by

FRANK H. T. R HODES
The University of Michigan
ILLUSTRATED BY

REBECCA MERRILEES

and

RUDY ZALLINGER

GOLDEN PRESS

NEW YORK

Western Publishing Company, Inc.

Racine, Wisconsin

FOREWORD

How l ife arose a n d h ow m a n developed are two q u es

tio n s that are as old a s man h i m s e l f, a s the cre a t i o n
a c c oun ts o f m a n y civi l i z atio n s b e a r witn ess. But a n c i e n t
a s i s t h i s con c ern , t h e i m p l icatio n s o f m a n ' s re l a ti o n s h i p
t o t h e wo r l d o f l ivi n g th i n g s are a s s ig n ifica n t i n t h e
S p a ce Ag e a s t h ey were i n t h e Sto n e A g e. Th i s boo k i s
a s i m p l e a c c o u n t o f m a n's search for t h o s e ori g i n s a n d
rel a ti o n s h i p s. I t d escri bes t h e h i storica l d eve l o p m e n t of
the pres e n t t h eory of evo l uti o n , or d es c e n t with m o d ifi
cati o n , the i n d i ca ti o n s that sup port it, its n a ture a n d
m ec h a n i s m , a n d its res u l t i n th e l o n g h i story of l ife.
The boo k c o n c l u d es with a secti o n on t h e m ea n i n g of
evolutio n , for the th eory of evo luti o n h a s h a d a p rofoun d
i m p a ct o n m a n ' s view of h i m s e l f a n d h i s re lat i o n s h i p to
th e worl d in w h i c h h e l i ves. Evo luti o n a ry th eory provi d e s
a powerfu l ex p l a n a t i o n of h ow l i fe d eve l o p e d , yet b e
yo n d i t, a n d un a n swered by i t, l i e s t h e ul ti m a te questi o n
o f w h y l ife d eve l o p e d . T h a t q uestio n , confro n ti n g a s i t
does t h e l a rg er s i g n ifica n c e of l ife, t h oug h t h e a b stra c
tive m ethod s of s c i e n c e provi d e n o a p propriate so l ution
to it, is neit her m ea n i n g l e s s n or inco n s e q uentia l. for in
our re s p o n s e to it, i n d ividua l l y a n d c o l lective ly, l i e s the
future of evol uti on , a n d with it the future of ma n k i n d.
I a m g ra teful to m y co l l ea g u e D r. A l fred S m it h w h o
k i n d ly rea d t h e m a n u sc r i pt o f t h i s book.
Fra n k H. T. R ho d e s

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C O N T E N TS
OVE RVIEW OF LIFE .

. .4-29

Di vers ity, develop m e n t a n d orig i n of l i fe; spon

ta neous g e n era t i o n ; c l a s s i f i cation; degrees o f
d i vers ity a n d d i scove ry of l i fe's lon g h i story;
deve l o p m e n t of t h e theory of evo l u t i o n from
Aristotle to La m a rc k to C h a rl e s D a rw i n a n d
Alfred R u s s e l Wa l l a ce; t h e voya g e of t h e Beagle
a n d p u b l icat ion o f O n the Origin of Species; t h e
w o r k of G r e g o r M e n d e l , lea d i n g to t h e laws o f
i n h e rita n c e a n d to the new synthesic t h eory of
evo l u t i o n .
.... 30-55
Con t i n u ity, u n ity a n d t h e n a t u re of l i fe; i n ter
depen d e n c e a n d i m p l icatio n s of s i m i la r i t i e s
a n a to m i c a l , e m b ryolog ica l , b i oc h e m i ca l , a n d
serolog i ca l ; ad aptations; geogra p h ical d i s t r i b u
tion; l i v i n g species; c h a n g es i n species; fos s i l s a n d
h i g h e r ta xa; m i ss i n g l i n k s a n d t h e fos s i l record .

I NDICATI O N S OF EVO LUTION .

. .56-1()3
I n herita n c e : Ce l l d i v i s i o n ; patte r n s a n d laws;
proba b i l ity and m e c h a n i s m ; g e n e s ; c h e m istry
(DNA a n d RNA), sources of varia b i l ity; reco m
b i nation a n d m u ta t i o n ; g e n e t i c d ri ft; i s o l a t i o n
a n d m i g ra t i o n .

THE PROCESS OF EVO LUTI O N .

N a t u r a l Selectio n : Nature of n a t u r a l selection

a n d its action i n living popu lations a n d fos s i l s ;
adaptation a n d m i m icry; sexual selectio n ; n a t
u ra l selection i n m a n ; m i s s i n g l i n k s .
T i m e: T h e geolog i c t i m e scale , rates of a n d
recipe of evo l u t i o n .
THE COURSE O F EVOLUT I O N .

104-150
T h e p r i m itive eart h ; orig i n of l i fe; fos s i l s-th e
o l d est; m a r i n e i n verte b rates-t h e oldest; l i f e o n
l a n d ; l a n d , va s c u l a r seedless a n d seed bea r i n g
p l a nts; a m p h i b i a n s ; t h e r i s e of a n d d o m i n a n c e
of t h e re ptiles; a doptive rad iation; b i rds; evol u
t i o n a n d g eog ra p h i c d i st r i but i on of m a m m a l s ;
pri mates a n d evo l u t i o n of h u m a n societies.

T H E M E A N I N G OF EVOLUT I O N

.... 151-155
I t s i m p l icatio n s ; perspective; t h e future of m a n .

M O R E INFORMAT I O N.

.156

I NDEX

.1 57

OVERVIEW OF LIFE
Ea rt h tee m s with l ife . living cre a t u re s exist from ocean
dept h s to t h e h i g h e st m o u ntain p e a k s, fro m e q u a to ri a l
j u n g les a n d h o t m inera l springs t o t h e frozen p o l a r
waste l ands, fro m t h e b l inding b rig h tness a nd a ri d i ty
of t h e desert to t h e d a rk intesti n e s of a ni m a l s . I n e a c h
envi ro n m e n t, unto l d nu m bers o f indivi d ua l o rg a ni s m s
i n h abit eve ry n o o k and cranny of t h e a va i l a b l e s p a c e .
Most a n im a l a n d p l a n t species conta i n a myriad
i n d ivid u a l s . Thus the s u rface layer of m ost m e a d ow
soil s conta i n s several m i l l io n a n i m a l s per a c re . Micro
scopic a n i m a ls and pla nts exist i n u n cou n ta b l e n u m b e rs .
O n e g ra m of soil may conta i n h u n d reds of m i l l i o n s of
l ivin g t h i n g s . B i rds a n d in sects exist i n populatio n s so
vast a s to con stitute loca l "plag ue" con d ition s . Aquatic
l ife is n o l ess p rol ific.
It is u n l i ke l y t h a t the e a rt h is u ni q u e in this respect .
I t h a s bee n c a l c u l a ted t h a t t h e re m a y b e m i l l i ons of
p l a nets in ot h er p a rts of the u n iverse ca p a b l e of s u p
p o rti n g s o m e form o f l ife.
Eac h e nviro n ment s u p ports
a n i m a ls .

distinctive co m m u n ity of pla nts a n d

6. Other invertebrates-2 1 ,000

5. Wormlik e phyla-38,000
4. Protozoans-30,000

3. Chordates-45,000
2. Mollus ks-45,000
1 . Art h ropods-900,000

MAJOR GROUPS (PHYLA)

OF ANIMALS-OVER
1 ,000,000 SPE CIES

DIVERSITY OF LIFE i s shown by t h e existence of

m o re t h a n o n e m i l l io n ki n d s (species) of a n i m als a n d
m o re t h a n 350,0 00 k i n d s o f p l a nts .
An i m a l s ra n g e i n size from a few thousa ndths of
a n inc h to m o re th a n 1 00 feet in l e n g t h . They represent
a vast variety of ways of l ife-p a rasites, pred ators,
herb ivores, swi m m ers, fl i e rs, crawlers, burrowers. S o m e
spend their l i ves fi x e d i n o n e spot; others u n dertake
sea sona l m i g rations of thousa n d s of m i les.
In spite of t h e m a ny ki n ds of a n i m a l s a n d p l a n ts,
t h ey rep rese n t o n ly a few ba sic g ro u p s ( p hy l a ) .

4. Algae a n d F u n g i-60,000
3. Mosses

and

L iverworts-

23,000

2. Ferns, Con ifers, et c.-1 0,000

1. Floweri ng

Pla nts-250,000

MAJOR GROUPS (PHYLA)

OF PLANTS-ABOUT
350,000 SPECIES

THE DEVELOPMENT OF LI FE h a s a lways b e e n o n e of

m a n's g re a t conce rns . A n c i ent s a cred writing s of

m a ny fa it h s d i s c u s s this q uestio n . T h e ea r l y c h a p t e rs
of the Boo k of Genesis, fo r exa m p le, d e a l with the
seq u e n ce of c re a t i o n , and Ad a m n a m e d t h e d iffere n t
k i n d s o f a n i m a l s . T h e nee d t o c l a s sify l iving t h ings w a s
p a rt l y p ra ctica l . So m e p l ants were poisonous, ot h e rs
e d i b l e . So m e ani m a ls were h a rm fu l , ot h e rs we re n o t .
Ea r l y m a n's s u rviva l d e pended o n h i s s k i l l i n recogniz
i n g each kind . Ma n 's d a i l y e x p e rience and re l i g i o u s
t ra d ition coinci d e d h e re : every a ni m a l a n d p l a n t t h a t
h e recognized re p rod u ce d " a fte r i t s own k i nd . "
Ma n's e a r l y life a s a h u nter b ro u g ht h i m in c lose
contact with a n i m a ls, and a n cient cave pa inting s b e a r
a record of h i s inte rest. Late r d o m estication of a n i
m a l s and h a rvesting of c rops inc rea se d thi s concern.
Creation of Ada m f r o m M i c h e l a n g e l o 's p a i n t i n g of Sisti n e C h a p e l
Cei l i n g .

r
)

ARISTOTLE, G reek p h i losopher,

was a ls o one o f the fi rs t a n d
greates t b i o l o g i sts. He wrote
exten sive ly o n the c l a s s ification
a n d s tructure of over 5 0 0 s pe
c i e s of a n i m a l s f r o m t h e M e d i
terra n e a n a re a . Ari s totle w a s a
gifted observer, a n d described
details of s uc h t h i n g s a s ch ick
em bryo logy. H e accepted the
sponta n eo u s g e n era tion of fl ies
from p u trefyi n g m a tter, b u t was
also concerned about the prob
lems of h eredity.

A R I ST O T L E (384-32 2 B. C . ), p u p i l of P l a to and tutor

of Alexand e r the Great, o b s e rved t h a t s pecies a p pea red
to be unc h a ng ing . Cows p ro d u ced onl y cows; horses
a rose only fro m horses. Between the two t h e re wa s a
clea r d ivision. S p ecies we re c h a ra cte rized by t h e i r re p ro
d u ctive isolation. Ind ivid u a l s d eveloped, a ccord ing to
A ristotle, by the c a p acity ( psyc he) of e a c h to confo rm
to the a rc h etype c h a ra cters of t h e s p e c i e s relations h i p s .
H e constru cted a "la d d e r o f Natu re" s h owing t h e unity
of p l an.
In contra st to A ristotle's "vita l i st" views, the De
m o c rita n s were " m e c h a n ists ." T h e y b e l i eved t h a t a n
o rganis m's a ctivity wa s the resu l t o f t h e intera ction
of the ato m s of w h i c h it wa s m a d e . Altho u g h vita l istic
and m e c h anistic controve rsies sti ll persist, the s u p posed
confl ict b etwe en s c i e n c e and relig ion being an exa m p le,
the two views a re ofte n co m pl e m e n ta ry, not c o m petitive .
In s o m e situ a tio n s , we need to u s e both ( p . 1 5 5 ) . T h e
d iscuss ion i n t h i s book is m e c h anistic ("how" t h ing s d e
velop, not "why") , but that do es n o t m e a n that l ife h a s
n o m ea n i n g a n d p u rpose .

THE ORI G IN OF LIFE

wa s

long regarded as a spon

taneous eve n t : l iving thing s a rose fro m nonlivi n g m at

ter. Althoug h the va rious g ro u ps of l iving thing s were
b e l i eved to h ave b e en created in d efinite s eq u e n ce, it
was su pposed that each k ind of ani m a l and p l ant
a rose " fu l l y fo r m ed " fro m the d ust of t h e e a rth . S u c h
a view involved n o obvious contra d i ctions . Flies, for
exa m p l e , cou l d be seen to d evelop fro m m a g g ots,
w h i c h a rose " s pontaneously" in decayi n g m eat. T h e
s pontaneous g enerati o n o f l iving t h ings b e ca m e a
universa l assu m ption. We sti l l s p e a k of d i rt " b ree d i n g "
ve r m in. T h e view w a s a l so econo m ical : it involved only
one category of expl anation . O u r c u rrent popu l a r
vi ews req u i re not only a n explanation for t h e origin
of l ife b u t a l so one for t h e origin of species.
Early views o n the o r i g i n o f l i f e i n c l u ded o n e t h a t s u g gested s h ee p
a rose from a pla nt. (Afte r Weinberg.)

of l iv i n g cre atures
from n onlivi n g m a tter beca m e increasing l y s u spect in
the seve nteenth c e n tu ry. Francesco Redi ( 1621-97), an
Itali a n p hysici a n , beca m e convi n ce d that the m a g gots
fou n d in m eat were d e rived n ot fro m the m e a t itself
but fro m e g g s l a i d by flies .
S PONTA N E OUS G E N E R A T I O N

f l i es, d e c o y i n g
m ea t , a n d m a g g ots

p l a ced a "dead s n ake,

s o m e fi s h , a n d a s l ice of vea l"
in four o p e n - m o u t h e d fl a s k s ,
and then placed the same
t h i n g s i n f o u r fl a s k s t h a t h e
c l o s e d a n d s e a l e d . F l ies con
sta ntly settl ed o n the m eat in
the open fl a s k s , w h i c h beca m e
w o r m y . N o wor m s a p peared o n
t h e m e a t i n t h e s e a led fl a s k s .
Knowi n g t h a t s o m e b e l ieved
air to b e e s s e n tia l for g e n e ra
t i o n , Red i repeated t h e ex per
i m ent, t h i s t i m e u s i n g a g a u z e
cover f o r t h e " c losed" fl a s k s
t o p rotect t h e m f ro m fl i e s b u t
a l lowi n g a i r i n s i d e . Ag a i n , n o
m a g g ots a p pea red o n t h e m eat.
This d i scredi ted the most fa m i l
i a r e x a m p l e o f s po n ta n e o u s
g e n e ra tio n .
R EDI

meat decoy ing,

b u t n o f l i e s o r m a g g ots

A R EFINED VE RSIO N of Red i's

experi m e n t was used by Pa ste u r

i n the m id - e i g htee n t h c e n t u ry
to d e m o n strate t h a t p u trefa ction
a n d f e r m e n ta t i o n depend o n
action of ai r - b o r n e o rg a n i s m s .

..,

Ope nf l ies a n d
m a g g ots o n
decoy i n g m e a t

Cove red w i t h
g a u zeno f l i e s or
m a g g o ts o n
d e c oy i n g m e a t

A C LASS IFICATI O N OF LIFE was devised by Aristo t l e

i n t h e fo urth c e n tu ry B . C . a n d stood und i s p ut e d for

n i n eteen c e n t u r i e s . This c l a ss i fi c a tion e m b r a c e d a
co m pl ete g ra d a t ion fro m t h e l owest to t h e h i g h est
o r g a n i s m-m a n .
Fifte e n t h and s i xteenth centu ry voya g es of d i scov
e ry a n d t h e inven tion of t h e m i croscope revea l e d a
d i ve rsity of a n i m a l a n d pl ant fo rm and functio n un
k n own to Aristot l e . With these n ew observatio n s,
c h a n g e s in classification too k p l a ce .
(1627-1705), a n En
g l is h natural ist, i n trod uced the
present idea of species and
h ig he r categories i n classifica
tion . Ray s h owed t h a t groups
of s i m i l a r s pecies co u l d be
classified i n to sets, w h ich he
called genera. This syste m is
t h e b a s i s for t h e i n tern atio n a l
o n e s ti l l b e i n g u s e d today.
JO H N RAY

( 1 707- 1 77 8 ), a
Swed i s h na t u ra l i st, devel ope d
the present syste m a n d method
of biological c l a s s i f icatio n (ta x
o n omy). H e u sed a u n iform
system of c l a s s i f ication a n d no
m e n clature. T h e 1 Ot h edition of
h i s Syste m a Naturae ( 1 758)
marks the beg i n n i n g of modern
tax o n o my.

CARL LIN N A EUS

RELATIVE AGE OF
CATEGORIES OF ANIMALS

s h ows a n in cre a s i n g sim i

l a r i ty of e a c h g ro u p from t h e kin g d o m to t h e s pe c i e s . N ote t h e
modern evo l u ti o n a ry bra n c h i n g in terpre t a tion o n t h e rig h t.

THE L I N N A E A N T Y P E CLA S S I FICATI O N

IN BIN OMIAL NOME NCLATUR E ,

t h e ba sis of the cla ssif i ca tio n

developed by Lin n a e u s , each
s pecies h a s two n a m es: t h e f i rst
is the g e n u s to which it be
longs; the seco n d is the spe
cies. T h e Com m o n Rave n , for

exa m p l e , is Corv u s cora x , w h i l e

t h e s o m ewhat similar Co m mo n
Crow is Corvus brach yrhynchos.
Lin n a e u s used this s h ort, a n d
internatio n a l l y u n d e rstood clas
sification to c l a s sify all of the
species k nown a t that ti m e .

L I N N A E U S and m ost of his contem pora ries a s s u m ed

that e a c h sp e cies was d i stinct and unc hang ing , t h e i r

d e g rees of s i m il a rity refl ecting s i m i l a rity to the a rc he
types, o r m o d els, u pon w h i c h e a c h h a d been c reated .

11

THE VARYING DEGREES OF D IVERS ITY shown

by

d ifferent species s u g g ested to so m e e i g h teenth c e n t u ry

stu d ents a conclu s ion bol dly d ifferent fro m that rea c h e d
by li n n aeus and m o s t o f h i s contem poraries. P e r h a ps,
it was a rg u e d , s pecies were not u n c h ang ing and i m
m u ta ble . P e r h a p s existing s pecies a rose b y a slow
m o d ification of e a r l i er for m s . P e r h a p s d e g rees of
s i m ilarity b etween s pecies reflected t h e i r d eg ree of
rela tions h i p to co m m on ancestra l fo r m s . Perha p s
c h ange, not constancy, w a s one essenti a l c h a r a cteristic
of s pecies . Perh a ps species have evolved , o r unfol d e d ,
r a t h e r t h a n h a ving a p pea re d fu lly fo r m e d . Perh a p s
t h ey a rose not fro m a sing l e c rea tive act b u t b y s l ow
p rocesses of c h ange ove r l ong p e ri o d s of ti m e .
ERASMUS DARW I N ( 1 7 3 1 - 1 802),

JEAN

g ra n d father of C harles Darwi n ,

w a s a ph ysicia n , poet, a n d
natura list. H e was impressed by
t h e exte n t of c h a nges in farm
wit h i n t h e lifeti m e of indiv i d u a l
a n ima ls (frog s , for example), by
the i nfluence of s e lective breed
ing i n horses and dogs, by
d iffe rences d u e to c l i mate, a n d
b y t h e close a ffi n i ties of t h e
m a m ma l s-wh i c h h e rea soned
i m p l i ed their commo n o ri g i n .

( 1 744- 1 8 29), Fre n c h s o l d i e r a n d

biologist, w a s t h e fou n d e r o f
t h e stu dies of i n vertebrate a n i
ma l s . He stressed t h a t n o a b
solute limits s e p a rated o n e spe
cies from a n oth er and that
s pecies reta i n con sta nt c harac
teristics o n ly i n u n c h a n g i n g en
v i ronments. When the environ
ment does c h a n g e , he a rg u e d ,
t h e i n creased u s e of some or
g a n s a n d the relative d i suse of

BA PTISTE

DE

.r

LAMARCK

others
lead
to
i n h e ri ta b l e
cha nges. T h e g i raffe's long n e c k ,
f o r exa m p le , co u l d be b e s t ex
plained by t h e l o n g -conti n u e d
habit of reach i n g u pward t o
feed on t h e l e a v e s of trees.
By La m a rc k's t h eory, the rela-

tive devel o p m e n t of a n y organ

respo n d s to i t s d eg ree of use.
La marck's
belief that ac
q u i red c h a racteristics c a n be
i n h e rited i s n o l o n g e r a ccepted ,
b u t h is recog n ition of evolution
was of major i m porta n c e .

La m a rck's views s u g g e s ted t h a t

g i raffes reach i n g u pw a rd be
i n c rea s i n g l y
ca m e
l o n ger
n ecked a n d tra n s m i tted t h i s
c h a racteristic t o t h e i r offs pri n g .

T h e conce pts o f evo l ut i o n p ro p osed by Era s m us

D a rw i n a n d by La m a rc k were n ot o n l y rejecte d b u t
were a l so ri d i c u l e d by t h e i r scientific co n te m p o ra ries .
Th is wa s beca u se of t h e excesses of s o m e i n te r p reta
tio n s p ro p osed by La m a rc k a n d h i s d i sci p l es, a n d a l so
beca use m a n ' s eve ry d a y e x p e rie n ce p rovi ded l it t l e
s u p p o rt fo r La m a rc k ' s t h e o ry of s p e c i e s deve l o p m e n t .
I n sp ite of circu m sta n ti a l evi d e n ce, n o o n e h a d yet
seen o n e species t u r n i n to a n ot h e r.

13

THE DISCOVERY THAT LIFE HAD A LONG HISTORY

d i d not com e until t h e e i g h teenth and nineteenth cen

turies, when it b e c a m e genera lly recogni zed t h a t fos
s i l s were the re m a ins of once-l iving ani m a l s and p l a n ts .
Fos s i l s ind i cated that m a ny s p e c i e s h a d beco m e extinct
and that m o st l iving species were of recent origin. If
species were i m m uta b l e, how cou l d these c h anges in
the pattern of l ife be exp lained ? D u ring the nineteenth
centu ry, two o p posing schools of thought developed .
CATASTROPH I STS a ttem pted to

recon c i l e the foss i l record with

the early c h a pters of the Boo k
of Genesis. T h ey reg a rded t h e
Flood of N o a h as the l a s t o f a
series of g reat worldwide ca
tas trophes, each of which d e
stroyed a l l l iving t h i n gs. After
eac h catastrophe, a n ew crea
tion took pla ce, in w h i c h the
earth was repopu lated by ani
m a l s a n d pla nts of new a n d
d ifferent species. These in turn
were d estroyed, and their fos
sil re m a i n s e n t o m bed in t h e
s trata o f the next cataclys m .

GRADUALISTS m a i n ta i n e d that
t h e foss i l record s h owed n o evi
dence of wor l dwide catastro
phes, a l t h ou g h it d i d s h ow m a ny
exa m p les of local eros ion s u rfaces
and c h a n g i n g e n v i ron m e n t s of
roc k d e p o s i t i o n . A l t h o u g h t h ese
c h a n ges a re often m a rked by
t h e c u toff of o n e k i n d of fossil
and its re p l a c e m e n t by a n other,
th i s was a p i e c e m e a l , loca l , i rreg
u la r process, not a worldwide
o n e . New species ori g i n ated,
accord i n g to g r a d u a l ists, by t h e
s l o w m o d i fication of a n cestra l
fo r m s .
G E O R G E S CUVIE R ( 1 769- 1 832 ),
a n outsta n d i n g Fre n c h a n atom ist
and pa l eontologist, studied th e
foss i l ve rte b rates of t h e Pa ris
Ba s i n . T h e s u ccession of d iffer
e n t species see m e d to h i m to
i m p l y a se ries of u n ivers a l ca
tastro p h es, t h e last of w h i c h was
the F lood o f N oa h . C u v i e r be
l i eved that some species s u rvived
to repo p u late the earth w h i l e
other students i n voked a new
creation after each of the catas
tro p hes. As m a n y a s 30 catastro
phes were proposed.

JAMES

H UTTO N

(1726-1797),

Scotti s h p h ys i c i a n , l a n d owner,
a n d a g r i c u l t u r i st, l a i d t h e fou n
dations of m od e r n g eology. He
recog n ized t h a t many rocks
were the res u l t of eros i o n a n d
deposition i n e n v i ro n m e n ts t h a t
h a d modern co u n terparts. T h i s
co ncept
of
uniformitaria nism
s o u g h t to e x p l a i n t h e fea t u res
of t h e earth i n term s o f pres e n t
processes.

(1797-1875), a
Scott i s h s o l d i er, l awyer, a n d
g eologist, p u b l i s h e d T h e Prin
ciples of Ge ology in 1830-33
T h e book, w h i c h ra n to twelve
e d i t i o n s , had e n o r m o u s i n fl u
e n ce . I n it, Lye l l esta b l i s h e d the
s c i e n c e of g e o l ogy, j u st i f y i n g
a n d a m p l i fyi n g H u tto n ' s con
cept of u n i form itaria n ism . Lye l l
f i rst used t h e word "evo l u t i o n "
i n its prese n t se n se.

C HARLE S LYELL

N ew d iscove ri e s l e d to t h e g ra d u a l rejectio n of c a ta s
t ro p h i s m . Fi rst, t h e n u m b e r of catastro p h e s re q u i red to
ex p l a i n t h e fo s s i l record ste a d i l y i n c re a s e d u n t i l t h e
w h o l e syste m b eca m e u n wi e l d ly. I t beca m e c l ea r, a l s o,
t h a t the rock reco rd coul d b e i n te r p reted s a t i sfa c to r i l y
i n te r m s of p r e s e n t- d a y, o b serva b l e g e o l o g i c p rocesses
ra th e r t h a n u n k n own catastro p h e s . In a d d it i o n , t h e "d i ____
1=-uv'_
-' i a l " ro c k s t h a t l a y ove r t h e s u rfa ce of m u c h of Euro p e
a n d North Am e rica a n d we re t h oug h t to b e t he re m a i n s
o f Noa h ' s Flood we re reco g n iz e d a s g l a c i a l d e p osits .
More a n d m o re evi d e n c e of conti n u ity ( o r evo lutio n ) of
fo s s i l s wa s d e m o n strate d . D a rwi n a n d Wa l l a c e p ro p ose d
a n a ccepta b l e m ec h a n i s m fo r t h e p ro c e s s of evo l utio n .
15

D ARWIN'S VOYAGE aboard t h e HMS

Beagle c h a n g e d the worl d ' s viewpoi nt i n regard to
evo l ution a n d the d evelo p m ent of s p ecies. U ntil t h e
p u b l i cati o n of D a rwi n's On t h e O rigin o f Species, i n

CHARLES

1859, t h e i d e a o f evo l utio n was g e n e ra l l y rejecte d .

D a rwin was born a t S h rewsbu ry, E n g l a n d , o n F e b ru

a ry 1 2 , 18 09, t h e s a m e d a y a s Li n c o l n. After two ye a rs
of m e d ica l tra i n i n g a t E d i n b u rg h , h e w e n t to Com
b r i d g e, w h e re h e g ra d ua ted i n 1 83 1 . Afte r h i s g ra d u a
t i o n , D a rw i n was a p pointed n atu ra l ist to t h e Beagle, a
240-to n , 1 0 - g u n brig, which was to u ndertake a s u rvey
voyag e to South Am erica a n d fro m t h e re on a ro u n d
t h e worl d . Th e voy a g e l a sted five ye a rs, a n d t h e i n
s i g h ts D a rw i n g a i n e d d u ri n g t h ose yea rs were t o b e
co m e t h e fo u n d a t i o n of h i s l i fe ' s wo r k . Da rwi n mode
i m p o rta n t co n t ri b u t i o n s to t h e g e o l ogy of South
A m e rica, the orig i n of cora l reefs, the re l a t i o n s h i p s
betwe e n l ivi n g a n d foss i l a n i m a l s , a n d t h e struct u re ,
a d a p tation , a n d geog ra p h ic distri b u t i o n of a n i m a l s .
I t wa s t h ese s t u d i es t h a t l a t e r fo r m e d t h e basis fo r h is
evo l utio n a ry th eory .
DARWIN too k the first volu me
of Lye l l 's n ewly p u b l i s h e d Prin
ciples of Geology o n the voya g e
a n d was d e e p l y i m pressed by i t .
Lye l l a rg u e d tha t t h e ea rth's
s u rface had b e e n s h a p e d by
s u c h n a t u r a l forces as river
erosion , vo l ca n i c e r u p t i o n s , a n d
c h a n g e s i n s e a leve l s . D a rw i n
u s e d s uc h i d e a s i n u n rave l i n g
t h e geo logy o f a re a s h e v i s i t e d ,
a n d t h ey i n fl u e n ced h i s t h i n k
ing a b o u t t h e orig i n o f s pecies .

