¡£¡¡posterior

end£»¡¡Ia¡£¡¡ischium£»¡¡Pb¡££»¡¡pubis£»¡¡T£»¡¡tibia£»

F£»¡¡fibula£»¡¡As¡££»¡¡astragalus£»¡¡Ca¡££»¡¡calcaneum£»

I£»¡¡distal¡¡portion¡¡of¡¡the¡¡tarsus£»¡¡i¡££»¡¡ii¡££»¡¡iii¡££»¡¡iv¡££»¡¡metatarsal

bones¡£





Taking¡¡all¡¡these¡¡facts¡¡together£»¡¡it¡¡is¡¡obvious¡¡that¡¡the¡¡view£»

which¡¡was¡¡entertained¡¡by¡¡Mantell¡¡and¡¡the¡¡probability¡¡of¡¡which

was¡¡demonstrated¡¡by¡¡your¡¡own¡¡distinguished¡¡anatomist£»¡¡Leidy£»

while¡¡much¡¡additional¡¡evidence¡¡in¡¡the¡¡same¡¡direction¡¡has¡¡been

furnished¡¡by¡¡Professor¡¡Cope£»¡¡that¡¡some¡¡of¡¡these¡¡animals¡¡may¡¡have

walked¡¡upon¡¡their¡¡hind¡¡legs¡¡as¡¡birds¡¡do£»¡¡acquires¡¡great¡¡weight¡£

In¡¡fact£»¡¡there¡¡can¡¡be¡¡no¡¡reasonable¡¡doubt¡¡that¡¡one¡¡of¡¡the

smaller¡¡forms¡¡of¡¡the¡¡Ornithoscelida£»¡¡Compsognathus£»¡¡the

almost¡¡entire¡¡skeleton¡¡of¡¡which¡¡has¡¡been¡¡discovered¡¡in¡¡the

Solenhofen¡¡slates£»¡¡was¡¡a¡¡bipedal¡¡animal¡£¡¡The¡¡parts¡¡of¡¡this

skeleton¡¡are¡¡somewhat¡¡twisted¡¡out¡¡of¡¡their¡¡natural¡¡relations£»

but¡¡the¡¡accompanying¡¡figure¡¡gives¡¡a¡¡just¡¡view¡¡of¡¡the¡¡general

form¡¡of¡¡Compsognathus¡¡and¡¡of¡¡the¡¡proportions¡¡of¡¡its

limbs£»¡¡which£»¡¡in¡¡some¡¡respects£»¡¡are¡¡more¡¡completely¡¡bird¡­like

than¡¡those¡¡of¡¡other¡¡Ornithoscelida¡£





Fig¡£¡¡7¡£Restoration¡¡of¡¡Compsognathus¡¡Longipes





We¡¡have¡¡had¡¡to¡¡stretch¡¡the¡¡definition¡¡of¡¡the¡¡class¡¡of¡¡birds¡¡so

as¡¡to¡¡include¡¡birds¡¡with¡¡teeth¡¡and¡¡birds¡¡with¡¡paw¡­like¡¡fore

limbs¡¡and¡¡long¡¡tails¡£¡¡There¡¡is¡¡no¡¡evidence¡¡that

Compsognathus¡¡possessed¡¡feathers£»¡¡but£»¡¡if¡¡it¡¡did£»¡¡it

would¡¡be¡¡hard¡¡indeed¡¡to¡¡say¡¡whether¡¡it¡¡should¡¡be¡¡called¡¡a

reptilian¡¡bird¡¡or¡¡an¡¡avian¡¡reptile¡£



As¡¡Compsognathus¡¡walked¡¡upon¡¡its¡¡hind¡¡legs£»¡¡it¡¡must¡¡have

made¡¡tracks¡¡like¡¡those¡¡of¡¡birds¡£¡¡And¡¡as¡¡the¡¡structure¡¡of¡¡the

limbs¡¡of¡¡several¡¡of¡¡the¡¡gigantic¡¡Ornithoscelida£»¡¡such¡¡as

Iguanodon£»¡¡leads¡¡to¡¡the¡¡conclusion¡¡that¡¡they¡¡also¡¡may

have¡¡constantly£»¡¡or¡¡occasionally£»¡¡assumed¡¡the¡¡same¡¡attitude£»¡¡a

peculiar¡¡interest¡¡attaches¡¡to¡¡the¡¡fact¡¡that£»¡¡in¡¡the¡¡Wealden

strata¡¡of¡¡England£»¡¡there¡¡are¡¡to¡¡be¡¡found¡¡gigantic¡¡footsteps£»