C h a rles Darw i n , aged 31

16

The route of the HMS Beagle is show n on t h e m a p a bove. I t is

proba b l e that Da rwin contracted C h a g a s ' disease du ring an inl and
journey in South A m erica, m a k ing h i m a s e m i - i nva l i d later.
VERTEBRATE S collec ted
by Darw i n from Arg e n t i n a a n d
elsewhere i n c l u d e d T oxodo n a
h eavy e l e pha n t-sized m a m m a l
t h a t looked m uc h l i ke a rhi noc
eros. D a rw i n concl uded ( wro n g
ly) t h a t i t s h owed t h e two
groups were c l o s e l y re l a t e d .
D a rwi n d iscovered foss i l teeth
of horses that had l ived a t t h e
sa m e t i m e a s Toxodon a n d h a d
b e c o m e extinct with it, a l tho u g h
s u rviving i n ot h e r pa rts of the
worl d . T h i s m a d e the idea of
catastro p h i c w or l d w id e e x t i n c
t i o n a p pear s u s pect.
FO S S I L

T H E S I M I LA R I TY of some fo ssil

verte brates, such a s t h e g i a n t

a rm a d i l lo- l i ke
Glyptodon,
to
form s s t i l l l i v i n g s u g g ested to
Darwi n the idea of descent by
evolution.

THE GALAPAGOS ISLANDS, l oca ted

i n the Pa c ific
a bo ut 600 m i les west of the coast of Ecu a d o r, a re a
d esolate g ro u p of 1 4 roc ky i s l a n d s , representi n g t h e
re m a i n s of exti n ct vo l ca n oe s . T h e i s l a n d s a re s e p a
rated fro m e a c h oth e r by deep water, a n d n o wi n ds o r
ocean c u rre n ts carry s m a l l a n i m a l s o r seeds fro m o n e
t o a noth e r . T h e g e n eral a bsence o f m a m m a l s h a s a l
lowed g i a n t tortoises t o g ra z e i n safety, l i z a r d s t o be
co m e s e a g o i ng, and fi n ches to exist i n n iches that else
w h e re a r e occupied by other species.
D a rw i n d i scovered that ea c h of t h e i s l a n d s , a l t h o u g h
h a vi n g very s i m i l a r c l i mates a n d e n v i ro n m e n ts a n d
b e i n g o n l y a b out 5 0 m i l e s a p a rt, h a s its own fa una
and f l o ra - s i m i l a r to but d isti n ct fro m t h o s e of the
n e i g h b o ri n g i s l a n d s . This s ug g e sted to D a rw i n that
t h e s i m i l a r species m i g h t h a ve d eve l o p e d from a co m
mo n a n ce sto r rat h e r t h a n e a c h h a v i n g b e e n c reated
sepa ra t e l y .
The isl a n ds a r e of rece n t o rig i n , a n d t h ei r fa u n a ,
de rive d fro m t h e S o u t h A m e rica n m a i n l a n d , i l l u strates
co l o n iz a t i o n of, a n d a d o ptio n to, a n e m p ty e n vi ro n
m e n t b y re l a tive ly ra p i d evo l uti o n .

I G U ANAS g row to f o u r feet l o n g

a n d a re fears o m e i n a p p e a ra n ce ,
b u t t h ey are h a r m l es s h e r b i vore s ,
feed i n g o n s e a weeds. F o u nd o n l y
\r'..:'i"J4f!: i n t h e G a l a p a g o s I s l a nd s , t h ey
i n c l ude two rel ated s p e c i e s , o n e
terrestr i a l , a n d t h e o t h e r mari n e.
T h e latter are pow erf u l s w i m m ers,
with we b b ed toes and a f l atte n ed
ta i l to a s s i s t i n s w i m m i n g. Ea c h
i s l a nd h a s i t s o w n ra c e , s h ow i n g
m i nor d i f fere n c e s f ro m o n e gro u p
to a n ot h e r.

w e ig h i n g up
to 250 pou n d s g ra z e on veg eta
tion, fi l l i n g a n i c h e occu pied
i n oth e r p l a c e s by m a m m a l s .
These tortoises a re fou n d o n ly
i n the Galapagos I slands, a n d
each m a j o r isla n d h a s its own
variety. The variation within a
stn g l e sp e ci e s of tortoise is so
s i m i la r to that fou n d betwe e n
species i n t h e Gal a p a g o s fi n c h e s
t h a t Darwi n w rote, " I m ust s u s
pect t h at (th e fi nc h species) a re
o n ly varieties."

G IANT TORTO I S E S

THE F I N C H E S of th e G a l a pa gos

Islands showed a g e n e r al s i m
i l a rity t o o n e a noth er a n d to
th ose of t h e m a i n l a n d of South
A m e rica, b u t t h e fi n c h e s of
each i s l a n d d iffered s l ig htly
fro m those of t h e n ext. T h e
1 3 d iffe re n t s pecies s h owed a
perfect g radatio n , fro m g ro u n d
l i v i n g , seed-eati n g f o r m s with
heavy, large beaks to tree
dwe l l i n g , i n sec t eatin g
fo r m s
w i t h long, p o i n t e d beaks.
-

D a rwi n w o n d ered w h y the

s pecies, i f created s e parately,
rese m b led one a n ot h e r a n d
those of t h e m a i n l a n d o f So u t h
A m erica, w h e re a s birds of t h e
Ca pe Ve rde I s l a n d s , a t t h e
same latitude i n the South At
l a n tic, res e m b l e d t h o s e of Afri
c a . " O n e m i g h t rea l l y fa n cy,"
w rote Da rwi n , " t h a t . . . o n e
s pecies h a d been ta ken a n d
m o d i fi e d f o r d iffere n t e n d s . "
T h e s e a re i l l u s trated o n p . 8 2 .

The G a l a pagos Islands, s h owing route o f H.M.S. Beag l e .

19

THE SEARCH FOR A MECHAN ISM h a d b eg u n . C h a r l e s

D a rwin retu r n e d w i t h the Beagle t o E n g l a n d i n Octo b er

of 1 83 6. The fol l owi n g July h e o p e n e d h i s fi rst n ote
book o n The Transmutatio n of Spe cies. He was t h e n
27 y e a rs o l d . D a rwi n h a d see n how s m a l l v a r i a t i o n s
co u l d b e s e l ected b y a rtifici a l b ree d i n g i n d o m estic
a n im a l s . Could the sa m e tra nsform ations within a spe
cies a lso occur between species so t h a t one u l t i m ately
g ave rise to a noth er? D a rwi n 's observations s u g g ested
that they could, but h e could not vis u a l i z e t h e m e th o d .

ALFRED

RUSSEL

WALLA C E

( 1 8 32- 1 9 1 3),
British
su rveyor
a n d natural ist, i n d e p e n d e n tly
s u ggested the t h eory of natura l
se lection .
A l ready
con v i n ced
of the fact of evo l u t i o n , he con
ceived t h e idea of natura l se
l ection w h i l e l y i n g s i c k with
feve r i n the Moluccas i n Febru
a ry, 1 85 8 . H e reca l l e d the Essay
on Pop ulation, by Robert Ma l
t h u s, w h i c h h e h a d read twe lve
years before. He w rote tha t he
saw its a p p l ica tion to evo l u
t i o n " i n a fla s h o f i n t u i t i o n . "
Wa l l ace w a s a l so a n out
sta n d i ng p i o n ee r in the study of
the geogra p h i c d i stri bution of
a n i m a l s and its s i g n ifica n ce for
the th eory o f evo l u t ion (p. 43).

MALT H U S (1 766-1834)
was a n E n g l i s h c l e rg y m a n a n d
that
eco n o m i st .
U n c o n v i n ced
man is p e rfect, and d i s b e l i e v i n g
t h e p robab i l ity of un ivers a l
p e a c e , e q u a l i ty, a n d p l e n ty,
pred i cted by the p o l i t i ci a n s a n d
u t i l ita rian phi l osophers o f the
e i ghte e n t h
c e n t u ry,
Ma l t h u s
wrote a n a n o nymo u s " Essay on
Popu lation " in 1 798. In it, h e
sla ted t h a t h u ma n pop u l ation
c a n not
e x pa n d
i n defi n itely.
Pop u l ations e x pa n d at a geo
m etric rate of i n crease w i t h
wh ich f o o d s u p p l i e s c a n n ever
keep pace. Fa m i n e , d i sease,
and wa r, Ma l l h u s a rg ue d , wi l l
l i m it t h e i n c rea s i n g size o f h u
m a n popu l a tio n s .
ROBERT

Natura l selection i m plies that

a n c estral g i raffe populations i n
c l uded n e c k s of various le ngths.
More of the l o n g e r - n e c ked gi
raffes s u rvived, a n d t h ey pro
d u ced i n crea s i n g n u m bers of
offs p r i n g that i n h erited t h e i r
pare n ts' lon g e r nec ks.

"I n Octob e r, 1838," wrote D arwi n , " I h a p p e n e d to

read Mal t h u s for amu seme n t. Bei n g wel l pre p a red to
appreciate t h e stru g g l e fo r exi ste n ce , wh i c h eve ry
w h ere goes o n , fro m l o n g conti n u ed o b s e rvati o n of t h e
h a b its of a n i m a l s a n d p l a nts, i t a t o n ce stru c k m e
t h at u n d e r t h e s e circ u m sta n ce s favo r a b l e va r i atio n s
wou l d te n d t o b e p r e s e rved a n d u n favora b l e o n es to
be d e stroye d . T h e re s u l t of t h i s wo u l d be t h e fo r m ation
of a new s p e c i e s . "
D a rwi n ca l l e d t h i s process " n at u r a l s e l ecti o n ." H e
arg u e d t h at t h o s e parti cular i n d ivid u a l s b ette r -a d apted
to t h e i r e n v i ro n me n t wou l d l ive l o n g e r t h a n t h e rest.
S i n ce t h e offs p r i n g wou l d s h a re t h e i r p a rents' c h a r
a cteri stics, over m a n y g e n erati o n s, t h o s e most favo r
a b l e wo u l d te n d to pre d omi nate. D a rwi n m u l l e d ove r
his theory, p re p a ri n g a bri ef outl i n e of it i n 18 4 2 , a n d
a l o n g e r a b stra ct two years l ate r. T h e s e were n ot p u b
l i s h e d u n t i l 1858 ( S ee p . 2 2 ). For t h e n e xt fourteen
years h e g a t h ered data for a fou r- vol ume treatise.
These vo l umes were n ever p u b l i s h e d.
21

THE ORI G I N OF S PE C I ES. I n t h e summer of

1858,

D a rwi n received fro m Alfred Russel Wa l l a ce a m a n

u s c r i pt e n titled " O n the Ten d e n cy of Va rietie s to
D ep a rt I n d e fi n itely fro m the Orig i n a l Typ e . " Wa l l a ce
h a d i n d e p e n d e n tly rea c h e d t h e conclusion that n at u r a l
sel ection h a d pl ayed a m a jo r rol e i n t h e o r i g i n of n ew
species. D i s m aye d , D a rwin offered to with d raw h i s own
m a n uscri pt, but a j o i n t paper by the two men was rea d
b efo re t h e l i n n a e a n Society of lo n d o n o n July 1 , 1858.
On Nove m b e r 24, 1859, D a rwin p u b l is h e d t h e
Origin o f Species-a b rief abstract, as h e c a l led i t , of
h is views . The book created a sensatio n . The f1rst e d i
tion of 1250 copies sold out o n the f1rst d a y of p u b l i
cati o n . Scie n tists w e re at f1rst d ivi ded i n t h e i r views .
Oth e rs, wro n g ly as it n ow a p p e a rs, reg a rd e d the boo k
as a d i rect c h a l l e n g e to rel ig ious b e l i efs. I n s u c h d iverse
f1el d s a s p h i l osophy, h istory, a nth ropology, pol itics,
and sociology, D a rwi n 's book ra ised p rofo u n d q ues
tions. Th e d e b ate was wid espread a n d i ntense .

THE B O O K w a s ca refully w rit

THE ORIGIN OF SPECIES
Bf !lEANS OF NATURAL SELECTION,

Bf

CHARLES DARWIN, K.A.,

_............... _ .. , ....................... ,
...._ .... _.......... ..... _, ___ .......... ""...
_,.. ....
. .......

LONDON:

.lORN MURRAY, ALBEM:ABLJ: IITREET.

22

ten a n d cog e n tly a rgued. T h e

fi rs t four c h a pters described t h e
res u l ts of d o m e s t i c s e l ection
and bre e d i n g a n d then devel
oped t h e idea of natu ra l se
lectio n . T h e fifth c h a pter, de
voted to t h e m echen isms of
va riation and i n h erita n ce , is
t h e o n ly part of the boo k that
has s i nce been d iscredite d . T h e
s ixth to te nth c h a pters d i s
c u ss e d possible o b j e ctions to
the w h o l e idea of evo l u t i o n ,
a n d t h e re m a i n i n g c h a pters
treated poss i b l e evid e n c e for
evo l utio n . S h ow n at left i s t h e
title p a g e of t h e 1 s t edition .

Cartoonist's view, i n 1 8 7 1 , of Charles Darw i n

Hux ley, ( rig h t ) w h o c h a m pioned his teac h i n g s .

( Left ) a n d T. H.

The i m porta n ce of D a rw i n ' s book on Th e Origin of

Species, i s d iffi cult to exa g g era t e . It h a s b e e n ca l l e d t h e
most i m p orta n t b o o k o f t h e n i n etee n t h c e n tury. More
t h a n a n y o t h e r b o o k , b efore or si n c e , i t esta b l i s h e d t h e
t h e ory o f evo luti o n -or "d esc e n t b y m o d ifica t i o n , " a s
Da rw i n c a l l e d it. I n t h is, D a rw i n i n itiated a tra n sform a
t i o n i n t h e stud y o f t h e org a n i c worl d a s profoun d a s
t h a t broug h t a bout i n t h e p h ysica l worl d b y N ewto n ' s
work w i t h g ra vitatio n . I n b i o logy, evo l ut i o n p rovi d e d
a p owerful n ew unify i n g pri n c i p l e , g ivi n g n ew m ea n
i n g a n d i nsig h t t o a ma z e o f confl i c t i n g d a ta a n d a l s o
a n ew i mpetus i n every fi eld of i n quiry .
T h e boo k's i nflu e n c e w a s n o t co n fi n e d t o b i o logy.
I f l ife had a h i s tory, so had m a n , so had l a n g u a g e, so
h a d cul ture . H a d t h ey too evolved? I f there h a d been
org a n i c evo l uti o n , had there a l so b e e n i n o rg a n i c evo
lut i o n in w h i c h the e a rt h , t h e sol a r system , the u n iverse,
m atter, a n d e n ergy itself had u n d e r g o n e c h a n g e?
A wate rs h e d of h u m a n thou g h t was esta b l is h e d .
Ma n's view of the wor l d , of l ife, a n d of h i mself would
n eve r be q u ite the s a m e a g a i n .
23

DARWIN'S THESIS for t h e ong m of new species

rested on th ree essenti a l fou n d ation s-two of t h e m

d e m o n stra b l e fa cts, t h e oth er a n i n ference.
Fi rst, D a rw i n stressed t h at variation existed t h rou g h
o u t t h e wor l d o f livi n g t h i n g s . N o two i n divi d u a l s of
the s a m e s pecies a re exa ctly a l i ke . In size, p ro po rti o n s ,
colori n g , m e ntal a b i l ity, d i s position, physiolog i c a l p ro
cesses, a n d m a ny other ways, each i n d ivid u a l i s u n i q u e .
Furth e r m o re, m a ny o f these a n d oth e r featu res a re
tra n s m itted fro m p a rent to offs p ri n g .
Seco n d ly, D a rwi n a rg u ed that every s pecies over
pro d u c es . More you n g are p ro d u ced t h a n ever su rvive,
for the n u m be r of i n d ivid u a l s in a popul ation s h ow re l
atively l ittle va riatio n . Th is overprod uction exists at
every l evel i n the plant and a n i m a l ki n g d o m s . " Even
s low- b reed i n g m a n , wrote D a rwin, " h as d o u b l e d ( his
n u m bers) i n twenty-five yea rs, a n d at t h i s rate, i n less
than a thousa n d yea rs there wou l d l itera l l y not be
sta n d i n g room fo r h i s progeny. There m u st therefore
be a very hig h rate of m o rta l i ty, a n d t h i s h a s been
s h own to be so. In m a ny s pecies of birds and i n sects,
98 pe rce n t of t h e i n d ivid u a l s die b efo re m a t u rity .
II

11

Swarm of locusts exemplifies a b u n d a n ce of l ivi n g thi n g s .

NATU RAL
SELECTION

GRAZI NG HORSES

Darwi n 's "re c i p e " for evo l ution was the i n teraction of variation,
ove rpo p u lati o n , and n a t u r a l s e lectio n ; i l l u s trated h ere i s evo l u tion
of h orses ( See p . 5 1 ).

Third ly, D a rwin a rg u ed that m a ny c h a racteristics

of p l a nts a n d a n i m a l s were a d a ptati o n s to the e n
vi ro n m ents i n w h i c h t h ey l ived . T h e p rotective col o r i n g
o f m a ny a n i m a l s wa s c l e a rly a d a ptive . Th e teeth of
a n i m a ls we re clea rly related to t h e i r d iet, as were th e
bea ks of b i rd s . Wh a l es, though m a m m a ls, were so
a d a pted to l ife in the seas that they h a d fis h l i ke
bodies. D a rw i n s u g g ested that these h a d come a bout
by natura/ selection of favora b l e d iffere n ces i n a nces
tra l o rg a n i s m s . Those best a d a pted to t h e i r e nviron
ment wou l d s u rvive l o n g e r and so p ro d u c e m o re off
spr i n g tha n those that were not. Th e offs p r i n g wou l d
i n h erit th e i r p a re n ts ' favora b l e c h a racteristics . N e w
species cou l d d evelop i n t h i s way.
Th e vigorous d e bate that fol l owed p ubl ication of
D a rwi n 's book saw g ra d u a l acce pta nce of h is views .
D a rwi n m is u n d e rstood t h e m e c h a n is m of variation a n d
i n h e rita nce, but h i s g e n e ra l theory h a s withstood th e
test of ti m e .
25

THE

LAWS OF I N H E R I TANC E, w h ich h a d e l uded Da r

w i n a n d Wa l l a ce , were d iscovered by Gregor Me n d el

( 1 8 2 2 - 1 8 8 4 ) , a n Au stri a n m o n k . Men d e l is con s i d e re d
t o b e the fo u n d e r o f m o d ern g e n etics. His wor k w a s
p u b l is h e d i n 1866, b u t re m a i n ed g e n e ra l ly u n k n own
u n t i l it was i n d e p e n d ently " red iscovered " by t h ree
biologists in 1900 .
Men d e l d e c i d e d to stud y the i n h erita nce of o n e o r
two re a d i ly recog n i z a b l e c h a ra cters i n t h e g a rd e n
pea-t h e s i z e a n d fo rm o f t h e peas, t h e i r fl ower color,
a n d so o n . He cross- pol l i n ated o n e fo rm with a n other
and t h e n ca refu l ly recorded the resu lts of this over
seve ra l g rowi n g seaso n s . ( p . 60 )
MEN DEL raised peas by cross
pollinating those with smooth,
round peas and those with
shriveled, wrinkled peas. He
discovered that they did not
produce a blending of the par
ent characters, as was g e n e r
ally believed, but that all the
new peas were smooth and
round.
He then used these
seeds to p rodu c e a n ot h e r cro p ,

covered t h a t t h ree-quarters of t h e
n ew g e n e ra t i o n were s m oo t h a nd
ro u n d a n d o n e q u arte r w a s
wr i n kled.
Mendel ca l l ed c h ara cters t h a t
c o u l d be m a s ked i n o n e g e n e ra
t i o n b u t a p pear i n a n ot h e r ( s u c h
a s w r i n k l ed p e a s ) recessive;
t h os e t h a t ove r s h adow t h e m ( s u c h
a s t h e s m o o t h , ro u nd p e a f o r m ) ,

cross-pollinated them, and dis-

Me nde l co n c l uded t h a t t h i s
de layed a p pe a ra n c e o f reces
s i ve c h a ra c t e r s m u st i m p l y t h a t
e a c h c h ara c t e r i s g over n e d by a n
i nde p e n d e n t factor ( w h i c h w e
n o w c a l l a ge n e) a n d t h a t t h e se
m u st b e pa ired in t h e pare n t b u t
n o t i n t h e g a m et e s.
Me nde l m a de t h ree m ajor
d i s cove r i e s : (1) t h a t c h ara cters
are g ove r n ed by p a i red , b u t i n di
vidu a l "fa ctors," (2) that t h e s e
factors m a y b e do m i n a n t o r re c e s
s ive, a n d (3) t h a t t h e s e f a c tors
c o m b i n e , w i t h o u t b l e n di n g , to
prod u c e c h ara cter i s t i c ra t i o s in
the later g e n era t i o n s.

dom inant.

Hugo de Vries, the discoverer of

mutations, and the evening prim
rose that he used in his studies.

MUTATI ON. Me n d e l h a d shown t h a t i n he rita nce was

p a rticu l a r a n d p red i cta b l e . But if this was so, how

cou l d a ny new features ever a rise? Th e a n swer was
fou n d p a rtly i n t h e a ction of n atural sel ection a n d
pa rtly i n the work o f a D utch bota nist.
Hugo d e Vries {18 4 8 -193 5 ) was Professor of Bota ny
at Am sterd a m . He stu died the m ec h a n i s m of i n h erita nce
of c h a racters i n t h e eve n i ng p r i m rose and beca m e in
creasi n g ly s u s picious of t h e t hen c u r rent view that
d iffere n t p a re n t a l c h a ra cters a lways ble n d e d in the
offs p r i n g a n d that all va riatio n s were s m a l l . H e stu d i e d
ove r 5 0 , 0 0 0 p l a n ts, a n d out of t h e i r several h u n d red
thousa n d fl owers, he d i s covered rare exa m p l e s that
were " sports . " They h a d giant size o r dwa rf size, o r
twice the n o r m a l n u m b e r of peta l s . W h e n b re d to
gether, they p rod uced s i m i l a r offs p ri n g . S u c h new
forms de Vries c a l l ed mutants, the c h a nges p rod u c i ng
th e m mutations. I n searc h i n g the l iterature, d e Vries
red iscovered t h e work of Me n d e l . These m utatio n s
p rovided th e g e n u i nely n ew c h a racteristics u p o n which
evol ution by n atura l selection was d e p e n d e nt.
27

T. H. Morgan, a pioneer American geneticist.

Walter S. Sutton, geneticist who

identified role of chromosomes.

T H E N EW SYNTHESIS OF EVOLUTIONARY TH EORY

came i n th e early yea rs of the twentieth centu ry,

m a rked by recog n ition of chromosomes, m i n ute th rea d
l i ke structu res i n t h e c e l l n u c l e u s, a s t h e c a r r i e rs of
h e red ita ry c h a racte rs . Th is d iscove ry, w h i c h a l s o
showed a l inka g e of c h a racters that Me n d e l h a d n o t
suspected, w a s m a d e independently i n 1 902 b y W . S .
Sutton and b y T. Bove ri. T. H . Morg a n ( 1 886- 1 9 4 5 ),
experi m e n t i n g with t h e fru it fl y, Drosophila, d e m o n
strated t h a t t h e g enetic d eterm ina nts were p r e s e n t i n
a d efinite l ine a r order i n t h e c h ro m os o m e s and cou l d
be " mapped . "
Many wo r k e rs b e ca m e co n v i n c e d t h a t i t was s u d d e n ,
sponta n eo u s , l a rg e-sca l e m utations t h a t w e r e t h e rea l
b a s i s of evolut i o n rat h e r t h a n , a s D a rwi n h a d s u g
g este d , t h e m ino r va riations . But why, oth ers o b j ected,
s h o u l d so m a ny c h a ra cters then b e a d a ptive s i n c e
m a ny m utations proved t o b e l eth a l rather t h an b en
efici al? T h e d i scove ry in 1 9 27 t h a t X- rays, te m pe rature
c h a n g es, g a m m a rays, a n d var iou s c h e m i c a l s could
i n d u ce m utati o n s proved that the g re a t m a jority of
them were m i n u te in t h e i r effects and t h erefore were
m o re l i kely to s u rvive .
28

Magnified Segment

C h rom osomes from the fruit fly Drosophila g reatly m a g nified. The
map s h ow s loca t i o n of g e n e s a l o n g part of c h ro m o s o m e l e n g t h .

T h e s i m p l e M e n d e l i a n concept of i n d e p e n d ent, p a r
ti cul ar g e n etic d ev e l o p m e n t h a s g iven way to ac
ce ptan ce of a n i n d ivi d ua l re pre s e n t e d by a ge ne co m
plex i n wh ich g e n e s a re l i n ked a n d i nte ract to g eth e r
T h e curre n t synthetic th eory of evo luti o n is b a s e d
on rig orous sta tistic a l an a l ysi s , stud y of t h e fo ssil
re cord , exp eri mental studies, an d o b s ervat i o n of n at
ural popul a tio n s . I t ac c e p t s as the bas i s for evo l utio n
i n d ivid ual variations, ari si n g fro m mutatio n a n d repro
ductive reco mbin atio n , an d ac ted upo n , fi l tered , c o n
serve d , int e n sifi e d or eli mi n ated by n atural s e l ection.
.

Ge netic variation i n Drosop hila expressed by striking d iffe re nces i n

fo rm. The fl y at left is t h e normal wi l d type.
vestigia l
norm a l

wing s

strap

twisted
abdomen

n o win g s

29

INDI C ATIONS O F E V O LU T ION

The p roof requ i red for a n y p a rticu l a r statem e n t va ries

with the n a t u re of t h e state m e n t . To p rove t h a t 2 + 2
= 4 i nvolves a n a ppeal to reas o n a n d
mathematical
l o g i c . T o p rove t h at a n ath l ete ca n r u n a fou r- m i n ute
m ile i nvolves an a p p e a l to experi m e n t-th e ru n n i n g of
a ca refu l ly m ea s u red d i stance u n d e r s pecified con d i
tio n s a n d with a c c u rate ti m e keepi n g .
B u t n o experi m e nt cou l d prove t h a t t he s a m e ath
l ete ra n a fou r- m i n ute m i l e o n Ju n e 2 0t h a yea r a g o .
P roof of t h a t wou l d i n vo lve a n a p pe a l to the record
boo k s a n d to witnesses. No experi m e n t ca n p rovi de
p roof of p a s t eve n t s . Other k i n d s of evi d e n ce a re
n eeded, a l t h o u g h o b servatio n a n d experi m en ts of ex
isti n g facts a n d p rocesses m ay s u p po rt the pro b a b i l ity
of a p a rticu l a r p a st event . Often p roof i n volves an a p
peal to everyd ay experi e n ce to p rovi d e t h e m o st eco
n o m i c a l expl a n atio n . You cou l d n ot p rove, for ex
a m p l e , that all of the s p a rrows livi n g tod a y desce n d e d
f r o m those l ivi n g t h ree h u n d red. yea rs a g o , b u t t h e b a l
a n ce of e x p e r i e n c e wo u l d s u p po rt t h a t i n terpretati o n .
"I wi l l b e l i eve i n evol ution , " Wi l l i a m Jen n i n g s B rya n
re m a r k e d , " w h e n I c a n sit i n m y g a rd e n a n d see a n
o n io n t u r n i nto a l i ly. " Cl ea rly, i f w e h a d to rely o n
that k i n d o f i n sta n t experience, evol ution co u l d n o t b e
p rove d . B u t n e it h e r c o u l d t h e g rowth of a n o n io n s e e d
i nto a n o n io n be p roved i n sta ntly. I t, too, i s a s l o w
s c a rcely p e rceptible eve n t . W e ca n, h owever, o b s e rve
p o p u latio n s c h a n g i n g a nd c a n a l s o o b s e rve the m e c h
a n is m s by w h i c h s u c h c h a n g e c o m e s a b o u t . The p roof
of evo l utio n a l so l ies in its u n ique position as the o n ly
a d equate expl a n atio n .fo r t h e orig i n of t h e d iverse fea
t u res s hown by l iv i n g th ing s .
30

Frog

tadpoles,

metamorphose

into frogs by the resorption of

the tail, loss of gills, and
growth of lungs and paired
limbs.
Such
radical
change
within a few weeks makes it
less difficult to visualize evolu
tion over countless years.

CONT I N U I TY of l ivi n g thi n g s i s p rovi d e d b y r e p ro

d u ct io n . I n d ivi d u a l s l ive , g row old, a n d d i e, but the i r

k ind i s p e rpetua ted i n the i r offs p ri n g . W e k n ow o f n o
evi d e n ce s u g g e stin g tha t l ivi n g o rg a n i s m s a ri s e i n a n y
othe r w a y tha n fro m p a re n ts of the sa m e s p e c i e s . I t
wo u l d be d iffi c u l t to p rove, fo r exa m p l e , tha t a l l f ro g s
a l ive tod a y m u st have d e s c e n d e d f ro m fro g s tha t l ived
1 , 0 0 0 ye a rs a g o , but a l l of our e x p e r i e n c e s u g g ests
that they ha ve .
Bu t if fro g s a l ways g ive b i rth to fro g s a n d cam e l s
t o ca m e l s , h ow d o n ew k i n ds ( s p e c i es) of an i m a l s eve r
deve l op? Two fea tures of c o n t i n u ity s u g g e s t pos s i b l e
a n sw e r s . Fi rstly, con ti n u ity betwe e n pa r e n t s a n d off
s p r i n g i n volve s both b roa d res e m b l a n c e s a n d in divid
ua l diffe re n c e s a n d varia tio n s . Wha teve r e x p l a n a t i o n
we s e l e ct m u s t ex p l a i n both fea t u r e s .
Se con dly, c o n tin uous c han g e wi thi n the l ifeti m e of a
s i n g l e i n d ivi d ua l a n i m a l i s very g re a t. I f s uch cha n g e s
c a n occur i n on e g e n e ration , it m ay w e l l b e t h a t on e
s p e c i e s could d evelop i n to a n other.
31

U NITY OF LIFE i s s h o w n by t h e fact that, i n s p i te of

t h e ir d iversity of fo r m a n d variety of h a b its , the n e a rl y
1 % m il l io n s p e c i e s of p l a nts a n d a n im a l s a l l s o l ve t h e
b a s i c p ro b l e m s of l ivi n g i n m u c h the s a m e w a y . They
rese m b l e o n e a n othe r i n co m positio n , c e l l u l a r stru ctu r e ,
l i fe p r o c e s s e s , a n d b a s ic patte r n s o f r e p ro d u cti o n ,
a d a pta b i l ity, a n d d eve l o p m e n t . They a l s o s h a re a co m
m o n u n ity in the e n d le s s interd e p e n d e n c e o f a l l l i vi n g
t h in g s . I f e a ch s p e c ie s i s a n e n tirely s e p a rate creati o n ,
why d o a l l sha re thes e b a s i c c o m m o n p r o p e rti es?

CELLULAR STRUCTURE i s a ch ar
acter ist i c of all living material,
and the cells are made of pro
toplasm. Most cells are only a
few thousandths of an i nch in
diameter, but a few are m uch
larger. T he yol ks of b ird eggs
are single cells.

In sp ite of some di fferences,

p lant and ani mal cel l s do have
a s i m ilar bas i c str u ct ure. Even
t h e simplest cel l co n s i s ts of
thousands of di fferent
m ole
cules t hat i nte ract toge t h er in
coordination.
A typ ical ce ll
str u cture i s shown below.

A N I MAL CELL
P ROTO PLASM

Oxyg e n
Ca r bon
Hyd rog e n
N i trog e n
P h o s p ho r u s
Pota s s i u m
Sulfur
Chlorine

32

76.0%
1 0.5%
1 0 . 0%
2 . 5%
0 .3%
0 . 3%
0.2%
0 . 1%

P R O T O P LASM is sh ared by a ll
l i v ing things. It is composed of
a distincti ve
comb ination of
large m o lecules of no n l iv ing
s u bstances, inc l u d ing carbohy
drates, fats, proteins ( including
enzy mes), a n d nucle i c a c i d s t h a t
are organized into a co l l o idal
m i x t u re in wate r. T he u n i q ue
p ropert ies of t h is mater i a l form
the bas is of l i fe.

META B OLISM i n c l u d e s t h e n u
trition , re s p i ra t i o n , s y n t h e s i s , a n d
excretion t h a t i s c h ara c t e r i s t i c o f
a l l l i v i n g t h i n g s . N o n - l i v i n g food
materi a l s a re c o n ve rted i n to t h e
org a n i s m ' s l i v i n g t i s s u e s , c e rta i n
of w h i c h b rea k d o w n t o provide
t h e e n e r g y t h a t i s v i t a l to t h e
processes e s s e n t i a l to l i fe. Meta b
o l i s m i n vo l v e s a c o n s ta n t f l ow of
e n ergy a n d m a t e r i a l s w it h i n a n d
betwee n a n org a n i s m a n d i t s
e n v iro n m e n t .

REPRO D U C T I O N of n e w d u p l i

cate i n d i v i d u a l s i s c h a racteristic
o f a l l l i v i n g t h i n g s . The con
t i n u i t y of form i n volved i n re
prod u c t i o n i s contro l l e d by the
activity

of

s e l f-du p l i ca t i n g

c h e m i c a l structu res c a l l e d g e n e s
( p . 56).

of newborn i n d i v i d
uals i s a co m m o n prope rty of
a l l living things.

GRO WTH

of a l l
living
t h i n g s i nvolves c o n t i n u i n g a d
j u s t m e n t t o a c h a n g i n g e n v i ro n

ADAPTAT I O N

m e n t. I n d i v i du a l
s po n s e s
include

adaptive
rea c t i o n

re
to

sti m u l i , i rritab i l ity, p h ysiolog i c

c h a n g e s , h ea l i n g of I n J U ri e s ,
a n d m ov e m e n t. Over l o n g pe
riods, po p u l a ti o n s show m o re
g e n e r a l a d a ptati o n s .

33

Sponges

Des pite t h ei r diversity, all livi n g t h i n g s s h a re co m m o n propertie s .

T H E NAT U R E OF L I FE i s u n de rs tood l a rg e l y in terms

of a s e r i e s of fu n d a m e n t a l pr o pe rties ( p p . 3 2 -3 3 ) .
N o s i m p l e d e fi n ition o f " l ife " i s pos s i b l e , p a rtly
beca u s e o f its co m p l exity a n d p a rtly b ec a u s e it is
un i q u e . But we c a n d efi n e l ife in term s of s o m e of its
s i m p l e r p r o p e rti e s . livi n g o rg a n i s m s co n s i st of u n i q u e
a n d com p l e x co m b i n ati o n s o f c e rta i n n o n l ivi n g m ate
r i a l s , a r ra n g e d in l a rg e m o l e c u l e s t h a t a re c a p a b l e of
g rowt h , r e p rodu cti o n , a da ptatio n , a n d t h e g a t h e r i n g
a n d u s i n g of exte r n a l foo d a n d e n ergy.
So m e of these i n d ivi d u a l p ro p e rties of l i vi n g thi n g s
a re a l s o p r e s e n t i n n o n l ivi n g th i n g s , b ut o n l y l iv
i n g o rg a n i s m s e x h i b it t h e m a l l s i m u lta n e o u s ly .
O t h e r k i n d s of d efi n it i o n s of l ife a re p o s s i b l e a n d
a re e q ua l l y va l i d . Sc i e n tific d efi n itio n s o r stu d i e s a re
l a rg e l y c o n c e r n e d wit h how l ife d eve l o p e d a n d how
i t i s m a i n ta i n e d . P h i l o s o p h i c a l and re l i g i o u s d ef i n itio n s
a re m o re c o n c e r n e d with why. T h e two k i n d s o f d efi
n itio n s a re u s u a l l y co m p l e m e n ta ry, not c o m p etitive .

34

I NTERDEP E N DENCE is a c h a racte ristic of a l l l iv i n g

thin g s . Eve ry in d ivid u a l exi sts a s p a rt o f a n i n terbreed
ing po p u l atio n that co n s i sts of m a ny g e n et i c a l l y simi
l a r in d ivi d u a l s . These po p u l ati o n s of i n d ivi d u a l s exist
wi thin c o m m u n ities of m a ny s pecies that i n te r a ct with
o n e a n othe r a s p rey a n d p re d ator, host a n d p a ra site,
co n s u m e r a n d p rod u c e r, a n d co m petitors fo r s p a c e or
food . The intera cti o n cuts a c ross the m a j o r divi s i o n s of
p l a n ts a n d a n im a l s ; thus , trees she l t e r b i rd s , i n sects
fe rtil i z e flowers, he r b ivo res co n s u m e g ra s s , fish s u p
port p a r a s ites, s e a a ne m o n e s shelter c l o w n fi s h , etc .
i n teract
with
C O M M UN I T I ES
t h e i r p h y s i c a l e n v i ro n m en t , c o n
stitu t i n g a n ecosyste m . C h a n g e s
i n ra i n fo l l , t e m p e ra t u re ra n g e ,
s o i l type , e l e va t i o n , l a t i tude,
depth of sea water, sedi m e n t i n
strea m s , a nd cou n t l e s s o t h e r
p h y s i c a l factors a l l i n f l u e n ce t h e

deve l o p m e n t
of
com m u n ities.
Orga n i s m s i n t u r n m a y m odify
t h e i r e n v i ro n m e n t, crea t i n g lo
c a l s h ade in forests , m odifyi n g
a n d e n r i c h i n g s o i l s , p reve n t i n g
eros i o n , a nd i n m a n y o t h e r
w a y s . T h i s i n te rde p e n de n c e p ro
vides i m porta n t data .

O x y g e n a n d c a r b o n c y c l e s s h ow i n terde p e n d e n ce of a l l l i f e .
CARBON- HYDROGEN
OXYGEN CYCLE
( on land )

T H E S I M I LA R I T I E S t ha t exist betwee n l ivi n g o rg a n is m s

a t a l l leve l s h a v e c e rta i n i m p l icati o n s . Offs p r i n g o f t h e

sa m e parents h a v e a m o r e o r l e s s close rese m b l a n ce to
o n e a n oth e r a n d to th e i r p a re n t s . Althou g h e a c h i n d iv i d
u a l is u n i q u e , m e m b e rs o f the sa m e s p e c i e s s h a re " o b v i
o u s " co m m o n featu res t h a t a re c o n s e rved a n d p e rpetu
ated i n re p rod uctio n . W e d o n ot ha ve tro u b l e recog n i z i n g
a l io n , fo r exa m p l e - o r even a d o g , d e s p ite t h e m a n y
va riati o n s tha t d o m estic b reed i n g h a s p rod uced i n d og s .
o f rese m bl a nce a l so

exist a m o n g related species.

Ocelots, p u m a s , bobcats, a n d
dom estic cats, f o r exa m p l e , a l l
have ce.rta i n b a s i c c h a ra cteris
tics i n co m m o n , and these a re
recog n ized in a n i ma l c l a ssifica
tion (taxono my) by g rou p i n g
t h e m a l l togeth er i n the s a m e
g e n u s-Fe lis . B u t g e n e ra , too,

DEGREES

e x h i b i t d e g rees of rese m bl a n ce
so that we c a n g rou p t h e m i n tq
fa m i l ies of s i m i l a r m e m bers.
S i m i l a r fa m i l ies a re g roup ed
i n to ord e rs, orders i n to classes,
a n d c l a sses i n to p hyla. Each
" h i g h er" g ro u p thus i n c l u d e s
m o r e forms, a n d these h a ve
prog re ssively fewer features i n
com m o n ( p . 1 1 ) .

The o rioles be low belong to a sing le g e n us, Icterus. They have dif
ferent. colors a n d geog ra p h i c ra nges, b ut they share m a ny co m m o n
features. T h e y a re m e m bers of t h e s a m e f a m i l y a s b l a c k b i r d s .

To rosa u r u s

Trice ra tops

Arrhin oceratops

T h e s e h o r ned c e ra to p s i a n d i no s a u r s s h ow how deg rees of rese m

,
b l a n ce s u g g e s t evo l u t io n a ry re l a ti o n s h i ps . T h e g e o l o g i c prog re s
s i o n i s a r ra n g ed f r o m b o t t o m to t o p . ( After C o l bert. )

T h e m e a n i n g of t h e va r i o u s d e g re e s of res e m b l a n ce
wa s a t fi rst t h o u g h t to l i e i n t h e i r a p p ro x i m a t i o n to
the a rc h e t y p e o r i d e a l fo r m , u p o n w h i c h e a c h s p e
c i e s h a d b e e n " d e s i g n e d " o r p l a n n e d . But to l ater stu
de n ts, t h es e c l u stered re lat i o n shi ps, often p i ctured a s
t h e b r a n c h e s of a t r e e ( a s a b ove ) s u g g ested o n ly d e
g r ees of r e l a t i o n s h i p , a l tho u g h the c l a ssification itself
wa s esta b l i s h e d before t h i s w a s recog n i zed ( p p . 1 0 1 1 ) . J ust a s t h e b r a n ches of a tree g row by conti n u
o u s d evel o p m e n t fro m a seed , e a c h b ra n c h bei n g
fo r m e d b y s l o w a n d a l m o s t i m pe rcepti b l e m o d i fication
of e a r l i e r b ra n ches fro m an i n iti a l ste m , so t h e b r a n c h
i n g pattern o f c l a s s i fi c a t i o n s u g g ested a co m m o n ori
g i n . T h e b r a n c h e s r e p re s e n te d d e g re e s of r e l a t i o n s h i p
t o t h e o rg a n i s m s o f t h e c e n t r a l a n cestra l ste m .
37

DEGREES OF S IM I LARITY between l i v i n g t h i n g s

a re

reflected by va r i o u s featu res . The overa l l form a n d

structure ( m o rpho l o g y ) o f a l l creatu res s how va ryi n g
d e g rees o f s i m i l a rity. W hen we s pea k o f a " d eer, " w e
th i n k of a p a rt ic u l a r k i n d o f a n i m a l, b u t the d eer fa m
i ly c o n ta i n s 2 0 d ifferent g e n e r a a n d m a ny species . Al
though they d i ffer in s iz e , a ntl ers, colo r, a n d g eo
g r a p h i c d i st r i b u t i o n , all m e m bers of the deer fa m i ly
sha re b a s i c features . Their s keleton s rese m b l e o n e a n
other, b o n e fo r b o n e ; their i nte r n a l o rg a n s a re si m i l a r;
a n d t h ey d i s p l a y m a ny s i m i l a r b e h avio r a l c h a racteris
t i c s . Thi s co m p rehe n s ive s i m i l a rity, s howi n g a u nity of
b a s i c fo r m but a d ive rs ity of i n d ivid u a l pattern, sug
g e sts the i r d e rivation f r o m a co m m on a n cesto r that pos
s e s s e d these co m mo n featu res .
DEVELOP
EMBRYO N I C
THE
M E N T of m a n y s pecies s h ows

startl i n g s i m i larities, even in

forms that h ave few rese m
b l a n ce s as a d u lts. T h u s a m a n ,
a p i g , a n d a c h ic ken have a
g e n e ra l s i m i l a r i ty d u r i n g their
develop m e n t. I f each s pecies is
e n tirely d i s ti nct fro m every

o t h e r species, i t m a kes n o s e n s e
t h a t they s h o u l d h ave s u c h e m
b ryo n i c rese m b l a n ces a n d t h e n
l o s e t h e m i n a d u l t l i fe . A l t h o u g h
t h i s e m b ryo n i c s i m i la rity i s less
than was c l a i m e d by late n i ne
tee n t h c e n t u ry zoologists, it is
a n i n d ication and an i m print of
their re mote k i n s h i p.

STRU CTU RES

HOMOLO G O U S S T R U CTURES i n

A N A LO G O U S

m a n y orga n i s m s s u g g est t h e i r
derivation f ro m com m o n a n ces
tors. The s ke l etons of cats,
horses, whales, bats, m i ce, a n d
m e n , for exa m p le, a l l have a n
ess e n t i a l l y s i m i l a r for m . The
structure of t h e ve rte brae a n d
t h e fused bones of t h e s k u l l a re
s i m i la r i n every verte brate,
fro m fish to m e n . So a re the re
lated n e rves, m uscles, a n d b lood
vessels. In less c losely re lated
species, h o mology i s less well
m a rked, s u g g esti n g their m ore
d ista n t com m u n ity of ori g i n .

s h ow a s i m i l a rity of f u n ction but

not of d eta i l e d structu re . T h e
w i n g s of a n i n sect perform the
sa m e f u n ction as those of a
b i rd , but they h ave a ve ry d if
fere n t structure . S u c h d i ffer
e n ces res u l t n o t from i n h eri
t a n ce fro m a co m m o n a n cestor
b u t from a d a ptation to s i m i l a r
If
e n v i ro n m e n t a l
co n d i ti o n s .
each s pecies h a d a s e p a ra te
orig i n , t h e n a n a lo g o u s s truc
t u res s h o u ld be m o re co m m o n
t h a n h o m o l ogous s tructures, b u t
t h e reverse i s true.

VESTIGIAL STRUCTUR E S deve l

o p w h e n a n org a n i s reta i n ed
eve n thoug h its ori g i n a l f u n c
t i o n i s reduced or lost. S u c h
structu res a re fou n d i n a l l a n i
m a l s . I n m a n , t h e e a r m u scles
a re usually nonfu n ctional, b u t i n
other a n i mals, s u c h a s t h e dog,
these m u scles m ove t h e ears
and d i rect them towa rd particu
l a r sou n d s . T h e h u m a n a ppen
d i x h a s n o obvious f u n ction
a n d i s a n u i sa n ce, b u t in other
a n i ma l s, t h e a p pen d i x i s m ore
stro n g l y deve l o ped a n d serves a n

i m porta n t d i g estive f u n c t i o n . I n
w h o l e s a n d i n s o m e s n a kes, ves
t i g i a l h i n d l i m bs o re preserved,
s u g g esti n g t h a t they o re t h e
rem m o n ts of a n cestra l structures.

B I O C H EMI CAL S I M I LARITIES a lso e x i s t betwee n re

l a t e d o rg a n is m s . The most stri k i n g feature of these si m i

l a rities is the way they confirm i n d e p e n d e ntly the vari
o u s g ro u p i n g s of p l a nts and a n i m a ls that were esta b
l i s h e d o n t h e b a sis of thei r overa l l for m . T h i s i m plies
t hat t h e c l a ssification that h a s been d eveloped (pp. 3 63 7) is n ot w h o l l y a rtifi c i a l but reflects the a n cestra l - d e
scen d a n t ( p hylog e n etic) relatio nsh ips o f org a n is m s .
Oth e r m o re g e n eral b i o c h e m i c a l si m i l a rities i m ply
t h e co m m o n k i n s h i p of all o rg a n i s m s . These i n c l u d e the
use of n u c l e i c acids as a g ents of heredity (p. 68), the
use of a p a rticu l a r phosph ate, ATP, i n e n ergy tra n sfer,
a n d t h e use by p l a n ts with c h l o ro p hyl l of this g reen
pig me n t a s a cata lyst i n p h otosynthesis.
BLOOD
P I GMENTS
d i ffer in
d iffere n t a n i ma l g roups. I n
ve rteb ra tes a n d some other a n i
m a l s , the blood has a r e d pig
ment, he moglobi n , which h a s a
res piratory f u n ction i n carrying
blood from the lungs or g i l l s
throug hout the body. I n a l l ar
th ropods, the respi ratory pig
ment i s a b l u e copper com-

pou nd, ca l l e d h e mocya n i n ; i n

m a r i n e wo rms, a g reen i ron
com pou n d , ca l l e d c h l o rocruo
rine. These p i g ment s i m i l a r i
ties con fi r m the re l a t i o n s h i ps
between m e m bers of t h e g ro u p s
esta b l i s h ed by o t h e r criteri a .
Prote i n s o f e a c h s pecies, a l
thoug h d i s t i n ct, s h ow co m p ar
a b le d e g rees of s i m i l a rity.

A serolog ical test (p. 4 1 ) i s made by m i x i n g seru m and a n ti seru m

a n d record i n g t h e h i g h e st d i l ution of seru m t h a t wi l l s t i l l g i ve a
wh ite r i n g of p rec i p itate. (After Boyd en.)
HOMOlOGO U S
TEST
beef se r u m a n d
a n ti - beef seru m

1 000

1 5 00

2000 3000 4000

6 0 0 0 8 000 con trol

TI D
DU D
E n d Point

HETEROlOGOU S
TEST
sheep serum a n d
a n ti- beef s e r u m

40

SEROLOG I CA L S I M I LARIT I E S a re m e a s u re d b y i m m u n ity

tests . If b l o o d fro m o n e s p e c i e s , s u ch as a cow, i s i n
jected i n to the b l o o d s t re a m o f a n othe r, s a y a g u i n e a
p i g , the g u i n e a p i g p ro d uces a p re c i p i t a t e , a n a n t i
seru m , tha t i m m u n iz e s i t a g a i n st cow ' s b l o o d . Whe n
this a n ti - cow s e r u m i s m i x e d with the b l o o d of o the r
a n i m a l s , it p ro d uces p reci p i ta t e s of va ryi n g i n te n s i ty
tha t corres p o n d to the n e a r n e s s of the o the r s p e c i e s i n
the sche m e o f c l a s s i fi c a t i o n . Th u s , a n t i - cow s e r u m
g i ves 1 0 0 p e rc e n t p re c i p i t a tio n w i th the b l o o d of a n
o t h e r cow, 48 p e rce n t w i th a she e p , a n d 2 4 p e rce n t
w ith a p i g . Thi s b i o c h e m i ca l i n d i c a t i o n o f co m m o n
a n ce s t ry i s a m ethod o f c l a s s i fi c a t i o n c o n fi r m i n g wha t
w a s esta b l i she d i n d e pe n d e n t l y b y c o m p a ra t i ve a n a
tom ica l s t u d i e s .
Serolog i c a l tests m a d e w i t h a n t i- h u m a n s e r u m g ive varyi n g per
c e n t a g e s of p r e c i p i ta t i o n . T h i s refl ects t h e q u a n t i ta t i ve deg re e s of
s i m i l a r ity betwee n m a n a n d ot h e r s pe c i e s .

G o ri l l a

O ra n g u ta n

K a n g a ro o

to the pa rti c u l a r e n v i ro n m e n ts i n
w h i c h they l i ve a re s h own b y a l l l ivi n g creatu res. S o m e
that a re so g e n eral t h ey may b e overlooked e a s i l y i n
c l u d e t h e won d e rfu l ly efficient b u t d isti n ct wi n g struc
ture of i n sects, b a ts, and b i rd s ( a l so those of t h e ex
tinct pterodactyls), the s h a pe a n d structure of fi s h , the
speci a l i z e d ste m s of d esert cacti, a n d cou ntl ess oth e r s .
Sti l l o t h e r a d a ptati o n s are m o re specific . O f the m a ny
exa m p les a m o n g b i rds, those of the wood pec kers were
fi rst d escri b e d by C h a rles D a rwi n .
Ada ptation i s so w i d es p read i n both p l a nts a n d
a n i m a l s th at, althoug h n ot a proof o f evo l ution, i t
sugg ests that n atural sel ectio n i s a very p ro b a b l e ex
p l a n ation fo r o rg a n i c d iversity.

ADAPTATI O N S

PROTECTIVE fo rm and colora

tion a re adaptatio n s s h own by
many a n i m als . T h e p u pa e of
some i nsects rese m b le thorns or
twigs. Others m i m i c less v u l n er
able species by color rese m -

Wood pecker

42

b l a n ces. The color of some a n i

m a l s , s u c h a s t h e c h a meleon,
changes with t h e color of the
backgro u n d . Experi m e n ts h ave
s h own the s u rviva l value of t h i s
coloration ( p. 8 4 ) .

A wood pecker h a s two l a rg e

toes d i rected backward so t h a t
i t s f o o t forms a n a n chorlike
h o l d . I t s stiff t a i l feath ers form
a prop a s t h e b i rd chisels with
its powerful beak. It extracts
insects with its l o n g , barbed
ton g u e . All of t h e 1 79 species
of wood pec kers h ave esse n
t i a l ly s i m i l a r structure .

-=--- Salomon Islands

New G & i nea ( 5 20 species

of b i rds)

. :.
.:
'

Fiji

Island

( 1 26)

1541

Henderson

Island

(4)

TH E NUMBES OF ISLA N D S P E CI E S a n d
t h e i r res e m b l a n c e to those o f t h e m a i n l a n d
decrea s e wit h i n crea s i n g d i s ta n c e f ro m t h e
la n d . T h e n u m be r of m a m m a l s p e c i e s s hows a
s i m i l ar decrea se, s u g g e s t i n g t h a t t h e species
were deri ved from t h o se o n t h e m a i n l a n d .

P R E S E N T LI M ITED D I S TR I BUT I O N O F MANY S P E C I ES, s u c h a s ta p i rs,

can be i n terpreted o n ly on t h e a s s u m ption t h a t they a re descenda nts
of m ore w i d e sprea d foss i l a n cestors, some of w h i c h hove bee n fou n d
i n i n termed iate a reas.

T. indicus

llr.'-.... ..
+

GEOGRAP H I C

PlPlieoiscene
tocenespecispecies es

. /
.

After De leer

D I STRIBUTIO N of many p lants a n d

a n i m a ls s h ows features t h a t c a n b e a ccounted fo r o n ly

by s u p p os i n g t h a t t h ey a re t h e d escen d a nts of co m
m o n a n cesto rs . T h e fa u n a s o f t h e G a l a pagos a n d C a pe
Verde I s l a n d s were m a jo r clues i n D a rwi n 's d evelop
m e n t of an evo l utio n a ry theory (pp. 18- 19) .
Alfred Russel Wa l l a ce noted t h a t l a rg e r g roups,
s u c h as orders, h ave a w i d e r g e o g r a p h i c d i stri bution
than do s m a l l e r g roups, s u c h a s fa m i lies o r g e n era .
S pecies most s i m i l a r a re fou n d i n a d j a c e n t a re a s, s u g
gesting their evo l ution fro m co m m o n a nc esto r s.
43

..

LIV I NG SPEC I ES of p l a n t s a n d a n i m a l s a re c h a r a c

t e r i z e d by t h e i r con sta n cy of g e n e r a l fo rm a n d t h e i r
g reat ran g e o f i n d ivi d u a l variatio n . Each s p e c i es
b re e d s " true " a n d i s r e p ro d u ctively i s o l ated from o t h e r
s pecies, even t h o s e t h a t are closely si m i l a r . Y e t n o two
i n d ivi d u a l s of t h e s a m e species a re i d e ntica l . We n ow
recog n i z e t h a t t h e i n h erited c h a racteristics of a l l l iv
i n g t h i n g s a re contro l l e d by t h e i r g e n e s a n d c h ro m o
s o m e s a n d t h a t t h e s e stru ctu res u n d e r g o s p o n ta n eo u s
m u tatio n s ( p . 7 4 ) . T h i s i n pu t of n ew c h a racteristics
m e a n s that ove r a long period of ti m e species a re not
Axed e ntities a s o n c e s u p posed . Both in n a t u r e a n d in
ca ptivity, we see evid e n c e of va riations with i n a spe
cies, s u g g esti n g t h e i r evo l utio n a ry c a p a c ity.
S E L E C T I V E B R E E D ING of do
m e s t i c p l a n ts a nd a n i m a l s i ndi
cates t h e g reat v a r i a b i l ity of
many s pe c i e s . I l l u stra ted h e re ,
a re dog s of t h e s a m e s pe c i e s .

Basset H o u n d

Dach s h u n d

Vendee Hound

--

Canis fomiliaris leineri

T h i s s u g g ested to D a rw i n t h a t
natural selection might be ana
l og o u s i n i t s a c t i o n t o ( a r t i f i c i a l )
do m e s t i c s e l e c t i o n a s a n a g e n t
of c h a n g e.

Egyptian Grey h o u n d

S t . H u bert Hou n d

Sa l u k i

Beagle

Talbot H o u n d

Afg h a n H o u n d

Sleuth Hound

PO PULAT I O N S s h o w loca l v a ri
a t i o n i n n a t u re . T h e s m a l l e s t
u n i ts , ca l l e d d e m e s, a re o n ly
partly
i s o l a te d
populations,
with i n w h i c h t h e re i s c l o s e g e n
e t i c s i m i l a r ity. V a r i a t i o n b e
twe e n d e m e s i s ofte n ra ndo m ,
b u t betwee n s o m e i t i s n o n -

ra ndo m , f or m i n g g raded c l i n e s
t h a t m a y s h ow corre la t i o n w i t h
differe n t
e c o l og i c
c o ndit ions.
Th u s loca l r a c e s o r s u b sp e cies
d e v e l o p , e a c h a d a p t e d to t h e
c o n di t i o n s of a part i c u l a r a rea
a n d i n terg rad i n g w i t h o n e a n
o t h e r o n l y i n o v e r l a p p i n g a re a s .