arranged¡¡in¡¡order¡¡like¡¡those¡¡of¡¡the¡¡Brontozoum£»¡¡and¡¡which

there¡¡can¡¡be¡¡no¡¡reasonable¡¡doubt¡¡were¡¡made¡¡by¡¡some¡¡of¡¡the

Ornithoscelida£»¡¡the¡¡remains¡¡of¡¡which¡¡are¡¡found¡¡in¡¡the

same¡¡rocks¡£¡¡And£»¡¡knowing¡¡that¡¡reptiles¡¡that¡¡walked¡¡upon¡¡their

hind¡¡legs¡¡and¡¡shared¡¡many¡¡of¡¡the¡¡anatomical¡¡characters¡¡of¡¡birds

did¡¡once¡¡exist£»¡¡it¡¡becomes¡¡a¡¡very¡¡important¡¡question¡¡whether¡¡the

tracks¡¡in¡¡the¡¡Trias¡¡of¡¡Massachusetts£»¡¡to¡¡which¡¡I¡¡referred¡¡some

time¡¡ago£»¡¡and¡¡which¡¡formerly¡¡used¡¡to¡¡be¡¡unhesitatingly¡¡ascribed

to¡¡birds£»¡¡may¡¡not¡¡all¡¡have¡¡been¡¡made¡¡by¡¡ornithoscelidan

reptiles£»¡¡and¡¡whether£»¡¡if¡¡we¡¡could¡¡obtain¡¡the¡¡skeletons¡¡of¡¡the

animals¡¡which¡¡made¡¡these¡¡tracks£»¡¡we¡¡should¡¡not¡¡find¡¡in¡¡them¡¡the

actual¡¡steps¡¡of¡¡the¡¡evolutional¡¡process¡¡by¡¡which¡¡reptiles¡¡gave

rise¡¡to¡¡birds¡£



The¡¡evidential¡¡value¡¡of¡¡the¡¡facts¡¡I¡¡have¡¡brought¡¡forward¡¡in¡¡this

Lecture¡¡must¡¡be¡¡neither¡¡over¡¡nor¡¡under¡¡estimated¡£¡¡It¡¡is¡¡not

historical¡¡proof¡¡of¡¡the¡¡occurrence¡¡of¡¡the¡¡evolution¡¡of¡¡birds

from¡¡reptiles£»¡¡for¡¡we¡¡have¡¡no¡¡safe¡¡ground¡¡for¡¡assuming¡¡that¡¡true

birds¡¡had¡¡not¡¡made¡¡their¡¡appearance¡¡at¡¡the¡¡commencement¡¡of¡¡the

Mesozoic¡¡epoch¡£¡¡It¡¡is£»¡¡in¡¡fact£»¡¡quite¡¡possible¡¡that¡¡all¡¡these

more¡¡or¡¡less¡¡avi¡­form¡¡reptiles¡¡of¡¡the¡¡Mesozoic¡¡epochs¡¡are¡¡not

terms¡¡in¡¡the¡¡series¡¡of¡¡progression¡¡from¡¡birds¡¡to¡¡reptiles¡¡at

all£»¡¡but¡¡simply¡¡the¡¡more¡¡or¡¡less¡¡modified¡¡descendants¡¡of

Palaeozoic¡¡forms¡¡through¡¡which¡¡that¡¡transition¡¡was

actually¡¡effected¡£



We¡¡are¡¡not¡¡in¡¡a¡¡position¡¡to¡¡say¡¡that¡¡the¡¡known

Ornithoscelida¡¡are¡¡intermediate¡¡in¡¡the¡¡order¡¡of¡¡their

appearance¡¡on¡¡the¡¡earth¡¡between¡¡reptiles¡¡and¡¡birds¡£¡¡All¡¡that¡¡can

be¡¡said¡¡is¡¡that£»¡¡if¡¡independent¡¡evidence¡¡of¡¡the¡¡actual

occurrence¡¡of¡¡evolution¡¡is¡¡producible£»¡¡then¡¡these¡¡intercalary

forms¡¡remove¡¡every¡¡difficulty¡¡in¡¡the¡¡way¡¡of¡¡understanding¡¡what

the¡¡actual¡¡steps¡¡of¡¡the¡¡process£»¡¡in¡¡the¡¡case¡¡of¡¡birds£»¡¡may

have¡¡been¡£



That¡¡intercalary¡¡forms¡¡should¡¡have¡¡existed¡¡in¡¡ancient¡¡times¡¡is¡¡a

necessary¡¡consequence¡¡of¡¡the¡¡truth¡¡of¡¡the¡¡hypothesis¡¡of

evolution£»¡¡and£»¡¡hence£»¡¡the¡¡evidence¡¡I¡¡have¡¡laid¡¡before¡¡you¡¡in

proof¡¡of¡¡the¡¡existence¡¡of¡¡such¡¡forms£»¡¡is£»¡¡so¡¡far¡¡as¡¡it¡¡goes£»¡¡in

favour¡¡of¡¡that¡¡hypothesis¡£



There¡¡is¡¡another¡¡series¡¡of¡¡extinct¡¡reptiles¡¡which¡¡may¡¡be¡¡said¡¡to

be¡¡intercalary¡¡between¡¡reptiles¡¡and¡¡birds£»¡¡in¡¡so¡¡far¡¡as¡¡they

combine¡¡some¡¡of¡¡the¡¡characters¡¡of¡¡both¡¡these¡¡groups£»¡¡and¡¡which£»