P. major m a jor

P.

rnajqr

intermedius

P. major minor

Z o n e of Overl ap
Z o n e of Ove r l a p

Geog ra p h i c a l D i s t r i b u t i o n of t h e Great Ti t .
Th e G re a t T i t of Europe a n d
As i a s h ow s deve l o p m e n t o f typ
ical geog ra p h i c r a c e s or va r i
e t i e s . T h e m o s t w i d e s p read race
i s P o r us major major, e x t e n d i n g
from B r i ta i n to Ea s t A s i a . P .
c i n e r e u s a n d P. m inor h a ve
m o re restri cted ra n g e s .
T h e i n te r b reedi n g i n I ra n o f
P . major w i t h t h e c e n t ra l As i a n
a n d I n dia n v a r i ety P . cin ere us
g ives a f o u r t h v a r i e ty-P. i n te r -

( Af t e r De B e e r )

medius.
I nterbree d i n g
of P.
cinereus a n d P. m inor in I n d o
c h i n a g i v e s a fi ft h , P . com m ix
I u s . B u t P. major a nd P . c i n er
eus occ u r tog e t h e r w i t h o u t i n

te r b ree d i n g
in
n o rt h c e n tra l
A s i a , a s do P. m i n or a n d P .
m a jor i n n o rt h e a stern C h i n a .
R e p rodu ctive i s o l a t i o n b e t w e e n
g e o g r a p h i c races s u g g e s ts o n e
mecha n i s m for the formation
o f n e w s pe c i e s ( p . 76 ) .

45

CHANGES IN SPECIES h a ve bee n observed eve n d u r

i n g the l i m ited t i m e that accu rate observatio n s h a ve

been reco rded . S o m e d isease- p ro d u c i n g bacteria h a ve
been s uccessfu l l y treated with d ru g s, but o n e of th e
s i de effects of this med icatio n h a s been the deve l o p
m e n t o f v a r i o u s dru g resista nt stra i n s of the bacte ri a .
Escherichia coli i s a co m m o n bacte r i u m that h a s d e
ve l o p e d p o p u l a tio n s e n ti re l y resista n t to strepto m y
ci n . T h ese resista n t g ro u p s a rise fro m m u ta tio n s . W h e n
t h ey a p p e a r, t h ey a re t h e o n l y g ro u p a b l e to s u rvive
a n d m u l ti p l y .

A l t h o u g h m o s t evo l u ti o n i s
pro b a b l y t h e res u l t o f s low,
c u m u l a tive c h a n ge , by t h i s
process of " prea d a ptatio n " i n
w h i c h a m u ta ti o n " e n co u n ters "
a favora b l e e n v i ro n m e n t, t h e
w h o l e c h a racter of a popu l a
t i o n m a y c h a n g e very ra p i d ly.
By a s i m i la r proces s , so m e s pe
cies of destructive i n sects have
d eveloped an i m m u n i ty to vari
ous i nsecti c i d e s . Sca l e i n sects of
t h e citrus reg i o n s of Ca l i fornia
have beco m e i n crea s i n g ly re
s i sta n t t o hyd rocya n i c a c i d , for
exa m ple.

sca le

ra ntii.

i nsect,

A o n idella

a u

MELA N I SM
ha s
been observed i n s cores o f spe
cies of moths d u ring the past
century. In
i n d u strial a reas,
many species have beco m e p ro
gressively darker, o r even b l a c k ,
w h i l e m e m bers of t h e s a m e s p e
c i e s i n rura l a reas re m a i n l i g h t
colored. T h i s d e m o n strates h ow
plastic a n d c h a ngeable many
species a re, eve n over short pe
riods of t i m e . The m e c h a n i s m
o f t h i s c h a n g e i s d iscussed o n
p. 84 . I n t h e p h otogra p h b e
l o w , t h e l i c h e n - covered tree
tru n k prov i d es concea l m e n t for
Pep pered
the
l i g h t-colored
Moth but m a kes the d a rker, i n
d u stri a l m e l a n i c form co n s pi c u
ous.
The
soot-covered
tree
tru n k from an i n d u s tri a l a re a
concea l s t h e d a r k form of t h e
Peppered M o t h b u t ma kes t h e
l i g h t form c o n s p i cuous.
I N DUSTRIAL

cred i t : H. B. D . Kettlewe l l

Peppered moth, Bis t o n b e t u la r ia, show i n g l i g h t a n d d a r k fo r m s

o n two d i ffere n t b a c k g rou n d s .

FOSS I L S PEC I ES a re d ifficu l t to reco g n ize b ec a use

the

test of re pro d u ctive isolation, by w h i c h l ivi n g species

a re d isti n g u i s h e d , c a n n ot be used . But we c a n reco g
n iz e i n fossils t h e s a m e d e g rees o f structu ral d iffere n c e
a s betwe e n related l ivi n g species . We c a n a l s o recog
n iz e i n d e p e n d e ntly deve l o p i n g fossil g ro u ps, a n d
t h ese, b y d efi n itio n , ca n be reg a rd e d a s s p e c i e s .
T h e foss i l record a l l ows us to observ e c h a n g es ove r
fa r l o n g e r periods of ti m e th a n a re eve r a va i l a b l e i n
livi n g p o p u l atio n s . I n fossils, w e c a n reco g n i z e evo l u
tio n i n a ctio n . Althou g h they tel l us l ittl e a bout t h e de
t a i l e d m ec h a n i s m s of c h a n g e, fossi l s d o p rovi de pow
erfu l evi d e n ce t h a t evo l ution h a s occu rred .

FORAMI N I FERA a re m i c roscopic

protozoa n s , most of w h i c h se
crete a s h e l l . I l l u strated i s a
m a r i n e g e n u s , Textularia, stud
ied i n rocks of Tert i a ry a g e (p.
98) i n New Zeala n d . When
traced t h roug h a period repre-

s e n te d by the a cc u m u l a t i o n of
500 feet of strata, t h ere is a
m a rked c h a n g e i n s h a pe for
each p o p u l a tion . Two s pecies
a re recog n ized. Horizo n t a l l i n e s
represe n t s ta n d a rd d ev i a t i o n
f o r eac h . ( After Ke n n ett. )

Fossil b iva lve cl a m s ( b e low) , fro m roc k s of P e n nsy l

va n ia n a n d P e r m i a n a g e ( p . 98 ) o f t h e m id c o n ti n e n t re
g ion of t h e U n ited States, s h ow successive d evel o p m ent
by desce n t of closely s i m i l a r s pecies of the g e n u s Mya
/in a . Each of the n u m bers o n g ra p hs represents a d iffe r
ent species, l isted b e l ow.
Form Ratio
of a n g les b/ a

Ratio of length

FIGURE A

to He i g h t

0.9

1 .0

1 .1

1 .2

Misso u r i a n

1.

M y a l i n a cop h a

2 . M . lepta
2 . M. iepta
M. w y o m i n g e n s is

3.

Desmoinesian

4. M . m iope t i n a

5 . M . p l iap e t i n a

6.
7.

M . cop e i

9.

M. p e t i n a

M. arbo/a

Ato k a n

8 . M . g /oss idea

1 0 . M . a v i c u loides

F I GURE B

Fi g u re A is a plot of t h e form
ratio of length to h e i g h t of the
shells, p lotted a g a i n s t the ra tio
of the a n g le b to the a n g le a
(see d ia g ra m ) . T h e rig h t - h a n d
l i n e represents i n - l i n e evol u t i o n ,
w h e r e new s pecies a rise by s u c
cessive modification of earlier
populations. T h e left- h a n d l i n e
re p res e n ts speciation by bra n c h -

i n g or s p l i tt i n g ra t h e r t h a n by
co n t i n uous c h a n ge.
Fig ure B s h ow s i n ferred evo l u
tionary descent a n d re lation
ships (phyloge ny) of species of
Myali n a . N u m bers refer to the
sa m e species a s t h os e i n Fig. A.
T h e n a mes a re those of s u cces
sive roc k d iv i s i o n s . (After New
ell and Moore.)

49

H I G H E R TAXA ( g e ne ra , fa m i l ies, etc.) of a n i m a l s

a n d p l a nts a re fou n d i n t h e foss i l record a l so t o a ri s e

by d e s c e n t w i t h s l ow mod ification from a n cestra l fo r m s .
T h i s is evol utio n . The fos s i l record provi d e s repeate d
evi d e n c e t h a t it i s t h e n o r m a l m ethod b y w h i c h n ew
g ro u ps of o rg a n i s m s orig i n ate .
C E R A T O PS I AN D I N OS A U RS (a l l
dra w n t o s a m e s c a l e ) l i ved i n t h e
Creta c e o u s Period ( p. 98), 7 0 m i l
l i o n years a g o. T h ey s h ow a n
overa l l i n crea s e i n s i z e a n d i n
t h e re l a t i ve di m e n s i o n s a n d co m

p l e x i ty o f t h e b o n y a r m or t h a t
covered t h e i r h ead a n d n e c k ,
Triceratops rea c h ed a l e n g t h o f
2 4 f e e t a n d we i g h e d u p to e i g h t
to n s. O n ly t h re e g e n e ra are
s h own. (Afte r Co l b e rt.)

O l D WOR L D

Equus

Styfohipparion

Hipparion

Pfio hippus

Anchitherium

GRAZERS
I

BROWS E R S

Orohippus

To R h i noceroses

1)

Epihtppus

H ORS E S p rovide a c l a s s i c e x a m

p l e of t h e evo l u t i o n of n e w g e n
e r a f r o m e a r l i e r o n e s over a
period of 70 m i l l i o n yea r s .

After S i m p s o
refl e cted
m odi fi c a t i o n s
L a te r
c h a n g e in d i e t from brow s i n g
to g r a z i n g . R i g h t u p p e r m o l a r
t o o t h s u rf a c e s a re s h o w n .

51

"MI S S I N G L I N KS," a s e v i de n ce t h a t o n e g ro u p de

veloped from a n o t h e r, were often d e m a n d ed by o p po

n ents of evo l uti o n i n e a r l i e r y e a r s of t h e evoluti o n a ry
d e bate. At t h e ti m e of t h e p u b l ication of On the Origin
of Species, very few of these tra n sitio n a l fo r m s were
k n own , b u t m a ny h ave since b e e n d iscove red . T h ey
b r i d g e m a ny of t h e m a jo r g ro u p s of existi n g o rg a n is m s .
I n th e ve rtebrates, for exa m p le, t h e re a re tra ns itio n a l
fo r m s b etwe e n fi s h a n d a m p h i b i a , a m p h i b i a a n d rep
tiles, reptiles and b i rds, and repti l e s and m a m m a l s .
They i n d icate t h a t these m a jo r g rou ps, d isti nct a n d
s epa rate i n l iv i n g forms, a rose fro m for m s th a t s howed
s o m e c h a racters i nte rmed iate betwe e n two g ro u p s a n d
oth e rs now restricted t o j u st o n e .
ARC HAEO PTERYX, a n a n cestra l

fos s i l b i rd from t h e J u ras si c of

Germa ny, had m a n y features of
the rept i l i a n g rou p fro m w hi ch
i t developed. Althou g h it had
the feathers of a b i rd , i t h a d a
repti l e l i ke too t h e d bea k a n d
clawed w i n g s . I t h a d b i rd l i ke

feet b u t re pt i l i a n vertebra e a n d
ta i l . I t h a d t h e w i s h b o n e of a
bird b u t a repti l i a n b ra i n .
A rchaeopteryx was i ndeed a
m os a i c or j u m b l e of va riously
d eve loped c h a racteristics that
were s u bs e q u e n tly restricted to
d i ffe rent g roups (p. 1 29).

Cynognath us

T H ER I O D O NTS
" beast
toothed ") were reptiles that
l ived i n Perm i a n a n d Triassic
times (p. 98). T h ey s h owed
many m a m m a l ia n c h a racteristics.
Cynognath u s
was a
typ i c a l
m e m be r of t h e g ro u p . An active
carn ivore, six feet l o n g , i t h a d
a l o n g s k u l l w i t h m a m m a l- l i ke
THE

d i fferentiation of the teeth i nto

i n c isors, ca n i nes, a n d cheek
teeth . I t had an " u pri g h t " m a m
m a l i a n postu re, a n d m a ny d e
ta i l s o f t h e s k u l l , vertebra e ,
h i ps, s h o u l d e rs, a n d l i m b s were
a l s o m a m m a l - l i ke . Ma m ma l s a re
b e l i eved to h ave deve loped
from these o r s i m i la r re pti l e s .

P l a ty p u s
FO S S I LS a re s u rviving
representa tives of a n c i e n t fos s i l
g ro u p s . T h e m o n otre m es-the
duckbilled platypus and t h e
sp i ny a n tea ters ( ec h i d n a s ) of
Austra l i a-a re very p ri m i tive
m a m m a l s t h a t reta i n m a n y typ i
cal repti l i a n c h a ra c ters i n t h e i r
LIV I N G

s k e l e to n s . T h e y l a y l e a t h ery,
l a rge-yo l ke d eggs a n d s ecrete
milk
fro m
sweat
m o d i fi e d
g l a n d s . S u c h a n i m a l s pro b a b l y
a rose fro m t ra n s i t i o n a l repti l
i a n - m a m m a l i a n form s . G i n kgos
and a ra ucarias a re p l a n t l ivi n g
fos s i l s .

53

T H E F OS S I L R E C O R D

s hows t h ree other g e n e ra l

featu res w h i c h s u g g est that species a rose b y c o n ti n u
o u s evol utio n . I t d i s p l ays diversificati o n , enviro n m e n t
fi l l i n g , a n d co m pl e x a d a ptatio n a l c h a n g e . Th ese a re
p recisely w h a t wou l d b e pred i cted, a priori, o n t h e
b a s i s o f t h e t h eory of evol uti o n .
THE

ADAPTAT I O N

OF

OR

GA N I S M S to a g reater ra n ge of

envi ron m e n ts
has
d eveloped
with t i m e . T h e o l dest orga n
is m s were confi n e d t o the seas,
but fresh waters, the l a n d , a n d
t h e a i r were successively colo
n ized . Deta i l s a re given on p.
1 1 8 and t h e fol lowi n g pages.

Mammals

Modern Amphibians

AQUATIC

Cartilaginous fishes

54

T h e h istory of t h e verte b rates

s h ows an i n cre a s i n g ra n g e of
ada ptation. Birds and a few
m a m ma l s and extinct re ptiles
h ave colonized the a i r; others
h ave " retu r n e d " to t h e a q uatic
l ife a b a n d o n e d by t h e i r a n ces
tors. Deta i l e d a d a ptati o n s have
deve loped i n each enviro n m e n t .

_j

l_j__l_j

----.

c
0

;;;

c
0

..,

1.

_ _

. .,

"

T h e c l a sses of vertebrates s h ow an i n c re a s e i n co m p l e x ity a o d d i

vers i ty w i t h t i m e . T h e w i d t h of t h e co l u m n s i n d i ca t e s t h e re l a t i v e
a b u nda n c e of e a c h g ro u p ; dotted l i n e s s h ow t h e pro ba b l e or i g i n
o f e a c h g ro u p . ( After S i m p so n )
T H E TOTAL N U M B E R of s p e c i e s
h a s s h own a steady i n c r e a s e
th rou g h g e o l og i c t i m e. I n sects ,
for exa m p l e , c o n s t i t u t e m o re
t h a n t h re e-qu a rters of a l l l i v
i n g s pe c i e s , yet t h ey d i d not
a p p e a r u n t i l the Devon i a n , 375
m i l l i o n yea rs ago ( p . 98). The
c l a s s e s (a bove) s h ow s i n crea s i n g
n u m b ers of s pe c i e s .

G R EATE R C O M P L E X I TY of o r g a
n i s m s w i t h t i m e h a s a c co m pa n i ed
t h e i n va s i o n of n ew e n v i ro n m e n ts .
"Co m p l e x i ty" i s a n a m b i g u o u s
q u a l ity, b u t m o s t wo u l d a g ree
that fish, a m p h i bians, reptiles,
a n d m a m m a l s represe n t s u c h a
s ca l e . Th i s i s a l s o t h e o r d e r of
t h e ir a p pe a ra n c e in the f os s i l
reco rd.

55

_ _

THE PROCESS OF EVOLUTION

I N H ERITAN CE p rovi d e s t h e c o n sta n cy of fo r m ( a l i o n

i s a l i o n i s a l i o n ) a n d va ri a t i o n i n d e ta i l ( b l u e e y e s ,
g re e n e y e s , b ro w n e y e s , b l a c k eyes ) t h a t a re c h a ra c
teristic o f a l l l ivi n g p l a nts a n d a n i m a l s . lobsters p ro
d u ce o n l y l o b sters; h u m a n s p ro d uce h u m a n s . B ut n o
two l o b sters a n d n o two h u m a n s a re eve r i d e n t ica l i n
a l l c h a r a cte rist i c s . H o w a re t h e s e two, co n sta n cy a n d
va riety, t ra n s m itted fro m p a re n ts to offs p ri n g ?
GENES, a s demon strated by
Greg o r M e n d e l (p. 26) and by
s u bs e q u e n t stu d e n ts of g e n et i cs ,
are t h e reg u lators that g overn
t h e deve l o p m e n t of c h a racteris
tics i.n new i n d iv i d u a l s . Genes
are m a d e of deoxyr i b o n u c l e i c
acid-DNA (pp. 72-75)-a n d
reproduce themselves exactly.
G e n e s are i n corporated i n
v i s i b l e structu res c a l led c h ro mosomes, e a c h of w h i c h conta i n s

m a n y g e n e s . Ea c h species h a s
a defin ite type a n d n u m be r of
c h ro m osomes. In all b u t t h e
m o s t s i m p l e orga n i s m s , t h e
c h romosomes a r e conta i n e d i n
t h e ce l l n uc l e u s . C h a ra cteri stics
of a n o rg a n i s m a re governed
by parti c u l a r g e n e s ; b u t i n d i v i d u a l g e n e s m a y i n teract w i t h
o n e a n ot h e r o r co m b i n e to
prod uce c o n t i n uous v a r i a t i o n
o f s o m e c h a racters .

T h e h u m a n c h ro m osomes s h own be low ca rry g e n e s that d eterm i n e

i n d ividual c h a racteristics. X a n d Y a re the s e x c h romosomes.

11

2 3 ,.;,.

of c h romosomes
i n order of s i z e

3<
6

KK 81

XI

II

' -
I
"'"' J'

:J-I
1-.f

7":\
'l

"'

lit

Kl
9

\
l

1"1
}I'
1.

_ .t t.

I1 3I

XX
10

IX
16

XK

\ 19

c h rom os o m e s i n s p e r m a tog o n i u m

J fi K&
4

i ii

a1 7x

1 1

20

Hl

A1 5ft
12

II

14

II

AM
21

18

11 5
22

I
y

II

spermatogon i u m

;:

rr t
p rmatocyte '

1 II

..

(f\
\!
...

THE DEVELOPMENT OF

11 1 1

fi st
oocy t e

. JI" , .
.,.,_
lfC

seco n d
spermatocy te$

.J.

...

' P "m'

II \.

111 1

.!

)(

J)

,J.

a re the s peci a l ized

re productive c e l l s of p l a n t s a n d
a n i m a l s . Most s p e c i e s prod uce
both male g a m etes (sperm) a n d
fe m a l e g a m etes (egg s).
Eac h g a mete c o n ta i n s o n ly
one c h ro m oso m e from each p a i r

" . .

fi rst
' ; polar
body

II.,I' ...........

-\.ve
pola

ootid

! ' / I
II

GAMETES

_...,

second
oocyte

\ sp r m atl ds J

'<,

o ogoniu m

S PERMS A N D EGGS

egg

fertil ized e g g
(zygote)

of the o rg a n i s m 's d i p l o i d c e l l s .
U n l i ke t h e body c e l l s , w h i c h
h ave a d i p loid o r 2 n c h ro m o
so m e n u m ber, t h e tota l n u m ber
of c h ro m os omes i n a g a m ete i s
n . These ce l l s w i t h u n p a i red
c h romosomes are c a l led h a p l o i d .

C H R O M O S O M E S of most species occu r in pa i rs so

that e a c h c e l l c o n ta i n s two s i m i l a r ( ho m ol o g o u s) c h ro
moso me s . Ea c h c h ro m o s o m e i n turn conta i n s h o m o l o
g o u s g e n e s . S u c h cel l s a re ca l l ed d i p l oi d . T h e d i p l o i d
n u m b e r i n d icates t h e tota l n u m b e r o f c h ro m os o m es t h a t
t h e s e c e l l s c on ta i n .
I n m a n , fo r e x a m p l e , m ost c e l l s conta i n 2 3 chromo
som es, h e n ce t h e u s u a l c h rom o so m e n u m be r (n) is 2 3 .
But s i n ce t h e c h ro m os o m e s a re p a i red i n d i p l o i d c e l l s,
the tota l d iploid n u m be r i n each c e l l is 2 n o r 46.

57

C E L L D IVIS ION-the consta nt m a n u fa ct u re of n ew

c e l l s-is essenti a l for t h e l ife a n d g rowth of a l l m u lti

cel l u l a r p l a n ts a n d a n i m a l s . I n j u re d or worn out cells
a re replaced by this p rocess, too . Th e new c e l l s a re
exact replicas of th e orig i n a l cel ls, conta i n i n g t h e sa m e
k i n d a n d n u m b e r of c h ro m oso m e s .
Th e p rocess o f p ro d u c i n g n ew c e l l s by d ivision of
the orig i n a l cel l is c a l l e d m itosis. In m ost c e l l d ivision ,
the p rocess is co m pl eted i n less t han two hours. Mi
tos i s c a n pro d u ce n ew i n d ivid ua l s as wel l as n ew body
cel l s . In m ost org a n is m s that reproduce asexually,
thE> new c e l l is ide ntica l to the p a re n t c el l .

STAGES OF M I T O S I S i n
clude :
Ia)
PRO PHASE,
fi rs t
stage o f m i tosi s , i s m a rked b y
thicken i n g o f c h ro m osomes i n
n ucleus a n d s e p a ration o f ce n
trioles from cen tra l body ( ce n
tros o m e ) a bove n uc l e u s . Fibers
from ce n trioles form a s ter. ( b )
S PI N D LE o f fi bers t h e n g rows
between the two s e pa ra ted a s
ters. N uclear m e m b ra n e d i si n
teg rates , a n d t h e t h icke n e d
c h ro m o s o m e s divide l e n g t h w i s e
i n to t w o c h ro m a ti d s . ( c ) I N
METAPHASE,
the
d u p l i ca ted
chromosomes leave t h e n u
cleus. They arra n g e t h e m s e lves
i n to pa i rs across the e q u a tor of
the e n l a rged s p i n d le. I d l I N
A N A PHASE, o n e o f each o f t h e
c h ro m os o m e s p a i rs , or c h rom a
tid, m i g rates to opposite poles
of t h e s p i n d le . l e l TELO P H A S E ,
fi n a l stage, i s m a rked by loss
of s p i n d le, division of centriole,
and division of p a re n t ce l l i n to
two identica l d a u g h te r ce l l s .

THE

58

MEIOSIS

2N

2N

OR

Division

I
N

2N \ Division
N

2N \
N

MEIOS I S is shown for a cell

with a d i ploid n u mber (2N) of
four c h ro m oso mes. The fl rst d i
vision i n volves t h e ra n d o m s h o r
i n g of exactly h a l f the n u mber
of c h ro mosomes betwee n the
daughte r ce l l s . In this reduc
tion d iv i s i o n , the s h o r i n g of t h e
orig i n a l c h ro m osomes m a y pro
d u c e e i t h e r of two co m b i nations
i n the haploid d a u g hter c e l l s ,
ea c h of w h i c h h a s N c h ro m o
somes-that i s , h a l f of the orig i
n a l n u m ber o f c h ro m o s o m e s .

._..
OR

2N \Division
N

The s e c o n d d iv i s i o n occ u rs by
m i tosis, in w h i c h each n ew
d a u g h ter c e l l d iv i des into two
f u rther h a p l o i d c e l l s , each a l s o
with N c h ro m osomes. These form
the g a m etes, or reprod u ctive
ce l l s , ea c h of w h i c h h a s one
c h romoso m e from each of the
ori g i n a l pa i rs. The u n i o n of two
ga metes, each of which has t h e
haploid o r N c h ro m oso m e n u m
ber, g ives the , offs p r i n g the
orig i n a l 2 N c h romoso m e n u m be r
o f t h e parents.

S EXUA L R E P R O DUCT I O N i n vo lves a seco n d a n d d if

fe re n t k i n d of c e l l d ivision , c a l l e d m e i o s i s . I n sexua l l y
re p rod u c i n g o rg a n i s m s, m eiosis is c o n f i n e d to t h e tes
tis of t h e m a l e a n d to t h e ova ry of t h e fe m a l e . A l l t h e
oth e r c e l l s o f t h e body i n c rease i n n u m b e r by m itos i s .
Meiosis i n volves two d isti n ct sta g es of c e l l d ivisio n .

59

T H E PATTERNS OF I N H E RITANCE we re fi rst d e m o n

strated by Gregor Me n d e l ( p . 2 6 ) with seve n e a s i ly

recog n i z e d c h a ra cters i n g a rd e n p e a s ( s i z e , s h a p e ,
col o r of flowers, a n d so o n ) . I n studyi n g t h e i r occu r
r e n ce t h ro u g h s u ccessive g e n e rati o n s, he d i scovered
t h a t c h a ra cters a re contro l l e d by factors ( n ow c a l l e d
g e n es) t h a t d o n o t b l e n d i n t h e offs p ri n g . H e a l so d i s
cove red t h a t i n t h e seco n d g e n e ration so m e c h a ra c
t e r s , s u c h a s s h o rt ste m s , yel low c o l o r , a n d w ri n k l e d
p e a s , m ay b e m a s ked b y others, s u c h a s l o n g ste m s ,
g re e n c o l o r , a n d rou n d s h a p e . The m a s k e d features
wi l l re a p p e a r i n the t h i rd g e n e ratio n . H e c a l l e d t h ese
c h a racters dominant a n d recessive and c o n c l u d e d
t h a t t h ey a re i n h e rited i n d e p e n d e ntly of o n e a noth e r .
THE I N HER I TED RAT I O o f o n e

c h a racter to a no t h e r w a s s h ow n
by Mendel to be c ons t a n t H e
c ross-po l l i nated p u re b r e d ro u n d
p e a p l a n ts (R) w i t h p u reb red
w ri n kled pea p l a n ts ( r ) . T h e Ft
(fi rs t fi l ia l ) g e n e r a tio n peas were
a l l rou n d , but t h ose in the next
g e n ehtt i on ( F2 ) i n cluded 25 per
cent wri n k l e d , as s h own h e re .

RR

1 . Pure
Domin a n t

60

ents

rr

Go m e t e s
F,

Rr

ar

Offs p r i n g
Rr

1 . Pure
Recessive

Me ndel co n c l u d ed that eac h

p l a n t h a d a p a i r of g e n e s for
each c h a ra cter, b e c a use s o m e
p l a n ts ( s u c h a s t h o s e of t h e Ft)
w i t h a do m i n a n t c h a ra c t e r p ro
du ced so m e offspr i n g w i t h t h e
recessive c h a ra cter. F t pl a n t s
a re s h ow n a s Rr o n t h e d i a
g ra m . P u rebred p l a n ts , a l so
w i t h pa i red g e n e s , a re RR or r r

Me n d e l reaso n e d t h a t t h ese
pai red g e n e s m u st s e p a rate to
form g a m et e s. At f e rt i l i za t i o n ,
t h ey u n i t e ra n do m l y w i t h ot h e r
g a m etes, p rodu c i n g a p redi ct
a b l e ra t i o o f c h a racteri s t i c s i n
t h e offs p r i n g. Ma n y f e a t u res m a y
be i n f l u e n ced b y m o r e t h a n two
k i nds of c o n t ra s t i n g c h a racters,
cal l ed a l l e l e s , in e a c h gene. Con
t i n uo u s l y varyi n g c h a ra c te r i s t i c s ,
l i k e h e i g h t, a r e a c co u n ted f o r
p a r t l y by t h i s a nd p a rt l y b y t h e
f a c t t h a t severa l d i f f e re n t pa i rs
of g e n e s m a y c o n t r o l t h e sa m e
c h a ra c t e r .

T h e gen oty p e of a n y o rg a n i s m i s its tota l c o m p l e m ent

of g e n etic m a te ria l s . T h e p h ysica l c h a ra cte ristics of an
o rg a n i s m a re called its phen otyp e . T h e g e n otype Rr
p rod uces t h e p h e n otype of ro u n d p e a s .
Beca u s e o f t h e d o m i n a n ce of
some c h a ra cters, o rg a n i s m s with
t h e same p h e n otype m ay h ave

TH R E E G E N OTYPES - R R , R r, a n a
rr-a re s h ow n i n p l a nts o f t h e F 1
g e n e ra t i o n a n p . 6 0 . A p l a n t w i t h
a pa i r of t h e sa m e g e n e s for a
part i c u l a r c h a ra c t e r ( R R a n d rr)
i s ca l l ed h omo z y g o us m ea n i n g
"sa m e pa i r." O n e w i t h a pa i r of
d i ffere n t g e n e s i n t h e g e n otype
( R r) i s h eterozygous, o r " d i ffe re n t
pa i r." G e n e s of t h e sa m e g e n e
pa i r - R a n d r, f o r e x a m p l e , a re
a l le l e s .

g e n otyp e s .
d iffere n t
Ro u n d
p e a s , f o r e x a m p l e , m a y h ave
g e n otypes of e i t h e r RR or R.
O n l y by s t u d y i n g t h e i r offs p r i n g
c a n we d i st i n g u i s h
b e twee n
t h e m . H o m ozyg o u s ( R R) p l a nts
wi l l b r e e d tru e , p ro d u c i n g a l l R
offs p ri n g . H e te rozy g o u s p l a n ts '
p ro d u ce b o t h R a n d r offs p ri n g .

BLENDING OF SOME CHARAC

T E RS m a y occ u r w h e n o n e a l l e l e
i s n o t co m p l etely d o m i n a n t over
the o t h er. In s h o r t h o r n c a t t l e , a
red b u l l ( R R ) a n d a w h i t e cow
(ww) p ro d u c e h e t e r o z y g o u s

ca lves t h a t a re roa n ( a m i x t ure

of red a n d w h ite h a i rs ) . C ro s s
i n g a roa n b u l l ( Rw ) a n d a
roa n cow ( R w ) g ives a p r o b
a b l e ratio of 1 w h i te , 2 roa n ,
a n d 1 red .

61

T H E LAWS O F I N H E R I TAN C E a re d e m o n strated below

b y the i nteractio n of two d ifferent c h a ra cters i n stud

ies of a co m b i n ation of pea p l a n t size (T
tall,
do m i n a nt; and t
dwa rf, recessive) a n d p e a s h a pe
( R = rou n d , d o m i n a nt; r = wri n kl ed , recessive) . C ross
i n g a p u re rou n d -seeded, ta l l p l a n t ( RT) with a pure
wri n k l e-s e e d e d , dwa rf p l a n t ( rt) g ives the results s h own .
Men d e l ' s s i m p l e ratios were obta i n e d by avera g i n g t h e
resu lts of c rossi n g l a r g e n u m bers o_f d iffe rent pla nts .
=

R = Ro u n d ( do m i n a n t )
r = Wri n k led ( recessive )
T = To l l ( do m i n a n t )
t = S hort ( recessive )

PARENTS

r R IT

R rT t

MEN D E L I A N RAT I O
9
3
3
1

Round-To l l
Rou n d-short
w ri n k led-Ta l l
w r i n k led-s hort

62

TA I LS

Prob a b i l ities a re of m a jo r i m p o rt a n ce i n u n de r
sta n d i n g g e n etic m ec h a n i s m s . Pro b a b i l ity is t h e li keli
hood of a p a rticu l a r even t h a p p e n i n g . I f you toss a
coi n , t h e re is a 50 percent c h a n ce that it wil l com e
down h e a d s . W e ca n ca l c u l ate t h e p ro b a b i l ity o f t h i s .
P is t h e p rob a b i l ity, f is t h e total n u m ber of ways i n
w h i c h t h e eve n t m ay o c c u r, a n d u i s t h e n u m be r of
ways in which so m e other u n favo ra b l e event m ay occur.
f
P =
-

f+

u .

If we toss a coi n 1 00 ti m es, it i s j u st a s li kely to com e

down h e a d s a s it i s tai l s for a ny g iven th row. S o the
p rob a b i l ity of t h rowi n g a h e a d ( PH) will b e :
P H eo d

f( Heads }

---

f( Heads } + U( rai ls )

50

50 + 5 0

50

1 00

0 5

I n g e n etics, g a m etes a re co m pa r a b l e to t h e coi n s ,

a n d t h e zygotes res u l t i n g fro m t h e u n io n of t h e
g a m etes a re c o m p a ra b l e to t h e h e a d s a n d ta i l s . I n m a n,
with 2 3 p a i rs of c h ro m oso m es, t h e re a re 223 o r
8 , 3 8 8 , 6 0 8 d i ffe rent co m b i n atio n s possi b l e i n t h e p ro
d u ction of s p e r m s a n d e g g s . S i n ce any o n e of these
s p e r m may ferti l i z e a ny of an e q u a l n u m be r of k i n d s
o f e g g s fro m a fe m a le, it i s theoretic a l ly possi b l e for
o n e pa i r of h u m a n s to prod u c e a s m a ny offs p ri n g a s
t h e wo r l d p o p u l a t i o n wit h o u t a n y t w o b e i n g i d e n ti c a l .
63

I N H ERITAN CE PROBAB I LITI E S ca n be ca l c u l a te d j u s t a s

p r o b a b i l ities a r e c a l c u lated i n fl i pp i n g a coi n . With

0.5 means a 50/50 or a
th e coi n , a va l u e of PH
o n e out of two p roba b i l ity of obtai n i n g a head o n
a n y o n e t h row . S i m i l a rly, t h e proba b i l ity ( PT) o f o b
ta i n i n g a t a i l i s a l so 0 . 5 . T h e s u m o f a n y d e p e n d e n t
p ro b a b i l i ties is always 1 . I n the c a s e of t h e coi ns, t h i s
1
ca n b e exp ressed a s P H + PT
Proba b i l ities ra n g e from 0 to 1 . 0 . P = 1 i m p l ies a
pa rtic u l a r eve n t is ce rta i n to ta ke p l a ce . P = 0 m e a n s
a n eve n t is i m possi b l e . T h e p ro b a b i l ity o f two o r m o re
i n d e p e n d e n t events occu r r i n g s i m u lta n eously is g iven
by t h e p ro d uct of their i n d ivi d u a l proba b i l ities, o r
P = P 1 X P2 I n t h i s fo r m u l a , P i s t h e p ro b a b i l ity o f
t h e eve nts occu r r i n g s i m u lta n eously, P 1 is t h e p roba b i l ity
of o n e eve n t, a n d P2 is the proba b i l ity of t h e other.
S o m eti m e s a single eve n t ca n have m o re tha n o n e
cause. B l a c k color , i n g u i nea p i g s b e l ow for exa m ple,
may be p rod uced by a l l black o r by b l a c k and wh ite
a l l e l es, if b l a c k is d o m i n a nt. T hen the p ro b a b i l ity of suc h
a n eve n t ( 2 ) t h a t m a y a r ise i n m o re t h a n o n e wa y is
the s u m of the p ro b a b i l ities fo r each of its ca u ses (p
a n d q ) . T h i s can be c a l c u l a ted by P2 = P P + PQ .
=

P u rebred parents

P u re bred parents

88 ( p u rebred b l a c k )

8b ( m i x e d b l a c k )

64

M i x e d parents

8 8 , Bb

or

bb ( p urebred w h i te )

C h e c k e rboard C a l c u l a t i o n of Possible G e n otypes

C H E CKE R B O A R D s h ows m et h o d
f o r ca l c u l a t i n g t h e g e n otypes i n
offs p r i n g of t w o g u i n e a p i g s
t h a t a re h e t e rozyg o u s f o r b l a c k ,
s h o r t h a i r. ( B = b l a c k , b =
b row n , S = s hort h a i r, a n d s =
l o n g h a i r. ) T h e f o u r poss i b l e
g a m etes f r o m e a c h a re a r
ra n g e d a lo n g t h e s i d e s of t h e
c h ec k e r b oa rd . T h e i r co m b i n a
t i o n s a re p l otted i n t h e s q u a re s .
S i n ce B and S are d o m i n a nt, a

b l a c k s h ort p h e notype c a n be
p ro d u c e d by BBSS, B b S s , B b S S ,
and BBSs.
I n t h e c o m b i n a t i o n , t h e re w i l l
b e 9 b l a c k s h orts ( 1 BBSS, 2
B b S s , 4 B b S s , a n d 2 BBSs), 3
b rown s ho rts ( 1 b b SS a n d 2
b bS s ) , 3 b l a c k l o n g ( 1 B B s s a n d
2 B b s s) , a n d 1 b ro w n l o n g ( b b s s ) .
T h i s rat io of 9 : 3 : 3 : 1 correspo n d s
to t h a t of t h e H a rdy-We i n b e rg
P r i n c i p l e, b e lo w .

o r fe rtilized e g g s,
a re f o r m e d by two i n d iv i d u a l
g a m et e s , e a c h o f w h i c h c a n b e
reg a r d e d a s a n i n d e p e n d e n t
" eve n t . " I f p i s t h e p ro b a b i l i ty
of a zygote w i t h a b l a c k
c h ro m os o m e a n d q i s t h e p rob
a b i l ity o f a zygote with a w h i te

t h e proba b i l i ty o f t h i s by ra n
dom mating i n a large popula
t i o n f r o m t h e for m u l a t h a t g ives
t h e p rod uct o f the male c o n t r i

ZYGOTES,

chro m oso m e , p + q = 1 .
A zyg ote w i t h o n e b l a c k a n d
o n e w h ite c h ro m os o m e m a y be
p ro d u ced in two w ay s-e i t h e r
pq o r q p , . e a c h g iv i n g t h e i d e n
t i c a l resu lt. W e c a n ca l c u l ate

bution ( p + q ) a n d the female

co n t ri b ut i o n
( p + q ) X ( p + q) =
( p + q) 2 = p 2 + 2 pq + q 2 = 1
This

re l a t i o n s h i p ,

called

the

H a r dy-We i n berg Pri nciple, shows

t h a t in a l a rge, s t a b l e p o p u l a
tio n , freq u e n c i e s of d i ffere n t
g e n otypes a re predicta b l e a n d
co n s e rva tive fro m o n e g e n era
tio n to a n other.

65

T H E M E C H A N I S M O F I N H E R I TA N C E revea l e d by the

stu d ies of M e n d e l and l a t e r g e n eticists s h owed t h a t

e a c h i n h erited c h a ra cte r, s u c h a s s e e d s h a p e o r col o r
i n peas, is contro l l e d by a pa i r of g e n e s . O n e m e m b e r
o f e a c h p a i r ( a n a l l e l e ) i s contrib uted b y e a c h p a re n t .
The g e n e s a re l ()cated o n vis i b l e rod l i k e c h ro m os o m e s ,
w h o s e b e havior s h ows a c l o s e pa ra l l e l t o th a t p re
d icted fo r g e n es (p. 6 0 ) . We h a ve a l rea d y s e e n how
these structu res d ivid e i n e g g and sperm fo r m a ti o n a n d
com b i n e i n fe rtil izati o n . B u t w h a t d ete r m i n es t h e sex
of the n ew i n d ivid u a l ?
SEX of the offs p r i n g is
deter m i n e d by s p e c i a l ized pa i rs
of sex c h romosomes. The fe m a l e
parent h a s one p a i r o f s i m i l a r
chromosomes (X c h ro m osomes,
p. 56), and t h e m a l e parent has
one pa i r of d i ssi m i la r c h ro mo
somes (X a n d Y). T h e d i v i s i o n of
the c h ro moso mes w h e n e g g s
a n d s p e r m s a re produced g ives
the eggs all X c h ro m osomes a n d

THE

t h e sperm h a l f X a n d h a l f Y
c h ro m o s o m e s . T h e ra n do m p a i r
i n g i n ferti l i z a t i o n res u l ts i n
either X X (fe m a le) o r X Y ( m a le).
O t h e r g e n e s , i n c l u d i n g those
t h a t produce h e m o p h i l i a a n d
some k i n ds o f color b l i n d n e s s ,
a re l i n k ed to sex c h ro m osomes.
These sex- l i n ked c h a racters c a n
be tra n s m i tted to t h e offs p r i n g
of a p parently " n o r m a l " parents.

S e x of a n i m a l s i s d e term i n ed by t h e sperm .
c h r o m o s o m e t h e offspri n g w i l l be m a l e .
,

66

I f i t co n ta i n s a Y

G ENES c o n t ro l i n h e rita nce, b u t w h a t c o n t ro l s g e n es?

How do they t ra n s m it their c o m plex g e n et i c i nfo r m ation

from one g e n e ra t i o n to a n o t h e r ?
C h e m ica l a n a lys is of c h ro m o so m e s h a s s h ow n t h a t
they co n s i st of fou r b a s i c co m pou n d s : two p rote i n s ,
one with a re l a tively low m o l ec u l a r we i g h t a n d t h e
other m u c h h i g h e r, p l u s two n uc l e i c a c i d s, d eoxyribo
n ucleic a c i d ( D N A ) and r i b o n u c l e i c a c i d ( R NA ) . The
co m b i n a t i on of t h e s e fo u r m o l e c u l es fo r m s chromatin ,
the " s tuff" of w h i c h c h ro m osom es a re m a d e . DNA
ca rries and c o n t ro l s g e n etic i n fo r m atio n .
DNA was f i rst s u s p ected t o be
t h e " b a s i c co m po n e n t" of g e n e s
w h e n i t w a s d i scovered t h a t t h e
n u c l e i of b o t h re p ro d u c t i v e a n d
body c e l l s of a n y p a rt i c u l a r s p e
c i e s c o n ta i n e d i d e n t i c a l a m o u n ts
of D N A for ea c h c h ro m o so m e
set . O n e stra i n o f t h e bacte
r i u m P n e u m ococc us cou l d be
tra n sfo r m e d i n to a n o t h e r by i n
fec t i n g i t w i t h a p u r i f i e d e x t ra c t

from d e a d cel l s of t h e s e c o n d
stra i n . T h e " t ra n sf o r m e d " stra i n
b r e d t r u e , a n d t h e ex t r a c t was
l a t e r i d e n t i f i e d a s DNA. T h i s
n o n l i v i n g D NA f r o m a s e c o n d
stra i n t h e re f o r e h a d t h e a b i l ity
to t r a n s f o r m the h e re d ita ry m e c h
a n i s m of b a cte r i a c e l l s . S i m i l a r
tra n sfor m a t i o n s i n d u c e d i n ot h e r
bacte r i a s h o w t h a t g e n e s a r e
co m posed o f D N A.

T h e ge netic tra nsfo rmation of Pneumococcus Typ e I I cel l s i nto Type

I l l c e l ls by the addition of an extract from d e a d Type I l l cel ls.
EXPER I MENTAL C U LT U R E

nutrie nt, p l u s
Type I I cells

sterile nutrient

type I l l
cells a d d e d

type I I c e l l s
tra n sfo r m to t y p e I l l

t y p e I l l cells a d d e d

no colonies

CONTROL C U LT U R E

67

( de
o xyri bonucleic a c i d ) , w h i c h controls hered ity b y reg u
l a ti n g i n structi o n s of g rowth and form from cell to cell
a n d fro m parent to offs p ri n g . DNA i s present in all l iv
i n g creatu res.
In structure, the DNA molecule is a double helix, re
sem b l i n g a l a d d e r that has been repeatedly twiste d .
Ea c h r u n g of t h i s m olecu l a r l a d d e r is m a d e u p o f a
pai r o r two fro m fou r c h e m i c a l bases-a d e n i n e , thy
m i n e, g ua n i n e, a n d cystosi n e . The size a n d structure of
these bases is such that each " ru n g p a i r " always con
sists o f either a d e n i n e a n d thym i n e o r o f g u a n i n e a n d
cystosi n e . It i s the seq u e n ce o f t h e pairs i n the ru n g s
that provi d es t h e code b y w h i c h g rowth i n stru ctio n s a re
tra n s m itted . From these fou r basic code substa n ces, a n
a l m ost i n fi n ite va riety of seq uences can d evelop.

T H E CH EMICAL KEY TO INHERITAN CE IS D NA

T h e portion of a DNA molecular m o d e l h e re s h ows t h a t each ru n g

of t h e h e l i x ladder i s b u i l t of a pa i r of c h e m ical bases.

adenine

cystosine

guanine

Model of

DNA Molecu le

thymine

T h e s p l itt i n g of a DNA ladder a n d t h e reco m b i n ation of t h e un

z i pped r u n g s with a p p ropriately fitti n g n ew base u n its, i d e ntical
to t h e i r origi n a l partners o n the run g , p rod uces two n ew m ol e
c u les. H a l f of each new m olecule is derived from t h e orig i n a l
pare n t molecule.
"

"

DNA con t ro ls g rowth by h a v i n g the u n i q u e a bi l it y

to u n z i p d own the m i d d l e of t h e m o l e c u l a r l a d d e r,
thus freei n g the p a i red e n d s of each c h e m i c a l run g .
The exposed com pou n d s then l i n k u p with si m i l a r n ew
u n its that a re d e rived from the o r g a n is m ' s food sup
ply. As t h e s p l it of t h e m o l ec u l a r l a d d e r conti n u es,
each half b u i l d s an exact rep l i ca of itself by this l i n k
a g e . The rig i d seq u e n ce of pairi n g i s preserved by t h e
fit o f the o ri g i n a l structure. I n t hi s way the DNA mole
cu le c a n repro d u c e itself over and over a g ai n .
69

PROT E I N MAN UFACTURE . RNA ( ri b o n u c l eic a ci d ) h a s

a structu re m u c h l i ke that of D NA . U n l i ke D NA, h ow

ever, it is not c o n fi n e d to the c e l l n uc l e u s . It is t h e es
se ntial m esse n g e r in the m a n ufacture of p rote i n s by
cells. Prote i n s a re i nvolved i n every body p rocess . They
fo rm the basic m ate rial of l iv i n g o rg a n i s m s . They a re
very l a rg e m o l e c u l es, m a d e u p of various co m b i n atio n s
of 20 essential k i n d s o f a m i no a c i d s .
R N A i s m a n ufactured fro m D NA, fro m w h i c h i t dif
fers i n h avi n g o n e m o re oxyg en atom i n its sug a r
( ri bose) a n d i n havi n g u racil i n stead o f thy m i n e i n t h e
m o l e c u l a r stru ctu re. S l i g htly d ifferen t " m esse n g e r " R N A
m o l e c u l e s c o n t r o l t h e m a n ufa ctu re of e a c h of the m a ny
k i n d s of protei n . Each is i m pri nted with a c o d e d tem
pl ate o f its structure. Selected a m i n o a c i d s a re t h e n
tagged by sti l l o t h e r d i stin ctive " tr a n sfe r " RNA m o l e
c u l e s , e a c h i m pri n te d w i t h a structure t h a t e n a b l es i t
t o b r i n g t h e p a rti c u l a r a m i n o ac id t o t h e m esse n g e r
te m p l ate . W h e n these a m i n o a c i d s a re a rra n g ed a n d
c o m b i n e d i n correct seq u e n ce, R N A m a y s e p a rate fro m
newly p rod uced p rote i n a n d perfo rm its ta s k a g a i n .
T h e m e s se n g e r RNA bar carries t h e cod e , w h i l e each of t h ree
s m a l l e r co m po n e n t s of t h e RNA u n i t is s t r u c t u red to a c c e p t a pa r
t i c u l a r a m i n o a c i d s h a p e . I n t h e bac k g ro u n d is a DNA h e l i x .

Tra n s f e r R N A u n its locate ( Le f t ) a n d tra n s port t h e i r ( Ri g h t ) vari

ous a m i n o acids toward t h e coded RNA te m p l a t e bar.

Tra nsfer RNA u n its u n ite with a

messe n g e r u n it op posite t h e i r
pa i red p a r t n e r c o m p o u n d s , s o
arra n ged as to b u i l d a n a m i n o
a c i d s e q u e n c e i n to a part i c u l a r
p rote i n m o l e c u l e .

T h e new prote i n m o l e c u l e
cons tructed f r o m a seq u e n ce o f
a m i n o a c i d s , separates f r o m t h e
R N A . M o s t p rote i n s consist o f
h u n d reds of a m i n o a c i d u n its
that are a rra n g e d tog ether.

71

WHAT ARE THE S O U RCES OF VARIABILITY? If g e netic

e q u i l i b ri u m i n any p o p u l ation is so conservative, h ow

does c h a n ge eve r take p l a c e ? How c o u l d evo l utio n i n
p l a n ts a n d a n i m a l s occur? W hat a re the so u rces of
variati o n s in p l a n ts a n d a n i m a l s ?
No two i n d ivid u a l s a re eve r exactly a l i k e . T h e s e
i n d ivid u a l d i fferen ces a re a s recog n i z a b l e i n d o g s o r
i n g o l d fi s h a s t h ey a re i n m a n . S o m e d i ffere n ces a re
acq u i re d d u r i n g life th roug h b e h avio r o r fro m t h e e n
vi ro n m e n t . D i eti n g , o r l a c k of i t , i nfl u e n ces wei g ht . R e s
p i rato ry d iseases m ay be pro d u ced by s m o ki n g o r b y
l ivi n g i n a p o l l uted a t m o s p h e r e . But m a ny i n d ivi d u a l
c h a racteristics a re h ered ita ry, n o t enviro n m en ta l . These
i n c l u d e s u c h phys i c a l features a s eye c o l o r, co m p l ex
i o n , color b l i n d n ess, a n d h e i g h t . How do t h ese d if
feren ces a rise?
Variations of this kind a re pro d u ced in two ways :
by a s h uffl i n g a n d red i stri buti o n of g e n e s d u ri n g re
( reco m b i n atio n } a n d
by sponta n eo u s
pro d u ction
c h a n ges i n g e n e structure l e a d i n g t o n ew c h a racteris
tics ( m utatio n } .
Crowd sce n e i l l u s trates vari a b i l i t y of o u r ow n speci es.

RECOMB I NAT I O N i s the ch ief so u rce of v a r i a tion . I t

m ay occu r e i t h e r b y a s i m p le reco m b i n at i o n o f the

c h romosomes w h e n ferti l i zati o n between p a rents ta kes
place o r by a c ross i n g -over that i n volves an exc h a n g e
o f g e n etic m ateri a l between c h ro m osomes . Reco m bi n a
tion i n volves t h e fo r m a t i o n o f n e w g e n otypes by the
res h u ffl i n g of e x i s t i n g o n es ( p. 6 4 ) . I n o rg a n i s m s t h a t
reprod uce a s e x u a l ly, m uc h less pote n t i a l va riati o n is
poss i b l e t h a n in those that reprod uce sexu a l ly. Th is
has put an evo l u ti o n a ry " p re m i u m " on sex u a l rep ro
d uctio n . Sex u a l re p ro d uct i o n i s the m a j o r so u rce of
the wide va r i a t i o n in p l a n t a n d a n i m a l s pecies .
of chromo
somes s o m e t i m e s takes p l a c e
duri n g m e iosis, t h e formation
of sex c e l l s . This results from
c h ro moso m e stra n d s stic k i n g to
g e t h e r so that an e x c h a n g e of
genes ta kes p l a ce
betwee n
c h ro m oso m e pairs. Where these
c h ro m osomes a re joined and the
CRO S S I N G-OVER

exchanges

occur

a re

called

chiasmata. Ea c h cross-ove r dou

b l e s t h e n u m ber of k i n d s of
g a m etes and modifies the l i n k
a g e of g e n es , t h u s prov i d i n g a n
i m porta n t sou rce o f varia b i l i ty.
I n the i l l u stration of cross i n g
over below, d iffere n t colors rep
resent d ifferent g e n e s .

CROSSING-OVER

73

Horned Hereford

Polled H e reford

Po l led H ereford cattle l a c k horns beca u s e of t h e effects of m u ta t i o n

of a g e n e w h i c h reg u l a tes horn s ecretio n .

MUTATIONS a re c h a n g e s i n g e n e t i c materi a l that

pro d u ce n ew c h a ra cte risti cs. The c h a n g e m ay b e o bvi

ous (as exa m pl es, wi n g for m in fl i es, horn l ess pol l e d
He refo rd cattle, s h ort- l eg g e d A n co n s h eep, o r double
flowers), o r it m ay be a n i n co n spicuous and subtle
c h e m i c a l a n d p hysiological d iffe re n ce . Particu l a r g e n es
m ay m utate i n m o re t han o n e way, a n d m a ny can re
ve rse the d i rection of c h a n g e .. Each g e n e has a d is
ti n ctive m utation rate . Some wi l l m utate o n ly once per
m i l l ion g a m etes, for exa m ple, while others u n dergo
m utati o n s m o re than 500 ti m es as freq uently. The rate
of m utation ca n be i nfl u e n ce d by va rious factors, such
as te m p erature c h a n g es, ra d iation, a n d c h e m ical sti m
u l a n ts . C o n s p i c uous m utation s a re m o re ofte n h a rmfu l
to t h e org a n i s m t h a n a re t h e s m a l l e r o n es .
Mutation see m s t o be caused b y s l i g ht i m perfection s
i n the self-copyi n g c h e m ical structu res o f the D NA
molecules that m a ke u p the g e n e s . They m ay affect the
n u m ber and structures of the DNA molecules of the
genes. They m a y a lso affect the n u m be r a n d structure
of c h ro m osomes. Occasion a l l y the c h ro m oso m es d ivide
but the cell does n ot, resu lti n g i n a whole new set of
c h ro m osomes ( polyploidy). The n ew m uta n ts b reed true
and do not revert to the orig i n a l typ e .
74

DRIFT was discove red by t h e Ame rica n

gen eticist Sewa l l Wrig ht, who studied the math e m atics
of population g e n etics. The g e n e pool-that is, the
tota l gene contrib ution of a particul a r popul atio,n to its
offspri n g-is g reatly i n fl u e n ced by the size of t h e pop
u l atio n . In very s m a l l populations, such a s those iso
lated from their parent g ro u p, cha nce pl ays relatively
a m uch g reater rol e i n p rod ucing g e n etic c h a nge,
som eti m e s lea d i n g to n o n - a d a ptive c h a n g e s . Although
the real ity of g e n etic d rift has been confirmed i n lab
oratory experi m en ts, its role i n evol ution i s sti l l not
clear. It m ay b e of some i m porta nce i n small popula
tions that l ater i n c rease i n size and m ay acco u n t for
some of the puzzl ing, pers iste nt n o n - a d a ptive o r neu
tral c h a n g es observed i n d iversified wild populatio n s .
Va ri a b i l ity t h u s a rises fro m fo u r possi b l e s o u rces
reco m b i n ation, crossi n g -over, m utation, a n d g e n eti c
d rift. Of these, reco m b i n ation is by fa r t h e most im
porta n t . Once it was thought that variabil ity a l o n e
cou l d p ro d u ce evoluti o n a ry c h a n g e, without a i d from
a d d itio n a l factors . Statistical a n a lysis showed , how
ever, that o n l y natural sel ection cou l d a ccount for t h e
perpetuation a n d refi n e m ent o f t h e e n d less a d a pta
tions shown by livi n g t h i n g s . Varia b i lity is not the
whole of t h e evol utionary reci pe, but i t is t h e essenti a l
raw m aterial o n which everything else d epen d s .
GENETI C

75

I S O LATION of g e n e pool s d ist i n g u i s h e s s pecies from

races a n d d e m es, w h i c h a re p o p u l ations ca p a b l e of

i n terb re e d i n g when t h ey co m e i n to conta ct . The de
ve l o p m e n t of isolation between o n ce i n te rb reed i n g
g ro u p s i s a n i m po rta n t fa cto r i n evo l utio n . O n c e i so
l ated, e a c h p o p u l ation wi l l u n d ergo i n d e p e n d e n t,
g r a d u a l g e n etic c h a n g e u n t i l it i s n o l o n g e r co m p ati b l e
w i t h t h e g ro u p with w h i c h it o n ce i n terbre d . S i n ce c l i
m a tic a n d e c o l o g i c c o n d itio ns u n d e rg o s l ow c h a n g e s ,
th e re is a d yn a m ic i n tera.ction betwe e n th e m a n d s p e
c i e s d istri b u tio n . Ma ny now- d ivi d e d ra n g e s of s p e c ies
reflect i so l atio n of orig i n a l ly w i d e s p re a d p o p u l a t io n s .
I s o l a t i o n m a y a r i s e i n d i ffe r e n t w a y s . I t m a y b e g e o

1 9) .

g r a p h ic, a s i n s o m e i s l a n d p o p u l a ti o n s (p.

R e l a ted

species t h a t d o not ove r l a p i n territory a re c a l l e d a l

lopatric;

those that do,

sy m p a t r i c .