as¡¡they¡¡possessed¡¡the¡¡power¡¡of¡¡flight£»¡¡may¡¡seem£»¡¡at¡¡first¡¡sight£»

to¡¡be¡¡nearer¡¡representatives¡¡of¡¡the¡¡forms¡¡by¡¡which¡¡the

transition¡¡from¡¡the¡¡reptile¡¡to¡¡the¡¡bird¡¡was¡¡effected£»¡¡than¡¡the

Ornithoscelida¡£



These¡¡are¡¡the¡¡Pterosauria£»¡¡or¡¡Pterodactyles£»¡¡the¡¡remains

of¡¡which¡¡are¡¡met¡¡with¡¡throughout¡¡the¡¡series¡¡of¡¡Mesozoic¡¡rocks£»

from¡¡the¡¡lias¡¡to¡¡the¡¡chalk£»¡¡and¡¡some¡¡of¡¡which¡¡attained¡¡a¡¡great

size£»¡¡their¡¡wings¡¡having¡¡a¡¡span¡¡of¡¡eighteen¡¡or¡¡twenty¡¡feet¡£

These¡¡animals£»¡¡in¡¡the¡¡form¡¡and¡¡proportions¡¡of¡¡the¡¡head¡¡and¡¡neck

relatively¡¡to¡¡the¡¡body£»¡¡and¡¡in¡¡the¡¡fact¡¡that¡¡the¡¡ends¡¡of¡¡the

jaws¡¡were¡¡often£»¡¡if¡¡not¡¡always£»¡¡more¡¡or¡¡less¡¡extensively

ensheathed¡¡in¡¡horny¡¡beaks£»¡¡remind¡¡us¡¡of¡¡birds¡£¡¡Moreover£»¡¡their

bones¡¡contained¡¡air¡¡cavities£»¡¡rendering¡¡them¡¡specifically

lighter£»¡¡as¡¡is¡¡the¡¡case¡¡in¡¡most¡¡birds¡£¡¡The¡¡breast¡¡bone¡¡was¡¡large

and¡¡keeled£»¡¡as¡¡in¡¡most¡¡birds¡¡and¡¡in¡¡bats£»¡¡and¡¡the¡¡shoulder

girdle¡¡is¡¡strikingly¡¡similar¡¡to¡¡that¡¡of¡¡ordinary¡¡birds¡£¡¡But£»¡¡it

seems¡¡to¡¡me£»¡¡that¡¡the¡¡special¡¡resemblance¡¡of¡¡pterodactyles¡¡to

birds¡¡ends¡¡here£»¡¡unless¡¡I¡¡may¡¡add¡¡the¡¡entire¡¡absence¡¡of¡¡teeth

which¡¡characterises¡¡the¡¡great¡¡pterodactyles¡¡£¨Pteranodon£©

discovered¡¡by¡¡Professor¡¡Marsh¡£¡¡All¡¡other¡¡known¡¡pterodactyles

have¡¡teeth¡¡lodged¡¡in¡¡sockets¡£¡¡In¡¡the¡¡vertebral¡¡column¡¡and¡¡the

hind¡¡limbs¡¡there¡¡are¡¡no¡¡special¡¡resemblances¡¡to¡¡birds£»¡¡and¡¡when

we¡¡turn¡¡to¡¡the¡¡wings¡¡they¡¡are¡¡found¡¡to¡¡be¡¡constructed¡¡on¡¡a

totally¡¡different¡¡principle¡¡from¡¡those¡¡of¡¡birds¡£





Fig¡£¡¡8¡£Pterodactylus¡¡Spectabilis¡¡£¨Von¡¡Meyer£©¡£





There¡¡are¡¡four¡¡fingers¡£¡¡These¡¡four¡¡fingers¡¡are¡¡large£»¡¡and¡¡three

of¡¡them£»¡¡those¡¡which¡¡answer¡¡to¡¡the¡¡thumb¡¡and¡¡two¡¡following

fingers¡¡in¡¡my¡¡handare¡¡terminated¡¡by¡¡claws£»¡¡while¡¡the¡¡fourth¡¡is

enormously¡¡prolonged¡¡and¡¡converted¡¡into¡¡a¡¡great¡¡jointed¡¡style¡£

You¡¡see¡¡at¡¡once£»¡¡from¡¡what¡¡I¡¡have¡¡stated¡¡about¡¡a¡¡bird's¡¡wing£»