G e n et i c

isolation

m a y a r i s e b etwe e n e i t h e r k i n d of p o p u l a t i o n . I t m a y
i n vo l v e e c o l o g i c a l , b e h a v i o r a l , m o r p h o l o g i c a l , o r p h y s
i o l o g i c a l d i ffe r e n c e s , a n y of w h i c h m a y p r eve n t m a t i n g .
Eve n i f m a t i n g d o e s t a k e p l a c e , v a r i o u s i n t e r n a l c e l l u
l a r o r d ev e l o p m e n ta l b a r r i e r s m a y p r e ve n t f e r t i l i z a t i o n
o r p r o d u ce n o n vi a b l e , w e a k o r s t e r i l e h y b r i d s .
is
I S OLAT I O N
GEOGRAPH I C
s h ow n b e l ow i n s c h e m a t i c for m :
(A) w i d e s p r e a d s pe c i e s w i t h n o
g eog ra p h i c v a r i a t i o n , (B) d i f fer
ent popu l a t i o n s d ev e l o p in ex
t re m e s o f ra n g e s, ( C ) p a rt i a l
barrier
develops,
g eog ra p h i c

(D) b a r r i e r bec o m e s a tota l pre

ve n tative to i n t e r bree d i n g of
two p o p u l a t ion s ( E) cu m u lative
g e n et i c d i ffere n c e s b e co m e so
g reat that g e n et i c s e pa r a t i o n
prod uces two species, eve n i f
t h e b a r r i e r i s e l i m i n a ted.
,

M I GRAT I ON of po p u l a tions i n to new a re a s may l ea d

to perm a n e n t co l o n i zatio n . I n so m e cases, d i fferent

a reas may h ave very closely s i m i l a r c l i m a tes a n d ter
ra i n s b u t b e s e p a rated by such b a rri ers a s oceans,
deserts, o r m o u n ta i n s . Over long periods, a few i n
d ivid u a l s m a y c ross these ba rriers a n d beco m e esta b
l ished i n th e i r new e nv i ro n m e nt. The n ew c o l o n y will
i nterbreed in isolation fro m t h e parent population a n d
m a y i n t i m e d evelop i nto a n e w s p e ci e s.
S o m e of t h e g reatest tra n sfo rmations i n the h istory
of l ife h ave a ri s e n by m i g ratio n . These i n c l u d e t he col
o n i zation of t h e l a n d by p l a nts and a n i m a l s .
M I GRA
I N TERC O N T I N E NTAL
T I O N of some Plei s tocen e m a m

m a l s is s h ow n h ere. Asia a n d
North A m erica were periodi
c a l l y joined b y t h e Beri n g

Straits. T h e " fi l ter" effect of

these i s l a n d co n n ecti o n s co n
fi n ed s o m e m a m m a l s ( ci rcled )
to t hei r ori g i n a l co n ti n e n ts .
( After Kay a n d Co l bert. )

N AT U R A L S E L E C T I O N is the seco n d m a j or co m po n e n t

o f t h e evoluti o n a ry p rocess. Left t o t h e m selves, l a rg e

a n d ra n d o m l y i n t e r b reed i n g p o p u l atio n s w i l l m ove to
wa rd g e n etic e q u i l i b r i u m (p. 6 5 ) . N a t u ra l s e l ectio n is
t h e n e t re s u l t of a l t t h e p hysica l a n d b io l o g i c a l e n v i ro n
m e n ta l factors t h a t t e n d t o d i sturb t h i s e q u i l i b r i u m a n d
c h a n g e a p o p u l a tio n ' s g e n e poo l . N a t u r a l s e l ecti o n d o e s
not i t s e l f c re a te n ew va riatio n s ; it sel ects , w i n n ows, a n d
p r e s e rves e x i s ti n g va riati o n s . The g e n et ic co m po n e nts of
a p o p u l ation dete r m i n e what i t m a y b e co m e ; s e l e c t i o n
d ete r m i n e s w h a t it w i l l becom e . I n he rita n c e p rovi d e s i t
with its potent i a l ; sel ection tra n sfo r m s pote ntia l to
a ctu a l ity.
THE ACTI O N of n atural se l e c

tion depends on t h e tende ncy

of a l l species to p rod uce more
offs pri n g than can norma l ly sur
vive (p. 24) in a n environ ment
of
l i m ited
capacity.
Better
ada pted org a n i s m s w i l l l ive
longer tha n the l ess wel l ada pt
ed, hence prod uce a g reater
n u m ber of descenda nts. The
descendant popu lation will i n
c l u d e a n i n c reas i n g ly large pro
portion of i n d ividua l s that have
features
favora b l e
inherited
from their parents.
Natura l selection is rarely a
"stru g g l e for existe n c e " be-

tween two c o m pet i n g i n divid

uals. Predation and direct com
petition
are
on l y
two
of

many factors i nvolved, includ

i n g such other t h i n g s as mo
bil ity, physiological and struc
tura l efficiency, resista nce to
d isease, a n d sexual vigor.
Natura l selection produces
d ifferential rep roductio n . It is
the "su rviva l of the fittest" o n ly
in th e sense that the fittest
produce more offspring. In ef
fect, it not o n ly e l i m i nates the
less we l l a d a pted characteris
tics but also p rod uces positive
res u lts.

Stro n g b u l l-m a n y offs pri n g

W e a k b u l l-no offspri n g

SELECTIO N

PRESSURE
varies
from season to season and from
place to p lace. It is influenced
by such fl uctuating factors as
cli mate, population size, food
supply, a n d m i g ration. Selection
genera l ly does not lead to long
term stability because so many
of these and other factors un
dergo more or less continuous
change a n d are re i n forced by
such major changes as colon iza
tion of a new environ ment, iso
IQtion, or change in the existi ng
geogra phy of a n a rea.
Natural selection does not
operate as a stea m-roUer effect
on one c h a racter, but as a
subtle series of i n teracti ng
" com promises" t h a t affect t h e

whole organism a n d may influ

ence many differe n t c h a racteris
tics. Selection is a c u m u lative
process. Even over a s h ort pe
riod, its effects can be s ubsta n
tial ( p. 8 4 ) . Over l o n g periods
its effects can be enormous.
The increasing ada ptation of
a population to its environ ment
is the res u l t of natural selection.
Each n iche is genera l ly fi l led
by on ly a single species, but
a n u m ber of s pecies may share
the sa me area by their va rious
adaptations to speciflc food
suppl ies or to habitats. The bet
ter adapted leave more off
spring, g radually cha nging the
character of the desce ndant
popu lation.
Con trasti n g effects of m utation
( B ) a n d n a tura l selection ( C)
are s h own on a " norm a l " dis
tribution curve ( A ) of a char
acter in a large populatio n .
( After Hardi n . )

Carn ivore

79

PRO OF OF NATURAL SELECTION ca me long after

D a rwin presented t h e theory i n The Origin of Spe

cies. D a rwin c o m p a red it with a rtificial d o m estic
selectio n and d e m o n strated its p roba b l e causes a n d
a p p a re n t effects, but the p rocess itself h a d not b e e n
d e m o n strated . M a n y l a b o ratory stud ies h ave s i n c e
b e e n c o n d ucted t h a t i l l u strate both the operatio n a n d
the effects o f natural selection . I n these, the biolog ist
atte m pts to d u p l icate and to isolate ce rta i n natural e n
viro n m e nts a nd t o a n a lyze their i nte racti n g p rocesse s .
M I C E a n c:l o the r wi l d a n i
m a l s , color c h a n g es a re ofte n
closely associated w i t h t h e pre
d o m i n a n t color of the b a c k
g rou n d so i l a n d vegetat i on
w h e re t h e a n i m a l s l ive. I n
Florida, for e x a m p l e , t h e deer
mouse (Peromyscus m a n iculatusJ
s h ows a co n t i n uous color tra n si
tion from very light o n t h e cora l
s a n d of isla n d reefs to dark i n
i n l a n d a re a s of d a rker s o i l .
I s t h i s t h e res u l t of n a tura l
s e lectio n ?
Research ers worked w i t h two
color va rieties-g ray a n d b uff.
Two c on t ra s t i n g b a c k g rou n d s
IN

were set u p i n a large d a r k

e n e d cage. T h e s o i l i n t h e cages
m o re o r l ess matched t h e color
of e a c h of the s h a d e s of m ic e .
E i g h t m i ce, four of e a c h color,
were e x posed to a barn owl for
1 5 m i n u tes. T h e b a c k g rou n d s
were a l ternated with ea c h e x
experi
p e r i m e nt. After 8 8
ments," it w a s fou n d that t h e
o w l h a d t aken 1 07 of t h e " n o n
m a tc h i n g " m ic e a n d on ly 62 o f
t h e bette r c o n cea led ones. T h i s
s h ows how powerful a se lective
factor p rotective
backg rou n d
coloration ca n b e , eve n over a
s hort period.

6 2 M a tc h i n g

1 07 N o n - m a tc h i n g

1 1 20

1 100
I
I 80
I
I 60
I
I 40
I
I 20
I

V i a b i l ity of seven stra i n s (A-G) of h o m ozygous Drosoph ila fruit

fl ies with d e leterious g e nes, before and a fte r t h e e x peri m e n t
described below. Th e via b i l ity is shown a s % o f n o r m a l viabil ity
( 1 00 % ). B l a c k col u m n s i n d i cate t h e v i a b i l ity before t h e expe r i m e n t;
the red col u m n s , v i a b i l ity of u n treated stocks after 50 g e n e ra
tions; t h e yellow col u m n s , of stocks i r ra d iated with X- rays after
50 g e nerati o n s . (After Dobz h a n s ky and Spass ky.)
POORLY

ADAPTE D

A N IMALS

are rarely fou n d in nature, b u t

t h e effect of natura l selection
upon
such
poorly a d a pted
form s ca n be observed u n d e r
experi m e n t a l c o n d i t i o n s . Dobz
h a n s ky and S p a s s ky ra i s e d sa m
p l e s o f t h e fruit fl y D ro s ophila
that
were
h o m ozyg o u s
for
d i ffere n t
c h ro m oso m e
seven
co m b i nations (A-G). Ea c h was
as soc iated with some a b nor
m a l ity s u c h a s reta rded deve l
op me n t or defor m e d w i n g s ,
l e g s , a n d a b d o m e n s-t h a t re
s u lted in a low via b i l ity.
Ea c h of t h ese stocks was
ra ised for fi fty g e n e ra t i o n s a n d
kept i n c rowd ed c u l t u re bott l e s
t o i n crease s e l ection pressure.

The populations were s a m p l e d

a t reg u l a r i n te rv a l s to s t u d y t h e
c h ro m os o m e s of repre s e n tative
i n d iv i d u a l s . In most cases, th ere
wa s a ra pid a n d m a rked de
c l i n e in the a b u n da n ce of t h e
deleterious co m b i n a t i o n s . The
experi m e n t i n c l u d e d e x pos u re
of o n e s a m p l e of each sto c k to
X - ra y ra d i a ti o n , b u t th ese pro
duced n o s ig n ifi c a n t d iffere n ces.
O f t h e 14 stocks, 1 1 s h owed
m a rked i n crea s e in v i a b i l i ty, 2
decl i n e d i n v i a b i l ity, a n d 1 re
m a i n e d u n c h a n g e d . Of the con
trol g roups, u n m i xed w i t h the
stro n g e r n a t u ra l pop u l a t i o n s , 8
s h owed a m a r k e d deterioratio n ,
6 we re u n ch a n g e d or s h owe d
only s l i g h t i m p rove m e nts.

81

NATURAL POPU LATI ONS a re a l so stu d ie d to dete r

m i n e the effects of n atura l s e l ectio n . One exa m p l e is

how n atura l s e l e ct i on m i g ht h ave p ro d u ce d the .d iffer
e n ces that exist b etween the va rious k i n d s of D a rwi n ' s
fr n c h e s o n t h e Ga l a p a g o s I s l a n d s ( p . 1 8 ) . Ty p i c a l
fr n c h es , s u c h a s s p a rro w s a n d c a rd i n a l s , h a ve s t ro n g
c o n ica l be a k s t h a t a re u s e d t o c r u s h se e d s . B ut t h e G a
l a p a g os fi n c h es s h o w a g reat va riety of b e a k s . S o m e
a re sto u t a n d co n i c a l , o t h e rs s l e n d e r, a n d sti l l ot h e rs
a d a pted fo r q u ite d iffe re n t d iets .
T h i s d ivers i ty wa s probably
p rod uced by the presence of
m a ny vaca n t ecolog ical n i c h e s
w h e n t h e fi n c h e s fi rst beca m e
esta b l is h e d on t h e Gala pagos.

I n c reas i n g n u mbers of b i rds

feed i n g o n a l i m ited seed s u p
p l y provi d e d t h e sel ective pres
s u re that favored those that
coul d e x i st o n new food .

DARW I N 'S FI NCH ES ( GA L A PAGOS I S LANDS )

82

The g rou n d - l ov i n g fi n ches re

ta ined t h e i r seed-eating h a bits,
a lthough
i n d iv i d u a l
s pecies
show m a rked variation i n beak
proportions t h a t see m related
to parti c u l a r types of seed
prefere nces .
Other
fi n ches
turned to k i n ds of food a n d
hab its t h a t a re c h a racteristic o f
species o f oth e r fa m i l ies o n
t h e conti n e n ts . S o m e beca m e
tree-dwe l l ers, i n c l u d i n g types
that a re w a r b le r l i k e i n se ctivores
and oth ers t h a t a re pa rrot l i ke
veg eta r i a n s . Sti l l oth e rs beca me
special ized
for feed i n g
on
cacti. An insectivore developed
a strong bea k t h a t it. uses to
bore i nto bark. Lacking t h e long
ton g u e of true wood peckers ( p .
42), it uses a c a c t u s s p i n e t o
extract the i n sects fro m t h e b u r
rows.
All fi n c hes are rat h e r s i m i la r
i n t h e i r g e n e ra l s i z e a n d h ave
d u l l p l u m a g e , probably because
of t h e d a r k volca n i c roc ks that
outcrop over m u c h of their ter
ritory.
The Ga l a pagos fi n c hes prob
ably a rose fro m an a n ce stra l
South A m erican f or m , b u t they
n ow d iffer so g reatly from a l l
e x i s t i n g m a i n l a n d fi n c h e s that
their a n cestry c a n not be recog
n i zed. This m a r ked d i ffe re n ce
s u g g ests
the
that
fi n ches
rea c h e d t h e i s l a n d s earl i e r than
oth e r s p e c ies, which are m u c h
c l o s e r t o t h e m a i n la n d for m s .
T h e a n cestra l fi n c h e s pres u m
a b l y rea c h e d t h e i s l a n d s b y be
ing " a c c i d e n ta l l y " carried t h e re ,
perh a ps a i d e d by ocea n i c cur
re n ts . It s ee m s most u n l i ke l y
t h a t t h e Ga l a p a g os I s l a n d s w e re
ever j o i n e d to t h e m a i n la d .

I n contrast t o t h e diversity
of t h e Ga l a pagos fi n c h es , Davi d
Lack has s h own t h a t i n the
n e i g h bori n g Cocos I s l a n d t h ere
is only a s i n g l e species of fi n c h ,
a l t h o u g h t h e i s l a n d prov i d e s
v a r i e d h a b itats a n d l a c k s m a ny
o t h e r typi ca l s pecies o f co n
t i n e ntal b i rd s . Th i s see m s t o re
s u l t fro m the s i n g l e i s l a n d of
Cocos l a c k i n g the n u m bers of
s m a l l , isolated e n v i ro n m e n ts t h a t
a re provi ded by t h e a rc h i pela g o
n a t u re of t h e Galapagos I s
l a n d s . ( See m a p p. 1 9 )

Tool-u s i n g Finch

( Amarhnyncluis pallid us)

N AT U R A L S E L E C T I O N I N A C T I O N is s h ow n b y t h e

p e p p e red m ot h ( p . 47), w h i c h i s a n exa m p l e of a

s p e c i e s t h a t h a s s h own a stri k i n g c h a n g e i n t h e fre
q u e n cy of a d a r k ( m e l a n i c ) fo rm in the last c e n t u ry .
T h i s m o t h wa s we l l k n own to t h e m a n y a m a t e u r
e n to m o lo g i sts i n B r i ta i n i n t h e ea rly yea rs of t h e 1 9t h
centu ry . U n t i l 1 8 4 5 , it was k n own o n l y i n t h e " p e p
p e r e d " fo r m , w h i c h h a s d a r k m a rk i n g s o n a l i g h t w i n g
b a c k g ro u n d . I n t h a t yea r, a d a r k fo r m w a s d i s covered
i n t h e i n d u stria l c ity of Ma n c heste r. At that ti m e t h e
d a rk fo r m m a d e u p l e ss t h a n 1 p e r c e n t o f t h e tota l
p o p u l a ti o n . With i n fifty yea rs i t m a d e u p 9 9 p e r c e n t
of t h e p o p u l a tion i n t h e Ma n c h e ste r a re a . T h e b l a c k
m o t h i s n ow t h e p re d o m i n a n t fo rm ove r m u c h o f E n g l a n d ,
a n d t h e o ri g i n a l p e p p e red fo r m i s a b u n d a n t o n l y i n
n o n - i n d ustri a l a re a s w h e re p o l l u t i o n h a s n ot b l a c k e n ed
t h e trees o n w h i c h t h e m o t h l ive s . I n s o m e of t h e n o n
i n d u s t r i a l ea ste r n a re a s o f t h e cou n t ry w h e re heavy s m o g
is ca r r ied i n , t h e b l a c k fo r m a l s o d o m i n a tes t h e ni oth
p o p u l a ti o n .
Rece n t s t u d i e s i n M a n c h ester
and other i n d u s t ri a l cities w h e re
stri n g e n t a n t i - p o l l ution o rd i
n a nces h ave b e e n e n force d ,
h ave s h ow n a revers a l of t h e
tre n d tow a r d s d a rker for m s ,
a n d a s l o w but m a rked i n
crease i n a b u n d a n ce o f t h e
l i g h ter peppered form s .
P E P P E R ED MOTH D I ST R I BU T I O N
I N B R I TA I N

( red ) M a j o r i n d u s tr i a l cities
o ( w h i te ) l i g h t form pred o m i
nant

( b l a c k ) d a r k form pred o m i
nant

( g ray ) i n te r m e d i a t e popu
lation

84

I N DUSTRIAL

MELAN I SM

(the

p red o m i n a n ce of d a r k varieties)
i s a l so s hown by a bout 70
oth e r species of m ot h s in
Europe. In the U n i ted Sta tes, the
Pittsburg h reg io n s h ows a com
para b l e predo m i n a n ce of once
rare black forms in a l m ost 1 00
species of m oths.
T h e c a u s e of i n d u strial m e l
a n i s m l ies i n t h e i n teraction o f
a d o m i n a n t g e n e , p ro d u c i n g t h e
b l a c k m u tatio n , a n d n a t u r a l s e
lection. H . B. D . Kettlewe l l h a s
d e m o n strated t h e i m porta n c e of
natura l selection by studyi n g
rates of b i rd predation o n t h e
two form s of t h e p e p pered
moth . H e released k n own n u m
bers of m a rked moths of e a c h
f o r m i n t w o a re a s a n d a n
a lyzed t h e n u m b e r of e a c h
f o r m that h e recaptu red by a t
tra c t i n g t h e m to a l i g h t a t

n i g ht. I n t h e i n d ustria l Birm i n g

h a m a rea, w h e re t h e l oca l pop
u lation has 90 percent black
forms, h e released 477 bla ck
a n d 1 37 l i ght i n d iv i d u a l s . He
reca ptured 40 p e rcent of t h e
b l a c k for m s b u t o n ly 1 9 percent
of the p e p pered.
In the u n poll uted coa sta l a rea
of Dorset, from a l most 1 ,000 of
t h e two colors of m o t h s re
leased, 6 perce n t of the b l a c k
a n d 1 2 . 5 perce n t of t h e l i g h t
form s w e re reca pture d . So i n
p o l l uted a r e a s , t h e b l a c k - w h i te
reca pture ratio w a s 2 : 1 . I n u n
pol l u ted a re a s i t w a s exactly
reversed-1 : 2 .
I n both a reas, caref u l ob
servation and fi l m i n g of b i rd s
eati n g t h e m ot h s from tree
tru n ks con fi r m ed
that these
ratios were t h e res u lts of rela
tive predation.

85

FOS S I LS a l s o d e m o n st r a te the effects of natura l se

l ectio n . B . K u rten h a s s hown its effects i n the E u ro p e a n

cave bea r (Ursus spelaeus) t h a t i n h a b ited north e r n E u
rope d u r i n g t h e P l e istoce n e . Ku rte n coll ected fos s i l s
from caves i n t h e Od essa reg i o n o f the U . S . S . R . B y
c o m p a r i n g these with s keleto ns of th e c l osely related
l ivi ng b e a r (Ursus arctos), h e wa s able to d iv i d e each
of the fossil popul atio ns i nto g rowth sta g e s . The fos s i l s
fro m a l l l o c a l ities s h owed stro n g s e p a ratio n i nto s i m i l a r,
yea rly g rowth stages, prob a b l y because t h e caves i n
wh i c h they were collected were i n h a bited o n l y d u ri n g
a n n u a l " h i b e r n atio n . " The a n a lysis o f these g rowth
stages d e m o nstrates the effects of natural sel ectio n .
Y O U N GEST STAGES of t h e cave

bear eq u a l in size to n ewborn

cubs of l i vi n g bears . a re rare,
proba b l y beca use of the f ra g i l
i ty of t h e b o n e s . T h e m o s t
a b u n d a n t g rowth s t a g e h a s a l l
t h e per m a n e n t teeth d ev e l o p e d ,
eq uiva l e n t to a b o u t 4 o r 5
m o n t h s i n l ivi n g bears, s u g gest
ing birth d u r i n g t h e period of
w i n ter d o r m a n cy, but a h i g h
m o rta l i ty o f c u b s toward t h e

end of this time. The next

g rowth s t a g e i s 1 . 4 y e a r s , i n
dicati n g t h e n e x t a n n ua l " h i
ber n a t io n . " A t 4 yea rs, bears
were f u l l y g rown . Life e x pecta
tion a t bi r t h w a s a b o u t 3 . 5
ye a r s ; m a x i m u m a g e , 1 8 years.
Divi d i n g the tota l n u m be r of
i n d ivi d ua ls i n a g iven age
g ro u p by the s u m of these a n d
a l l o l d e r i n d i v i d u a l s g i ves t h e
m o rta l i ty i n d ex f o r t h e g ro u p .

P l ei stoce n e Cave bear

86

1 000

- Ma n
Cave bear

------ o

1 00

- 1 00

.
1 00 2 0 0 300 4000

Age in % deviation from meon longevity

Survivors h i p c u rves from b i rth

for cave bea rs and m a n , s h ow
ing the s i m i l a rity of t h e c u rves .
( After K u rten . )

24

26

28

30

32

Mola r M ' Paracone I n d e x

Selecti o n pres s u re o n form of

paraco n e of seco n d molar teeth
of cave bear i s i n d i c a ted by
red u c t i o n of v a r i a t i o n a n d de
ve l o p m e n t of a rela tive l y s m a l l
e r paraco n e w i t h i n creas i n g
a g e . ( After Kurte n . )

Mola r M'
of Cave b e a r

TEETH of fossi l c a ve bea rs a re ve ry se n sitive to n a t u ra l

selection, because s u rviva l d e p e n d s o n s uccessful feed
ing p rio r to h i bernatio n .
K u rten stu d ied the form
of t h e seco n d molar teeth (M2) . The ratio 1 O O x the
l e n g t h of the l a rg est cusp ( pa ra co n e ) to the tota l l e n gt h
o f the seco n d m o l a r s h owed a sig n ifi c a n t c h a n g e , b e
c o m i n g s m a l l er with i n c reasi n g a g e . The you n g e r a g e
g ro ups were f a r m o re va ria b l e th a n th e o l d e r, b u t
fo r m s w i t h less wel l - a d a pted teeth we re e l i m i n ated .
O l d e r tooth m e a s u re m e n ts g ave c o m p a r a b l e resu lts.
S a m ples fro m oth e r caves g ave s i m i l a r b u t not i d e n t i c a l
resu l ts, s u g g esti n g d iffe rences i n s e l e ctive p ress u re
betwee n d iffere nt l o c a l e nviro n m e ntal n ic h es i n w h i c h
the b e a rs l ived .

87

ADAPTATION

i s t h e con t i n u i n g resu l t of n a t u ral

s e l ection . Ma n y o rg a n is m s a re very p recisely a d a pted
to p a rti cu l a r n i c h e s o r to p a rticu l a r ways of l ife . The
b i r d s of Hawaii p rovi d e a c l a s s i c i l l ustration of how a n
a ncestral g ro u p beco m es a d a pted t o s p e c i a l n i c h e s i n
a n e w e nviro n m e nt. Hawa i i cons ists o f a g ro u p o f iso
l a te d vol ca n ic isl a n d s situated i n the m id - Pa c ifi c . L i k e
m ost ocea n i c is l a n d s, the n u m b e r o f l a n d b i rd s is s m a l l .
Th e h i g h d e g re e o f specia l i zation o f s o m e of t h e b i rd s
m a kes t h e m v u l n e r a b l e t o e nv i ro n m e n t a l c h a n g e .

Ko uoi okepo

LOBELIA

Hawaii akialoa
(Hemignalhus obscurus ohscurus)

o r drepa n i d i d s ,
a re a fa m i ly of b i rds fou n d only
i n Hawa i i . L i ke D a rwi n 's fi n ches
i n t h e Ga l a p a g os I s l a n d s (p.
8 3 ) , t h ey h ave a d a pted t h e m
se lves to a wide d i v e r s i ty of
conditions. The ori g i n a l for m s
se e m to h ave f e d o n the i n
sects a n d n e ctar of s h ortbe l l e d
flowers. T h ey p roba b ly h a d

S I CKLEB I LLS,

88

s h ort, s l e n d e r beaks. O t h e r
fo r m s s how d i versity of a d a pta
t i o n s to d i ffere n t d iets, i n c l u d
i n g s o m e w i t h re m a r k a b l y l o n g ,
curved b e a k s f o r feed i n g o n
the l o n g , t u b u l a r fl owers of
t h e Hawa i i a n l o b e l i a . Some of
the various for m s of s i c k l e b i l l s
a re i l l ustrated . N i n e of t h e 22
k n own species a re n o w extinct.

Pseu donestor

Psittirostra kona

H e m i g n a thus o b s c uru s procerus

Hemignathus lucidus

P ro b a b l e evo l ut io n of o n e g ro u p of d re p a n i d i d s from
a n a n cestor, p o s s i b l y s i m i l a r to t h e H o n eyea t e r ( p . 8 8 )
i s s h own a b ove . D repa n i d i d bea k s s h ow a d o p ti o n to
va r i o u s d i ets . H e mign a t h us o b scurus u s e s its e l o n g a ted
b e a k c h iefly to p ro b e fo r i n sects in b a r k caviti e s , a l
t h o u g h o t h e r s p eci e s u s e t h e i r l o n g " s i c k l e - b i l l s " c h i efly
to p r o b e l o b e l i a f l owers for n e cta r. The n ow ext i n ct H .
lucidus had a s h o rt l ow e r m a n d i b l e , w h i c h i s r e d uced
even fu rth e r in H . wilso n i, which uses it, wood p e c k e r
fa s h io n , a s a c h i s e l . T h e l o n g tu b u l a r to n g u e s of i n sect
eati n g s pe c i e s refl ect t h e i r d eve l o p m e n t f ro m n e cta r
fee d i n g fo r m s . Pseudon estor has a p a r rot l i k e b i l l , w h i l e
seed-eati n g s p e c i e s , s u c h a s Psittirostra k o n a , h a v e h e a vy
fi n c h - l i ke bea k s .
89

MIMICRY i s rel atively com mon

i n i nsects a n d in
so m e fl owers . In i n s ects it see m s to h ave a risen c h i efly
fo r defense; in flowers, for po l l i n ation . The i n fl u e n ce of
n atura l selection in the deve l o p m e n t of m i m icry is
s h own by the d i stribution of m i m i c s pecies o n ly i n
a reas where t h e i r models a re co m m o n . Where the
m o d e l s a re a b u n d a nt, the m i m ics s h ow g reater vari
a b i l ity, w h i c h c a n be exp l a i n e d o n ly by assu m i n g th a t
t h e l owe r predato r fa m i l i a rity with t h e m o d e l s i n s u c h
a reas h a s p ro d uced l ower selection p ressure o n t h e
m i m ics. Develop m e nt o f m i m ics d e p e n d s n o t o n " acci
d e n ta l " p a ra l l e l m utatio ns but on a series of i n teracti ng
ge nes that h ave u ndergone strong sel ection .

THE S L I P P E R O R C H I D o f Eu rope

and the M i d d l e East e m its a

perf u m e that attracts a wasp
wh i c h pol l i nates it. Other re
lated
Ophrys
orc h i d s
have
bri g h t, i n sectlike flowers t h a t
aid i n a ttracti n g m a l e wasps.

I N S E CTS h ave developed v a r

ious protective s h apes a n d col
o rs . The h a r m less b u m bl ebee
m o t h , for exa m p le, i s a m i m i c
of t h e s ti n g i n g b u m b l ebee. The
tree hoppers below look l i ke
thorns.

Tree h opper on
rose stem

Ophrys orchid

a n d pol li n ating

wasp

90

S EX UAL S ELECT I ON i s t h e s u m of va rious cha ra cters

by wh ich o n e p a rtner, gen erally the m a l e , of a spe

cies attracts a m ate a n d repels riva l s of the sa m e sex.
S i n ce it i n creases the i n d ivi d u a l ' s repro d u ctive produc
tivity, it con stitutes a d i sti n ct ki n d of sel ecti o n . Ela bo
rate m a l e p l u m a g e a n d song in b i rds, courts h i p d i s
pl ays, m o re i n ten se m a l e coloratio n, g reater s i z e a n d
a n tlers i n various a n i m a ls-these a re so m e o f t h e c h a r
a cteristics t h a t m ay resu lt. Th e rel ative i m porta n ce of
sexua l selectio n a s a component of natural selection is
sti l l not clea r . D a rw i n was convi n ced it was of m a jor
i m po rta n ce, but su bseq u e n t writers h ave been less sure.

MALE S U PERB LYREBI RD of A u s

tra l i a sta n ds w i t h i t s e l a bora te

ta i l feathers exte n ded a n d q u iv
ering, as it perform s its cou rt
s h i p d i s p l ay. T h e f e m a l e re
s e m b l e s the m a l e i n color but
l a c k s the special ized tai l feath
ers. You n g male b i rds are l i ke
the fe m a les. T h ey do not de
velop the ta i l feathers u nt i l t h ey
a re a bout t h ree yea rs old.

Each male esta b l i s h es a l a rg e

territory, a n d t h ro u g h o u t t h i s
a rea, h e b u i l d s a series o f
m o u n d s of ea rth , each a bout a
ya rd i n d i a m eter. H e patro l s
h i s territo ry by v i s i t i n g each
mou n d and pe rfor m i n g his d i s
play on it. T h e d i s play beg i n s
w i t h a voca l perform a n c e that
i n cludes a series of m i m i c k i n g
so u n d s a n d c a l l s .

91

N AT U RAL S ELECT I O N I N MA N i s re d uced i n i ts o b

vious effects by t h e stru ctu res of society a n d b y the

e l a b o rate patte r n s of fa m i ly c a r e . Even so, m a n y body
fe atu res show t h e i n fl ue n ce of n a t u r a l selectio n . S o m e
g e n e c h a n g es a l so d e m o n strate i t s effects. S i c k l e- c e l l
a n e m i a , a d isease c a u s e d by a s i n g l e g e n e , is m a rk e d
b y d i sto rtion of t h e b lood c e l l s fro m t h e i r n o r m a l d i s c
s h a p e t o s i c k l e s h a pe w h i c h b l o c k s t h e fl ow of b l o o d i n
c a pi l l a ries a n d t h u s cau ses a n e m i a . S o m eti m es d eath
results when sufferers a re s h o rt of b reath o r wor k at
h i g h a ltitu d e s . In West and Central Afri c a , w h e re m a
l a ri a i s e n d e m i c , po pulations with a s i c k l e- g e n e e q u i
l i b ri u m freq u e n cy of u p t o 2 0 perce n t a re com m o n .
A m o n g desce n d a nts of th ese people i n the U n ited
States, freq u e n cy of sickle-cell a n e m ia has d ropped to
9 perce n t in two ce nturies. How has this co m e a bout?
T h e fre q u e n cy of a sickle-ce l l g e n e i n Africa i s p lotted be low a s
p e rce n t of pop u l a t i o n . H ig h freq u e n c i e s a re re stricted t o e q u a tor ia l
a re a s i n w h i c h tertia n ma laria is a n i m porta n t c a u s e of death .
N o r t h a n d s o u t h of t h i s b e l t , m a la ria is less c o m m o n a n d i s
b e n i g n . S i m i l a r h i g h f r e q u e n c i e s occur i n m a l a r i a l a reas of S i c i ly,
G reece, T u r key, a n d I n d i a . (Afte r All ison.)

Eq u a tor
1 0- 1 5%
5 - 1 0%
0-5%

92

S I CKLE G E N E w a s demon
strated b y b iologist Ver n o n
I ng ra m to h ave t h e a b i l ity t o
change o n e of t h e t h re e h u n
d red a m i n o a c i d u n its i n t h e
h e m o g l ob i n m o l e c u l e s t h a t m a k e
u p t h e r e d b lood c e l l s . A t o n e
point i n t h e c h a i n of n i n e te e n
d i ffere n t a m i n o a c i d s t h a t m a ke
u p the prote i n , va l i n e is s u b
stituted for g l u ta m i c a c i d . T h e
res u l t i s t h a t n or m a l red b l ood
corpuscles beco m e s i c k l e -s h a p e d .
THE

s i c k le genes
have c o m pa ra t i v e l y l i tt l e a d
verse effect u n d e r n o r m a l co n
d i t i ons, b u t t h e h o m ozyg o u s
co n d i tion has far m o r e seriou s
effects, ofte n ca u s i n g blood
c l ott i n g or p re m a t u re deat h .
Why d o s i c k l e g e n e s sti l l per
s i st i f n a t u ra l s e l e c t i o n i s effec
tive? I t so h a p p e n s t h a t s i c k l e
s h a pe d h e mog l o b i n i s resista n t
t o i n fection b y t h e m a l a r i a l
parasite
Plasmodium,
which
feeds o n r e d b l ood c e l l s . I n
m a l a r i a l a reas, t h e refore, a se
lective e q u i l i b r i u m e x i sts, with
t h e heterozyg o u s s i c k l e - ce l l i n
d iv i d u a l s re s i sta n t a n d th u s
favore d . T h e h o m ozyg o u s i n
d i v i d u a l s (w i t h
2 s i c k l e - ce l l
H E TEROZY G O U S

Amino acids
in normal
hemoglobin

Amino acids
i n s i c k l e cell
hemoglobin

g e n e s) often d i e. A n e q u i l i b r i u m
tends to occur betwe e n t h e
n u m be r of peo p l e w i t h n o
s i c k le - c e l l g e n e ( t h e s e a re t h e
" n orma l " i n d i v i d u a l s) w h o d i e
of m a la r i a a n d t h o s e w i t h two
s i c k l e - ce l l g e n e s w h o d i e of
a n e m i a o r b l ood c l o t t i n g . I n
t h i s c a s e , a n a p pa re n t l y h a r m
f u l g e n e t i c effect m a y be pre
served beca u s e of side b e n e fi t s .
I n m a l a ri a - f re e U . S . , t h e co n
d i t i o n h a s n o s u rv i v a l v a l u e .
T h u s , n a t u ra l s e l ec t i o n repre
se nts not a b l i n d , c r u s h i n g wave
of exti n c t i o n but the reso l u t i o n
of confl icti n g e n v i ro n m e n t a l d e
mands on a population. It in
volves s u b t l e i n te r p l a y of i n
tern a l a n d e x ter n a l i n fl u e n c e s .

93

( p . 5 2 ) c o n fi r m t h e a ction of n at u ra l
s e l ectio n a n d d e m o n st ra te t h e way i n w h i c h i t o p e r
a tes . O n ce i t was a rg ued t h a t n a t u ra l sel ecti o n co u l d
n o t poss i b l y h ave p rod u ced t h e e l a b o rate co m p l e x of
c h a nges requ i red to b r i n g a bout s u c h m a j o r evo l utio n
a ry c h a n ges as the d evelop m e n t of a m p hi b i a n s from
fish o r b i r d s from reptiles. Tra nsitio n a l fossil for m s, o r
" m issing l i n ks, " s h ow h o w the p rocess took p l a c e .

M I S SI N G LI NKS

A Devo n i a n lobef i n crossopte

ryg i a n fish c l i m bs a s hore.

Labyri nthodont a m phibian, a de

s ce n d a n t, goes a s h ore c l u m s i ly.

RANDOM

52-53) s how that c h a n g e from

one m a j o r g ro u p to a n other was
a piece m e a l process. T h e re was
not one big j u m p or a syn
c h ro n ized deve l o p m e n t of a l l
t h e va rious c h a racters i nvolved .
I n A rchaeopteryx, some featu res,
s u c h as t h e bra i n , c l aws, teeth ,
a n d ster n u m , were p r i m itive a n d
repti l i a n ; others, such a s w i n g s
a n d fea t h e rs a n d t h e g e n era l
body s h a pe, were a l ready f u l l y
b i rd l i ke . T h i s j u m b l e d mosaic
evo l u t i o n i s exactly what we
should ex pect i f natural se lec
tion h a s bee n an effective a g e n t
of change.

NATURAL

S E LEC

T I O N , the critics o n ce a rg u ed,

cou ld n ot have produced t h e

ela borate series of variations
that g ra d u a l ly t ra n sfor m e d rep
t i l i a n a r m s i nto b i r d w i n g s a n d
gave b i r d s a l i g h t a n d mod ified
skeleto n , feathers , and s i m i la r
fea tures . E a c h of t h e s e e n d less
m i nor
mod ifications, i t was
thought, m i g h t offe r n o special
advantage-not e n o u g h to bring
a bout the va r ious s u pposedly
i n terd e p e n d e n t a n d coord i n a ted
c h a n g e s that were i nvolve d .
B u t Archaeopteryx a n d other
tra n sitio n a l fos s i l forms (pp.

94

MOSAIC EVO LU T I O N is shown

wel l i n the i c h t hyosteg ids, pri m

itive a m p h i b i a n s of t h e D e
von ia n (p. 98). T h ey h a d many
fi s h l i ke f e a t u re s . T h e c h a rt g i ves

"'

75
1 00
1 00
1 00
1 00

a b reakdown of t h e i r a m p h i b
i a n vers us fi s h l i ke c h a racteri s
tics, with 1 00 poi n ts f o r f u l ly
developed a m p h i b i a n form a n d
0 for fu l ly fi s h l i ke for m .

a
li

.c

...

Skull tlructure
Eye._titlon of

limbs
Shoulder and hlp girdle
Skull articulation
Rlbt-form of

lchthyoste ga, a pri m itive De

vo n i a n a m p h i b i a n from East
Gree n l a n d . About 3 feet lo n g .
A reco n s tructi o n .

Skel eton of lch thyostega prob

a bly rese m b l es c l osely that of
the a n cestors from w h i c h
p h i b i a n s evolved.

D iplove rteb r o n, a l a byri nthodont

a m p h i b i a n of t h e Carbo n i ferous,
with many rese m b l a n ces to the
i c h thyostegids.

Eusth enopteron,
a
Devo n i a n
lobefi n
crosso pteryg i a n
fi s h
2 feet l o n g , h a s m a n y rese m
bla nces t o t h e ichthyostegids.

95

is the fi n a l co m po n e n t of the evo l u t io n a ry

recipe . Ea rly oppon ents of evolution conte n d e d t h a t
the age o f the ea rth, t h e n reg a rded a s l e s s t h a n 4 0
m i l l io n yea rs, d i d n o t a l low enou g h t i m e fo r t h e s l ow
process of c h a n g e that evol ution i nvolve d . These es
t i m ates were based o n t h e rate of coo l i n g of a s u p
pose d l y o n ce m o lten e a rth . But t h e d eve l o p m e n t of
othe r m et h o d s of dati n g s u g g ested a m u c h g reater
a g e, a n d t h e use of ra d io a ctivity n ow i n d i cates that
t h e e a rt h i s p ro b a b ly about 5 b i l l i o n yea rs o l d . This is
a m pl e ti m e fo r evo l ution to have taken p l a c e . T h e d e
ve l o p m e nt of a geologic t i m e sca l e puts the fos s i l rec
ord i n to perspective, a n d reveals the order a n d se
q ue n ce of t h e va riou s fo r m s of l ife .
TIME

THE GEOLOGIC
CLOCK
S h ows last
600 m i l l ion
years of earth ' s
h i s tory, each
hour representing
5 0 m i l lion years.

u s to a p p re
ciate the e n o r m o u s i n terva l of
t i m e t h i s represe n ts . S u p pose a
cos m ic h i storia n wri t i n g a h i s
tory o f t h e e a r t h beg a n t o
write a t t h e creation of t h e
e a r t h a n d wrote o n e l i n e every
t h o u s a n d years. If the boo ks h e
p rodu ced were t h e s i z e o f t h i s
o n e y o u a re rea d i n g , h e wou l d
b y now h ave pro d u c e d 94,000
boo ks.

ANALO G I E S h e l p

96

Most Fossils

ELEME NTS, such

as u ra n i u m a n d rad i u m , have
u nstable a to m i c n u c l e i that u n
dergo s p o n ta n e o u s breakdow n
at a consta n t, m e a s urable rate
to form oth e r, more sta b l e e l e
m e n ts . U ra n i u m , for e xa m p le ,
p rod uces a s e r i e s of " d a u g h
t e r " e l e m e nts a n d fi n a l ly y i e l d s
l e a d a n d h e l i u m . O n e g ra m
of u ra n i u m prod uces 1 / 7000
g ra m s of lead every m i l l i o n
yea rs. T h i s r a t e i s u n affected
by any c h a n ge s in heat o r
pres s ure. M e a s u re m e n t o f t h e
ra tio o f " o l d " u ra n i u m t o " n ew"
RADIOACTIVE

lead i n u ra n i u m m i n e ra l s pro
vides an i n d ication o f the age
of the rocks in w h i c h t h ey are
fou n d .
O t h e r r adi oa c t i ve e l e m e n ts
used in age m e a s u re m e n ts i n
c l u d e lead-thori u m , pota s s i u m
a rgon , ru b i d i u m -stro n t i u m , a n d
carbon . S t u d i e s o f m eteorites,
w h i c h seem to b e " left over"
fra g m e n t s fro m the d evelop
ment of th e solar syste m , a n d
t h e rate of ex pa n s i o n of t h e
u n iverse te n d to co n f i r m a f i g u re
of a bo u t 4.5-5.0 b i l l i on years for
t h e ag e of the e a rt h .
URAN I U M T O LEAD
B R E A K DOWN

Oldest
u n d isp uted
fossil

SCALI N G D O W N EARTH'S H I S
TORY i n to a model s i n g l e ca l

e n d a r year w i t h t h e orig i n of
t h e e a rt h on J a n u a ry 1 st a n d
t h e p res e n t day o n Dece m be r
3 1 st wou l d m a ke e a c h second
e q u i v a l e n t to 1 67 years a n d
e a c h m i n ute to 1 0,000 years.

The o l d e s t u n d i s p u te d fos
s i l s wou l d a p pe a r about J u ly
1 ; t h e o l d e s t a b u n d a n t fos s i l s ,
a b o u t Nove m b e r 1 8 . Ma n wo u l d
a p p e a r a t a b o u t 1 1 :50 p . m . o n
Dece m be r 3 1 . A l l record e d h i s
tory wo u l d f a l l i n t h e fi n a l 4 0
seco n d s o f t h e ye a r.

97

"'

PERIOD

Q UATERN A R Y

;t

63

62

T ERTI ARY

C R ET A C E O U S

1 36

71

J U RASSI C

1 90

54

225

35

TRIASSIC

PERMIAN

280

55

325

45

345

20

P E N N S Y LV A N I A N

D EV O N I A N

0
N

395

50

430

35

M I SS I S S I P P I AN

SILU RIAN

O RDOVICIAN

500

70

CAMBRIAN

570
z
<(
"'
"'

0.. u

70

P R EC A M B R I AN

4,030

THE G EOLOG I C TIME S C A L E

H E G E O LO G I C TIME S CALE d e

r- r-

r-

f--

f--

f--

f--

f--

t-

+-

---

---

---

--

--

--

--

--

--

--

----

_____.....___.
.
....____.
.
________

ope d fro m a study of stratified

rocks a n d their fossils, is d ivided
i n to fou r e ras, fos s i l s b e i n g a b u n
d a n t o n ly i n t h e l a st th ree . The
n a m es of m ost of t h e periods a re
d e rived fro m p l a ce s w h e re rocks of
t h a t a g e were fi rst stud ied . Thus,
we spea k of Devo n i a n fi s h just as
we spe a k of Rom a n a rc h itectu re .
Wh e n t h e i r rela tive s e q u e n c e is
k n own, both ca n t h e n b e fitted i n to
a n u m e r i c a l ti m e sca l e . Preca m
bria n t i m e covers a l m os t 9 I 1 Oth s
of a I I e a rth h i story .

RATES OF EVO LUTIO N v a ry e n o r m o u s l y fro m o n e

species t o a nother. The s m a l l b r a c h i pod Lingula, a n i n

h a b itc;mt o f wa r m , s h a l low seas, h a s sca rcely c h a n ged

i n t h e l a st 400 m i l l io n yea rs. But t h e s pecta c u l a r d i
versification o f m a m m a l s has ta ken p l a ce with i n t h e
l a st 60 m i l l i o n yea r s . Develo p m e n t of a g eolog i c ti m e
scale a l l ows u s to a n a lyze a n d i n terp ret t h ese d ifferi n g
rates, wh ich m a y b e expressed i n various ways .
IN
STR U CTU R ES,
rates
of
c h a n g e can b e m e a s u red for
some foss i l g ro u p s . In the evo
lution of early Tertiary h orses,
c h a n g e s i n d i m e n s i o ns of molar
teeth were o n ly 0. 1 5 mm per
m i l l ion years. T h is is about
equal to t h e d i a meter of a
h u m a n hai r . S i n g l e populations
ofte n conta i n e d u p to twe n ty
t i m es as m u c h variation i n tooth
d i m e n s i o n s . S u c h s l ow tra n s for
m ation ca n be observed o n ly i n
foss i l s .

TRA N S FO RMATI O N O F S P E C I E S

E C O L O G I C A L R E PLAC EMENT o f

R A P I D D I V E R S I FI C A T I O N of
Mioce ne horses f ro m b rows i n g
to g razi n g correspo n ded t o
t h e w i despread c h a n g e fro m
lowland forests to upla nd
pra iries i n North A m erica a n d
t h e f i rst appeara n c e of fos s i l
g rasses. T h i s i n teraction see m s
t o conf i r m t h e i m po rta n ce of
n a t u ra l selection (p. 1 0 1 ).

s o m e ext i n ct g rou ps b y oth e rs

of s i m i la r e n v i ro n m e ntal h a b i ts
s u g g ests that com pet ition be
twe e n the two g roups may have
been a factor in some cases of
exti n ct i o n . T h e width of each
co l u m n in t h e d ia g ra m i s p ro
portional to the d iversity of
t h e group.

1 00

fro m one i n to a n other for Ter

t i a ry m a m m a l s (p. 1 3 2) h a s
b e e n calcu lated to be a bout
500,000 yea rs. With such s low
rates, it is not surpri s i n g that
few exa m p les of t h e develop
ment of n ew species can be
observed i n l i v i n g populations.
S u c h rates also a l low a m pl e
t i m e f o r t h e operation of
" s low" evo l u t i o n ary
mech a
n i s m s , s u c h a s t h e selection o f
s m a l l va riations.

BROWSI N G TEETH
( low.crowned- no ceme n t )

G R A Z I N G TEETH
( H i g h crown ed-cemen t )

H Y PO H I P P U S

P A RA H I P P U S

ANCHITHERI UM

EVO LUTION O F H O RSES

( Teeth drawn to scale-After S i m pson )

PATTE RNS O F C H AN G E beco m e

e v i d e n t i n s o m e g ro u p s w h e n t h e
time factor is considered. The
ra pid d i ve rs i f i cation of h orses i n
M i o c e n e t i m e s represe n ted a
c h a n g e i n fee d i n g h a b i t f ro m
brows i n g to g ra z i n g , correspo n d
i n g to t h e w i d e spread c h a n g e
f ro m l ow l a n d forests to u p l a n d
pra i ries i n North A m e rica a n d
t h e f i rst a p pe a ra n ce of fo s s i l
g ra s s es. T h i s see m s to be a n i n
te raction co n f i r m i n g t h e i m por
ta n c e of n a t u ra l selection ( p . 5 1 ).

1 01

s h owi n g t h e i n te r p l a y
o f t h e va rious fa cto rs i n volve d , c a n b e s u m m e d u p fo r
a ny popu l a tion a s d e m o nstrated i n the d ia g ra m below.
Such a si m pl e recipe does not m ea n tha t evo l ution i s
itself s i m p l e o r t h a t it fol lows a c l e a r ly predicta b l e p a t
tern . The reverse is the case. The i n teraction of t h ese
va rious p rocesses pro d u ces an eno rm o u sl y c o m plex dy
n a m ic syste m . Both the co m p l exity a n d the potenti a l ity
fo r n ovelty of the evo l utio n a ry process a re i n d i cated
by t h e g reat d iversity of l ivi n g thi n g s .

A R E C I PE FOR EVOLUT I O N

I SOLATION :

VARIATION :
Genetic
Reco m b i nation

Over-Production
Limited Food Supply
NEW SPEC I ES

EVOLU T I O N S T I LL C O N T I N U E S .

Many g eo g ra p h i c races a re po
te ntia l ly new s pecies i n t h e
m a k i n g . Co n t i n ued i s o l a t i o n o f
r a c e s of t h e g o l d e n w h i s t l e r i n

1 02

t h e Solo m o n I s l a n d s
l o n g i n g to a s i n g l e

a l l be
s pecies,
Pachycephala pe ctoralis, wo u l d
pro b a b l y c o n v e r t t h e m i n to re
prod u c t i v e l y i s o l a te d s pecies .

MUTAT I O N S a re i m po rta n t i n
t h e evo l u t i o n a ry process, eve n
though so m a ny i n l i v i n g popu
lations see m to be h a r m f u l . I n
a population w e l l -a d a pted to a
part i c u l a r
e n v i ro n m e n t ,
the
most b e n efi c i a l m u tations have
ofte n a l ready bee n i n corporat
ed. In a d d i t i o n , m u tations with
a re
effects
visible
l i m ited
known to be co m mo n .

N o t a l l h a r m f u l tra its a re
e l i m i nated from t h e g e n e pool
of a population . Selection is
a l ways a c o m p ro m i se . Even
h a r m f u l c h a racteristics may have
s o m e be n e fi c i a l s i d e effects (p.
93). Prese rvation of such g e n e s
provides a reservo i r of pote n
t i a l c h a n g e that may be o f
critica l i m porta n c e if e n v i ro n
m e n ta l co n d itions c h a n g e .

COLOR CHANGES I N M I N K o r e t h e res u l t o f m utations.

FACTORS A C CO U N T I N G F O R
N E W S P E C I E S orig i n s e e m a d e

q u a t e to a ccou n t a l so for a l l
evol u t i o n a ry
change.
So m e
writers re.f er t o m i c ro-evol u t i o n
. a n d m a c ro-evol u t i o n , b u t t h e s e
are not f u n d a m e n ta l l y d i ffere n t.
C u m u l a t i ve devel o p m e n t of new

- 5

species leads to w h a t we later

classify a s n ew g e n era a n d
h i g h e r g ro u p s .
Sepa ra t i o n of desce n d a n ts
from a n ce s t ra l po p u la.ti o n s by
ti m e re i n forces g e o g ra p h i c d i f
fere n ces be tween co n te m pora ry
race s , as s h o w n below.

4
2

3
II
IV

ARAB I C N O ' S .
( S PEC I ES)

l i ne-Evol ution of o Species

ROMAN NO ' S .

C o n t i n u e s to E xist
Becomes Extinct

(GENERA)

1 03

T H E C O U RS E OF EVOLUTION

Pre-org a n i c evo l ution t h a t occu rred before t h e a p pe a r

a n ce of life o n e a rth left l ittle d i rect evi d e n ce of the
p rocesses i nvolved . Much s m a l l e r m o l ecu l es were i n
vo lved t h a n i n o rg a n ic evo l u ti o n , h owever, a n d the
process resu lted i n bod ies of m u c h g reater d i m en sions,
s u c h a s stars and g a l axies. The o ri g i n a l m ateri a l p ro b
a b ly co n s i sted of su b-ato m i c p a rticles, s u c h a s n e u
tro n s , p roto n s, a n d el ectrons, that l a te r p rod uced hy
d ro g e n . Much of the m ate rial in the vi s i b l e u n ive rse
see m s to be hydrogen, which cons ists of a s i n g l e p ro
to n . Heavier e l e m e nts were pro b a b l y produced from
the l i g hter hyd rog e n by a p rocess of neutron ca pture.
Each neutron t h a t was a d d ed yiel d e d a new isoto pe.
The p rocess by which hyd rog e n a g g reg ated to fo rm
bodies s u c h a s stars, i n which e l e m ent- b u i l d i n g too k
p l a ce, is sti l l t h e s u b j ect of speculation . It m ay be a
conti n u o u s p rocess i n w h i c h n ew hydrogen i s b e i n g
created con sta ntly. Altern atively the a p p a re n t exp a n
sion of t h e u n iverse m ay result fro m t h e " b ig b a n g " o f
a s i n g l e e p i s o d e of creation about 5 t o 1 0 b i l l io n years
a go. The u n iverse m ig h t u n d er go p u l s a t i n g m ove m e nts,
so t h a t t h e p rese n t e x p a n si o n wou l d b e fo l l owe d by
co n t ra ctio n . The " b ig b a n g " h y p o t h eses te n d s to be
t h e c u rre n t fa vo rite .
a n d t h e re s t of t h e s o l a r system pro b a b l y o r ig i n a ted by
the a g g r e g a t i o n of a c l o u d of cos m i c d u s t . Earth ' s s t r u c t u res s u g
g e s t t h a t i t formed fro m c o l d ra ther t h a n fro m m o l te n m a teria l .
E A RTH

THE PR I M IT IVE EARTH o n wh ich l ife o ri g i n a ted was

an enviro n m en t very d i fferent fro m a n y o n ea rth today.

Th ree l i n es of evi d e n c e s u g g est that the e a rt h ' s p r i m i
tive a t m os p h e re p r o b a b l y co n s i sted of hyd rog e n , he
l i u m , m et h a n e, a n d a m m o n i a . The present a t m o s p h ere
of n itro g e n , ca rbon d ioxi de, a n d oxyg en ca m e l ater.
METEOR ITES a pp e a r t o be ma
terial " l eft over" f ro m the orig i n
of t h e solar syste m . A n a lys is o f
their composition p rovides a n
i n d ication o f t h e poss i b l e " b u l k
co m pos i t i o n " o f t h e earth. Most
m eteorites a re m a d e of i ron
n i ckel or "stony" m ateri a l . A
few (the carbo n a ceous c h o n
dr ites) conta i n c a r b o n co m
pou n d s of extrate rrestrial orig i n .

3 4 - to n m e teori te
f r o m G r e e n Ia nd

A NALYS I S of
l i g h t f ro m other p l a nets s h ows
that the s i x " ba s i c " e l e m e n ts of
l i v i n g t h i n g s a re w i d e l y d i strib
uted. Hydrog e n , oxyg e n , carbo n ,
a n d n i trog e n a re a m o n g t h e
m o s t a b u n d a n t e l e m e nts i n the
solar syste m . Sulfur i s n i nt h ;

SPECTRO S C O P I C

p h o s p h o rou s , s i x teen t h . T h i s i m
p l i e s t h ey w e r e a l s o proba b l y
presen t in t h e p r i m i t i ve e a r t h .

Dark l i n es i n s pectru m a n a lysis

a re prod u ced by a bsorption by
coo ler g a s i n f ro n t of g lo w i n g
sou r c e .
TH E ATMOS PH E R ES of p l an ets
m ost d i stan t f r o m t h e s u n h a v e
p rob a b l y c h a n g ed t h e l ea st .
T h e i r a t m os p h e res i n c l ude h ydrog en , h e l i u m , m et h a n e ( C H 4 ) and,
in J u p i ter and S a t u rn , a m m on i a
( N H 3 ). Proba b l e c h an g e s in t h e
eart h ' s a t m o s p h e re (water o m i t
ted) a re b a s ed p a r t l y on a n a l o g y
w i t h t h e s e.

PA L EO Z O I C
....
Cl)

z 0
""'
<(

:;

o..

T H E O RI G I N OF LI FE m u st h a ve i n vo l ve d t h e deve l o p

m e n t o f protei n s fro m thei r a m i no-acid co m po n e n t s .

An experi m e n t b y Sta n l ey Mi l l e r a n d H a ro l d U rey d e m
on stra tes o n e poss i b l e way i n w h i c h t h i s m i g ht h ave
ta ken p l a ce o n the p r i m itive ea rth .
AMMO N I A, meth a n e, h y d rogen,

and steam were m i xed together

i n a closed c i r c u l a t i n g system

and then s u b j ected to a n elec

trica l d i s c h a rg e . Afte r severa l
days t h e c o n d e n s e d water was
fou n d to conta i n a m i xtu re of
a m i n o a c i ds. The effects of
l i g h t n i n g o n t h e p r i m itive at
mosph ere may have prod u ced a
s i m i l a r n o n - biological synt h e s i s
o f orga n i c m o l e c u l e s .
_

M i l ler- U rey a p paratus, show n i n

d i a g ra m matic for m .

1 06

I n orga n i c synthesis of other

molecules, s u c h a s carbohydrates
and n uc l e i c a c i d s , h a s also been
d e m o nstrated.
va rious
These
co m pou n d s were pro b a b l y pre
served o n the early e a rth by
t h e a bs e n ce of oxyg e n and o f
other l iv i n g t h i n g s .