that¡¡there¡¡could¡¡be¡¡nothing¡¡less¡¡like¡¡a¡¡bird's¡¡wing¡¡than¡¡this

is¡£¡¡It¡¡was¡¡concluded¡¡by¡¡general¡¡reasoning¡¡that¡¡this¡¡finger¡¡had

the¡¡office¡¡of¡¡supporting¡¡a¡¡web¡¡which¡¡extended¡¡between¡¡it¡¡and¡¡the

body¡£¡¡An¡¡existing¡¡specimen¡¡proves¡¡that¡¡such¡¡was¡¡really¡¡the¡¡case£»

and¡¡that¡¡the¡¡pterodactyles¡¡were¡¡devoid¡¡of¡¡feathers£»¡¡but¡¡that¡¡the

fingers¡¡supported¡¡a¡¡vast¡¡web¡¡like¡¡that¡¡of¡¡a¡¡bat's¡¡wing£»¡¡in¡¡fact£»

there¡¡can¡¡be¡¡no¡¡doubt¡¡that¡¡this¡¡ancient¡¡reptile¡¡flew¡¡after¡¡the

fashion¡¡of¡¡a¡¡bat¡£



Thus£»¡¡though¡¡the¡¡pterodactyle¡¡is¡¡a¡¡reptile¡¡which¡¡has¡¡become

modified¡¡in¡¡such¡¡a¡¡manner¡¡as¡¡to¡¡enable¡¡it¡¡to¡¡fly£»¡¡and¡¡therefore£»

as¡¡might¡¡be¡¡expected£»¡¡presents¡¡some¡¡points¡¡of¡¡resemblance¡¡to

other¡¡animals¡¡which¡¡fly£»¡¡it¡¡has£»¡¡so¡¡to¡¡speak£»¡¡gone¡¡off¡¡the¡¡line

which¡¡leads¡¡directly¡¡from¡¡reptiles¡¡to¡¡birds£»¡¡and¡¡has¡¡become

disqualified¡¡for¡¡the¡¡changes¡¡which¡¡lead¡¡to¡¡the¡¡characteristic

organisation¡¡of¡¡the¡¡latter¡¡class¡£¡¡Therefore£»¡¡viewed¡¡in¡¡relation

to¡¡the¡¡classes¡¡of¡¡reptiles¡¡and¡¡birds£»¡¡the¡¡pterodactyles¡¡appear

to¡¡me¡¡to¡¡be£»¡¡in¡¡a¡¡limited¡¡sense£»¡¡intercalary¡¡forms£»¡¡but¡¡they¡¡are

not¡¡even¡¡approximately¡¡linear£»¡¡in¡¡the¡¡sense¡¡of¡¡exemplifying

those¡¡modifications¡¡of¡¡structure¡¡through¡¡which¡¡the¡¡passage¡¡from

the¡¡reptile¡¡to¡¡the¡¡bird¡¡took¡¡place¡£







LECTURES¡¡ON¡¡EVOLUTION



III

THE¡¡DEMONSTRATIVE¡¡EVIDENCE¡¡OF¡¡EVOLUTION





The¡¡occurrence¡¡of¡¡historical¡¡facts¡¡is¡¡said¡¡to¡¡be¡¡demonstrated£»

when¡¡the¡¡evidence¡¡that¡¡they¡¡happened¡¡is¡¡of¡¡such¡¡a¡¡character¡¡as

to¡¡render¡¡the¡¡assumption¡¡that¡¡they¡¡did¡¡not¡¡happen¡¡in¡¡the¡¡highest

degree¡¡improbable£»¡¡and¡¡the¡¡question¡¡I¡¡now¡¡have¡¡to¡¡deal¡¡with¡¡is£»

whether¡¡evidence¡¡in¡¡favour¡¡of¡¡the¡¡evolution¡¡of¡¡animals¡¡of¡¡this

degree¡¡of¡¡cogency¡¡is£»¡¡or¡¡is¡¡not£»¡¡obtainable¡¡from¡¡the¡¡record¡¡of

the¡¡succession¡¡of¡¡living¡¡forms¡¡which¡¡is¡¡presented¡¡to¡¡us¡¡by

fossil¡¡remains¡£
СÌáʾ£º°´ »Ø³µ [Enter] ¼ü ·µ»ØÊéÄ¿£¬°´ ¡û ¼ü ·µ»ØÉÏÒ»Ò³£¬ °´ ¡ú ¼ü ½øÈëÏÂÒ»Ò³¡£ ÔÞһϠÌí¼ÓÊéÇ©¼ÓÈëÊé¼Ü