A LL LIVI N G T H I N GS tod ay de

pend, d i rectly or i n d i rectly,

u po n g ree n p l a n ts for t h e i r
foo d . T h e e a r l i e s t orga n i s m s
pro b a b l y " fed " b y s o m e fer
m e n tatio n - l i ke proces s u po n t h e
o rga nic " b roth" from wh ich
they a rose, b u t t h i s food
s o u rce was l i m i te d . C h a n g es i n
t h e earth ' s a t m o s p h ere ca used
both by s o l a r radiation and by
t h e effects of t h e e a r l i e s t or
g a n is m s produced a n e n v i ro n
m e n t w i t h i n creas i n g q u a n t ities
of n i trog e n and carbon dioxide.
T h i s proba b l y e n co u raged the
a l tern ative
of
develop m e n t
fee d i n g m ec h a n i s m s , i n vo l vi n g
fi rs t t h e s y n t h e s i s of m o re co m
p l e x m o l e c u l e s . T h e m ore s o
p h i s ticated process of p h oto s y n
thesis in which s u n light sup
p l i e s t h e e n e r g y f o r t h e co n
vers i o n o f a t m o s p h eric carbo n
carbo h y d rates
i n to
dioxide
ca m e l a ter. P h otosy n t h es i s re
l e a s e s o x yg e n , s o t h e e a r l y a t
m o s p h e re u n d e rw e n t tra n sfor
m a tion from a red u ci n g to a n
oxyge n a t i n g e n v i ro n m e n t .

s o l a r ra d i a t i o n w a s i n crea s i n g l y
f i l tered o u t a s r e l e a s e of f ree
oxyg e n p r o d u ced o I o ye r of at
m o s p h e r i c ozo n e . C ol o n i za t i o n of
s u rface waters , a n d l a t e r o f t h e
l a n d , d e p e n d e d o n t h e g rowi n g
effective n e s s
ozone
the
of
scree n . T h e " s u d d e n " a p pea r
a n ce of h a rd - bod i e d i n vertebrate
a n i m a l s in Early C a m br i a n t i m e s
m a y reflect t h e d ev e l o p m e n t o f
t h e ozo n e I o y e r , w h i c h p rovi d ed
a p rotec t i ve e n v i ro n m e n t.

R E PROD U C T I O N a rose by d u
p l i cation of I o r g e m o l e c u l a r a g
g re g a t i o n s by a u tocata l y s i s , i n
w h i c h e l e c t r i c a l l y a ct i ve co m
poun d s , s u c h a s p rote i n s , cou l d
prec i p i ta te d ro p l e ts of col l o i d a l
a g g r e g a t e s t h a t were c a p a b l e
of d eve l o p i n g i n to o s u rface
m e m b ra n e .
W h e n i n c rea s i n g oxyg e n i n
t h e a t m o s p h ere r e a c h e d a l evel
to a l low d e ve l o p m e n t o f res
s p i ro t i o n ,
harmful
u l tra v i o l et

bi I l i o n s of years

EVOLUT I O N OF L I F E
V

befo re pres e n t

-T__________________.______,

-0-

"'
w
z

<
-+------
:r:
...

z
0

....
;j
......

>
w
......

EVOLUT I O N O F EARTH'S
ATMOSPHER E & HYDROSPHER E

E a rl i es t
a n i mals

known

foss i I

m a r i ne

-1 -

* Fo s s i l g reen a l g a e ( m u l ti - ce l l e d ,
sex u a l reprod u ct i o n )

* Foss i l

bacte r i a

and

b l ue - g reen

a lgae

2-

F ree oxyg e n s u ffici e n t

res p i r a t i o n

for a n i m a l

F ree oxygen i nv a d es a t m osphere;

starts t o form oz o n e I oyer w h i c h
screen s
out
u l trav i o l et
rays
Wea th e r i n g of rock beg i n s

<

"'
0
......
0
""

_,

-<
u

oo

_, _

Q l

- ;j
.., _,

u:. o

<>< >
I>. W

* Ea r l i est known foss i l s ( s i n g l e

ce l l protists ( ba c t er i a ) a n d s i m p l e
p l a n ts ( b l ue- g reen a l g ae ) . Deve l
o p m e n t o f ce l l m e m b r a n e

F ree oxygen i n h ydrosph ere

creases; o x i d izes i ro n

-3-

Protei n and m a c ro m o l e c u l e s
A m i n o acids
S i m p l est c o m p o u n d s of
and N

Deve l o p m e n t of p h otosyn thesis; re

l eases free oxygen to hydrosphere
Fermentation
(?); adds
d i o x i d e to h ydrosphere

Co l l o i d a l CO!Jcervates

C, H , 0,

-4 -

in

c a r bon

Pri m i t i ve a t m osphere proba b l y

H 2 , H 2 0 , CH4 a n d N H 3 ; n o free
oxygen . I ntense u l traviolet radi
ation

FO RMAT I O N O F P L A N ET EARTH

Crit i c a l events i n t h e ea r l y h i story of l i v i n g m a tter a n d i n t h e deve l o p m e n t of

a t m o s p h e re and hydrosphere . denote foss i l s , the earl ier part of the h istory
being hypothetica l ( A fte r F l i n t and ot h e r s )

THE FOSS I L RECOR D i s the basis of o u r u n de rsta nd

i n g o f the h istory o f l ife a n d t h e course o f evol utio n .

Fossils a re t h e re m a i n s o f or i n d ications o f prehisto ric
a n i m a l s a n d p l a nts preserved i n the roc ks of the e a rt h ' s
crust. Fossi ls a re o f m a ny k i n d s a n d a r e fo rmed by
va rious p rocesses, but the c h a n ces of a n y o rg a n i s m be
co m i n g foss i l ized a re s m a l l . Thus, the fossil record is
a very i n c o m p l ete a n d rath e r b i a sed record of t h e
h istory of l ife . Recog n ition o f t h i s is i m po rta nt i n i nter
preti n g the fossil reco r d . Orga n i s m s l a c k i n g h a rd
pa rts, fo r exa m pl e, a re ra rely fou n d a s fossi l s . For t h is
reason records of the ea rly d evelo p m e n t of l ife a re
p a rtic u l a rly m e a g e r .
W H O L E A N I MALS A N D PLA NTS

are very ra rely preserved in t h e

fos s i l record . Wooll y m a m moths,
up to 1 0 feet ta l l , fou n d i n S i
ber ia a n d A l a s ka are e x a m ples
of such preservation by a deep
fre eze process.

O UTLI N E of soft parts of

org a n i s m s b u ried in fi n e m u d
i s s o m eti m e s preserved a s a
carbon fi l m , t h e m o re vo l a t i l e
com p o n e n ts d i s t i l led off b y h e a t
a n d pres s u re i n rock s . Exa m p les
a re t h e tri lobi tes and l eaves.

AN

Woo l l y m a m moth
preserved i n
froze n gro u n d

Ca m b r i a n
tri l o b i tes

Fo s s i l p l a n t
leaves fro m
Pe n n s y l va n i a n

Growth ri n g s a re preserved i n
s i l ica i n petrified wood.
This cla m s h e l l i s a l m ost u n a l
tered except f o r leach i n g .
MOST FOSS I LS consist of o n l y

the h a rd pa rts of a n i m a l s a n d
pla n ts , s u c h a s s h e l l s , bones,
teeth, and wood. In a few cases
these are a l most u na ltered, but
usua l ly they a re leached a n d
partly o r wholly replaced by
oth e r m i nerals, e s pecia l ly s i l ica
(Si0 2) and calcite (CaC0 3) . The
re p l a c i n g m i n e ra l s may some
ti mes preserve t h e orig i n a l m i
c rostru cture, a s i n s o m e s i l ici
fied wood, but this i s u n u su a l .

A m m o n ite, c a l ci u m carbo n a te ,
replaced b y pyri te ( FeS2 ) .

i m p res s i o n
or m o l d

IMPRESS I O N S A N D CASTS of
a n i m a l s a n d p l a n t s may be
formed in porous roc ks, s u c h as
san dstone, w h e n a l l the o ri g i n a l
com ponents a re d issolved away.
T h i s leaves a cavity, w h i c h may
later be ti l led by new m i nera l s ,
c a r r i e d i n sol u t i o n , to g ive o
cast of t h e orig i n a l o u t l i n e .
B U R R O W S , TRA I LS A N D TRACKS

may be preserved in s ed i m e n ts
t h a t a re l a ter co n so l i d a ted i n to
rocks.
STO N E ARTI FACTS a re the most

D i n o s a u r t r a c k s i n s a n d s to n e

c o m m o n re m a i n s of p re h i storic
man. They represent various
types of tools and wea pons.
P re h i s toric h a n d a x e

1 09

THE OLDEST FOSS I LS fou n d i n rocks est i m a te d t o b e

. a bo u t 2 . 7 b i l l io n y e a r s o l d , co n si st of si m pl e p l a n ts,
,
i n c l ud i n g l i m e-sec reti n g a l g ae, b a cteri a , a n d fu n g i .
Various o rg a n i c a m i no-acid res i d u es a re a lso k n own
frol?l these very a n cient rocks. Wel l - p reserved a n i m a l
fo is i l s fi rst a p pea r i n roc ks 6 0 0 m i l l i o n yea rs o l d .
Microsco p i c colo n i e s o f a l g a e
f r o m 1 .6 b i l l i o n years ago.
.Gu n f l i n t
Formation,
O n tario

x200)

Preca m b r i a n Fo s s i l s

O L D E S T PLANTS a re p re
served i n c h erts from North
A m erica, Africa, and A u s t ra l i a
a n d ra n g e i n a g e from 2 to
a b o u t 3 b i l l i o n years. They i n
c l u de fi l a m e n tous a n d s p h e r i c a l
al gae a n d bacte ria a n d ot h e r
m i c rosco p i c structures t h a t a re
n o t eas i l y classified. S o m e a re
c l osely s i m i l a r to l i v i n g for m s .
O t h e r m o r e w i d e s pread Preca m
bria n foss i l s i n c l u d e o p t i ca l l y
active orga n i c co m po u n d s o f
s u pposed org a n i c ori g i- n .
Stromatolites, w i d e ly d i stri b
uted in rocks of Preca m b r i a n
age, a re m ou n d l i ke, la m i n a ted
stru ctures, a few feet in d i a m
eter, fou n d i n calcareous roc ks.
They represent the deposits of
l i m e-secret i n g b l ue-g reen a l g a e.
THE

OLDEST
A N IMALS
a re
known from Ed iacara, South
Austra l i a , in Preca m br i a n rocks
that lie o n l y 500 feet b e l ow t h e
Ca m b r i a n . T h ey a re soft- bod ied
a n i mal s, i n c l u d i n g j e l lyfi s h , seg
m e nted wor m s, sea p ens, and
some a n i ma l s of u n k nown affi n i
t i es. I n con trast t o t h e oldest
pla nts, which a re pri m i t ive, t h e
o l d e s t a n i ma l s a re relatively ad
va nced type s, s u g g e s t i n g a l o n g
earl ier h istory.

THE

Seg m e n ted w o r m
fl o un d e ri
(Afte r Glaessner)
Spriggina

1 .5

in.

Jel lyfish
Medusian mawsoni

about 1 i n .

(Afte r Glaessner)

Ob olella, lower Ca m b r i a n
brach iopod A b o u t 0.2 i n .
.

0/ene//us, a lowe r C a m b r i a n
trilobite. Le ngth to 9 i n .

T h e " s u d d e n " a p p e a ra n ce o f fossi l a n i m a l s a b o ut

6 0 0 m i l l i o n yea rs a g o i s o n e of t h e m a jo r evo l utio n a ry
pro b l e m s . I t h a s been variously s u g g ested t h a t : ( a ) n o
Preca m b ri a n a n i m a l s existe d ; ( b ) Preca m b r i a n a n i m a l s
d i d exist but l a c k e d h a rd p a rts a n d were n ot fossi l i zed ;
( c ) Preca m b ri a n a n i m a l foss i l s have be e n d estroyed
by erosion a n d meta m o r p hi s m ; a n d ( d ) Preca m bria n
a n i m a l s were co n fi n e d to isolated, oxyg e n - ri c h a re a s
a n d a r e a s y e t u n d i s cove red o r u n exposed a s fos si l s .
N o n e o f t h e s e expl a n ati o n s i s n ecessarily u ntru e .
E a r l y C a m b ri a n d iversification o f a n i m a l s exte n d e d
over 3 0 m i l l i o n years, h e n ce w a s n o t rea l ly " su d d e n . "
I t seems prob a b l e that a n i m a l s d i d not orig i n a te i n
l ate Preca m b r i a n ti m es, t h a t t h e i r ea rli est representa
tives were soft- b o d i e d fo r m s of restricted d istri b ution ,
a n d t h a t t h e later w i d e s p r e a d a p p e a ra n ce o f h a rd
bod ied fo r m s i n C a m b r i a n ti m es m a y m a r k a res p o n s e
t o s o m e e n v i ro n m e ntal c h a n g e, s u c h a s t h e atmo
s p h e r i c co m position o r cut-off of u ltravi o l et ra d i ation
(p. 1 0 6 ) . I t w a s pro b a b l y r a p i d because of t h e rel a
tive " e m pti n e s s " o f m a n y enviro n m ents t o a n i m a l l ife,
a n d t h e stro n g sel ective pressure that t h e d evelo p m e n t
o f h a rd pa rts by a ny o n e g ro u p wou l d exert o n others .
111

M i.d d l e Ca m bri a n sea based on s peci m e n s f ro m B u rgess S h a l e of

Bri t i s h Co l u m bi a : ( 1 ) j e l l yfi s h , ( 2 ) s p o n g e , ( 3 ) trilobite, ( 4 )
worm , ( 5 ) brach iopod, ( 6 ) x e n o pod a r t h ropod

I NVERTEBRATES were t h e m ost disti n ctive

a n i m a l s of C a m b ri a n , Ordovi c i a n , a n d S i l u r i a n ti m es
a period of so m e 200 m i l lion yea r s . Althou g h verte
b rate fra g m e n ts a re fou n d in Ordovic i a n rocks, they
were ra re u n t i l D evo n i a n t i m e s . The ea rliest i n ve rte
b rates ( p . 1 1 0) i n cl u d e d jel lyfish, sea pens, a n d seg
m e nted wo r m s , b u t C a m b r i a n fa u n as were d o m i n ated
by trilob ites, riow extinct a rth ropods . Spong es, s n a ils,
e c h i n o d e r m s, and s m a l l horny b ivalved brach iopods
were a b u n d a n t i n s h a l low seas. In the Ordovician,
cora l s , b ryozo a n s ( m oss a n i m a l s ) , a n d m a n y n ew
k i n d s of b ra c h i o p o d s a n d t ri l o bites a p pe a re d . P ro
tozoa n s we re ra re. S q u i d l i k e cepha lopods, so m e 1 5
feet l o n g , d eveloped . I n t h e S i l u ri a n , e u rypte r i d s, a r
t h ropods to 6 feet l o n g , l ived i n del ta s a n d estua ries.
MAR I N E

1 12

Represe n t a tives of a l l t h e m a jo r l ivi n g i n verte

brate p h y l a a n d n ea rly a l l the c l a sses were esta b l i shed
by Ordovi ci a n ti m e s . S i n ce t h e n , the m a j o r patter n s of
inverteb rate l ife i n the seas h ave c h a n g e d l ittl e . A few
m a jor g roups h ave becom e exti nct, d iffe re nt g eo
g ra p h i c a r e a s a n d d iffere nt e nviro n m e nts h ave sup
ported d iffere n t fa u n a s, and g e n e ra and spec ies h a ve
shown va ried p a tterns of mod ificatio n a n d exti n ctio n .
EARLY

PALEOZO I C

A N IMALS

s h owed many a d a ptations to

d ifferi n g modes of l ife. They i n
c l uded fixed benthic for m s , such
as cora l s a n d brac h iopod s ; va
g rant benth i c types, s u c h as
starfi s h and s n a i l s ; free-swi m
m i ng for m s , s u c h a s ce p h a l o
p o d s a n d e u rypterids; a n d free
fl oating forms, such a s j e l lyfi s h .
Comparable d ivers ity existed
in feed i n g h a b its. The p hyto
plan kton , on w h i c h m a ny l i v i n g
marine i nve rte b rates f e e d , h ave
s i l iceous a n d calcareous hard
parts. These types a re u n k n ow n

i n the Ea rly Pa leozoic, perhaps

beca use their foreru n n e rs were
soft- bodi ed. T h e h a rd parts of
Early Pa leozoic i nve rteb rates a re
com posed of various m i nerals.
Ca m brian for m s consist c h i efly
of phosphatic, s i l iceou s , a n d
ch iti nous m a te ria l s , but calci u m
carbonate beca m e t h e pred o m i
n a n t s h e l l s u bsta nce i n O rdov i
c i a n t i m es . Little i s y e t k n own
of the s i g n ifica nce of this b io
c h e m ical
evolution.
O r i g i na l
s h e l l com position i s ofte n mod i
fled b y s u bseq u e n t alteration
duri n g fos s i lization ( p. 1 0 8 ) .

A Devo n i a n cora l reef : ( 1 ) trilobite, (2) cephalo pod, (3) bryozoan,

(4) brac h i opod , (5) cora l , (6) cora l , (7) cora l
------

THE O LDEST VERT EBRATES a re fra g me nts of a rm o re d

fi s h fou n d i n roc k s of Ordovici a n a g e i n Wyo m i n g a n d

el sewhere. Fish re m a i n ra re a s fossils unti l l ate S i l ur
i a n ti m e s . They b e co m e d iversifi e d a n d a b u n d a nt d u r
i n g t h e Devo n ia n .
The o r i g i n o f ve rtebrates i s obscure. Th ey belong
to th e Phyl u m C h o r d ata, conta i n i n g so m e m e m b e rs
t h a t l a c k a vertebral col u m n ( a corn wo rms, sea s q u i rts,
l a n celets, a n d t h e i r k i n ) tho u g h t h ey do h ave a sup
porti n g n otochord and other featu res s h a red with the
" h i g h e r " vertebrates . Larva l acorn wor m s show stri k
i n g s i m i l a rities to l a rva l ech i n o d e r m s, sugg esti n g that
both g roups may h ave a rise n fro m a com mo n b u t un
known a n cestra l stock.
primitive
g roup of fi s h , a re represente d
today o n ly b y t h e h a g fi s h a n d
l a m preys. Agnatha l a c k the true
jaws a n d paired fins typical of
most l i v i n g fish.
Ma ny of these oldest a n d
most d iversified o f the early fi s h
had a bony arm or. Cal led ostra
coderms (" bony s k i n "), they

AG NATHA, t h e most

ra rely exceeded a foot i n

l e n g t h . T h ey l ived ch iefly in
stre a m s and estu a ries w h ere
pre s u m a bly they fed on botto m
m ud s or on suspended mate ria l .
They a re n ot k n own i n roc ks
you n g e r than the D evon ian, per
h a p s beca u se th ey were soft
bod ied-l ike their l iv i n g repre
sen tatives.
Pteraspis

U p p e r S i l u r i a n to De
von i a n . About 6 i n .

B irkenia

A n S i l u ri a n a g n a t h a n
f i s h a bo u t 4 i n . l o n g .

Drep a n aspis

1 14

lower Devon i a n .

T o 1 ft.

Climatius, U pper S i l u r i a n to De
vonian, was a spiny a c a nth o d i a n
"shark" with rhomboid s c a les, 2
spines on bac k , a n d 5 pa i rs of
ve ntra l fins. L e n g t h 3 i n .

w a s a j o i n ted
necked m a r i n e a rt h rod ire to 30
ft. lon g . It was the largest verte
brate of Devon i a n l i m es.
D u n k l e os t e u s

( late S i l urian to
Per m i a n ) are the o n ly verte brate
class to have beco m e extinct.
They reached their peak i n the
Devo n i a n a n d a re rare in
you nger Pa l eozoi c ro'cks. Placo
derms d i ffer fro m Agnatha i n
havi ng pa i re d fi n s a n d pri m i tive
jaws, i m porta n t fea t u res i n
later d iversification o f verte
brate s .
bot h
i n cluded
Placoder m s
freshwater a n d m a r i n e form s,
such as t h e 30-fool, joi nted
necked arlhrod ires; s m a l l fresh
water s p i ny a c a n t h o d i a n s ; a n d
mass ively a rmored , strong-finned
a n t ia rch s .
PLACODERM$

shark jaws
and teeth

a s h a rk f r o m the
U p per Devonian, with a stream
li ned naked body. To 4 ft.

Clacloselache,

Teeth of C arch a r o clo n, a

40- to 50-foot shark from
the Mi o cene
S H A R K S A N D RAYS belong to

the C h o n d richthyes, a class of

predaceous, cart i la g i nous, open
g i l led fish. Sharks s h ow m a ny
ada ptations to life i n t h e open
ocea n s , i n c l u d i n g strea m l i n i n g ,
wel l - developed teet h , a n d spiny
skin scales. S o m e l i v i n g s harks
rea c h a l e n g t h of 50 feet. The
earliest m e m bers of t h e g roup,
which ap peared in Devo n i a n
t i m es , l ived i n f r e s h water.

Skates and rays, fl a tte n ed for

botto m - d we l l i n g existe n c e , h ave
fl a tte n e d teeth for cru s h i n g
s h e l ls.
Loss of t h e bony a r mor, de
velop m e n t of efficient jaw sus
pension, and more flexible fins
pro v ided both s h arks and bony
fi s h with an advanta g e over
t h e i r placod e r m a n cestors. I so
lated teeth a n d spines a re the
m ost co m mo n fos s i ls.

1 15

Rece n t

EVO L U T I O N O F F I S H

Te rti a ry
Bon y R a y - Fi n n e d Fi s h

Cretaceo u s )
J u ra s s i c

L u n g fi s h

Triassic
Perm i a n
P l acoder m s

:::::::1, =
Ord ov ici a n

Ag n a t h a

Crosso p teryg 1 a 1

Carti l a g i n o u s F i s h

....... . . . . . . ..

- -

- - - - -

---

'

...... . . .

BONY F I S H (Oste i c h t hyes) i n

c l u d e nearly a l l l i vi n g fres h
water a n d m a r i n e species. They
have stro n g , but flex i b le, bony
s ke l eto n s and either sca les or
plates. Most k i n ds have a n a i r
bla dder. Bony f i s h l i ve i n every
k i n d of a q u a t i c e n v i ro n m e n t
(eve n caves), a n d t h e y outn u m
be r a l l other vertebrates co m
bi ned, both i n n u m bers of spe
cies a n d of i n d iv i d ua l s . The

oldest m e m bers w e r e fres hwater

for m s from the M i d d l e Devo
n i a n . T h ey i n c l u d e f i s h with two
types of f i n s .
The ray-f i n ned f i s h were a
rare, f reshwater g roup i n the
Pal eozo ic, b u t t h ey beca m e the
do m i n a n t g r o u p i n t h e Mesozoi c
a n d Cenozo i c . T h e i r scales be
ca m e t h i n n e r, a n d t h e i r jaws
and s keleto n s s h owed progres
sive i m prove m e n t.

a Mi d d le Devo n i a n
fi n ned fish. L e n g t h a b o ut 1 1 i n .

ray

Cheirolepis,

I ,
,,

Deta i l of ray fi n , w i t h
typica l s u pporti n g b o n e s .

1 16

EVO L U T I O N O F AM P H I B I A N S

U ro d e l e s
Ste reos po n d y l e

To Repti les

..

... .

L a b y r i n t h o d o n ts

). .l e-f)o fpQ" d y I e

. . . . . . . . . . . . . . . . . . .. . . .

Am p h i bi a n s

A I R-BREAT H I N G

BONY

FISH

( C h oa n i c h thyes), a s m a l l e r g rou p
t h a n t h e ray-fi n n ed fi s h , have
i n te r n a l n os tr i l s that o p e n i n to
t h e m o u t h , a s do t h ose i n l a n d
l iv i n g vert e b r a t e s . L i vi n g fo r m s
i n c l u d e t h e l u n g fi s h ; t h ree g e n

era a re k n own, o n e fro m each

of t h e s o u t h e r n co n t i n e n t s . T h e y
have p owe rf u l fi n s , s u p ported
n ot b y a fan of s l e n d e r bones
as i n t h e ray-fi n n e d fish but by

a strong bony a x i s . T h e y u s e
t h e s e s t o u t fi n s to " wa l k " f r o m

p o o l t o p o o l d u r i n g t h e d ry sea
son.
Lobef i n s, t h e othe r m a j o r
g r o u p , i n c l u d es t h e l i v i n g m a
r i n e coelaca n t h s a n d t h e i r more
g e n era l i zed, freshwater, ca r n i v
o rous, Devo n i a n crossopteryg i a n
forebea rs. I t wa s t h ese t h a t g a ve
rise to the terrest r i a l verte b rates
( p p . 94-95).

Oste ole pis, from M i d d le Devo

nian, with t h ick, rhomboid sca les
a n d short, lobed fi n s. To 9 i n .

fr i n ge

Lobe fi n , show i n g t h e
strong s u p porting bo nes
from wh ich feet deve loped.

1 17

c o mp a r a ti v e l y late de
ve l o p m e n t . life p ro b a b l y o ri g i n ated in the s h a l l ow
seas, where t h e m a jo rity of i n vertebrate g ro u p s a re
sti l l restri cte d . Life o n the l a n d i nvolved m ajor c h a n g es
fo r these c reatu res that ori g i n ated a n d lived i n t h e
ocea n s . T h e m o d ificatio n s i n cl u d e d c h a n ges n ecess a ry
fo r p rotection a g a i n st d ryi n g u p , n ew m ethods of s u p
port i n a i r a s o p posed t o the m o re buoya nt water,
b reat h i n g oxyg en as o p posed to extra cti n g it from the
water , n ew sou rces of food a n d water, a n d n ew repro
d u ctive m e c h a n i s m s to assure ferti lization i n the ab
sence of water. Colon ization of rivers a n d l a kes was
only slig htly less for m i d a ble, for it involved d evelop
ment of m ec h a n is m s to p reve nt d i l utio n of body fl u id s

L I F E ON TH E LA N D w a s a

A RTH R O PODS h a v e exceeded

all o t h e r g rou ps in the d i ver
sity and n u m be r o f t h e i r terres
tri a l and flyi n g re p r e s e n ta t i ve s .
T h ey g a i n e d a " f l y i n g start"
b y t h e i r tou g h , f l e x i b l e outer
coveri n g and by t h e i r s t ro n g
a p p e n d a g e s . T h e o l d est l a n d
a rt h ro pods a re l a te S i l u r i a n

m i l l i pedel i ke f or m s t h a t m a y
h a ve b e e n p a r t l y a q u a t i c . I n
sects f i rst a p peared i n t h e D e
von i a n . By C a r bo n i ferous t i m e s ,
a va r i ety o f a rt h ro p o d s , i n c l u d
i n g p r i m i t ive w i n g ed i n sects,
coc k roa c h e s , s p i d e rs , and scor
pions,
had
a p pea red .
Most
g ro u p s a rose in t h e Mesozo i c .

A RT H ROPODS

that, in a l l a n i m a ls, conta i n d issolved salts p re cisely

adjusted to the osmotic balance of sea water.
land dwe l l i n g , i n spite of its problems, offered all
the a dva ntag es of an em pty envi ro n m ent. Beca use of
the deli cate i nterdependence of a l l livi n g t h i n g s, it is
not surprisi n g that both pla nts and a n i m a l s seem to
have colo n i zed t h e l a n d at about the s a m e ti m e d u ring
the Silu ria n a n d Devo n ia n . Th e i nvasion of the l a n d
a l m ost certa i n l y i nvolved t h e earlier i nvasion o f fresh
waters. Ma ny l iving g rou ps, wh ich a re essentially m a
rine, conta i n a few freshwate r colon ists (cla m s a n d
crustacea ns, fo r exa m ple), but o n ly t h e p l a nts a n d
three m a j o r g ro u p s o f a n i m a l s (snails, a rthropods, a n d
vertebrates) h ave beco m e fully established o n the l a n d .

h a v e established
them selves o n the land with
varying degrees of s u cces s . Most
a m ph i b i a n s a re l i m ited to a reas
near e nough to water to a l low
them to return to it to repro
d uce. Most reptiles a re restricted
to a reas from the tropical to the
te m perate zones. Ma m ma l s and
bird s a re more widely d i stri b
uted a n d ada pted. Some verte
brates, i n c l u d i n g turtles a n d
other ext i n ct rept i l e s , porpoises,
wha l es, and pe n g u i n s , have u n
dergone a secondary adaptation
to m a ri n e l ife (p. 1 2 8).

VERTEBRATE S

S N A I LS
have i nvaded fresh
waters a n d t h e l a n d . Some
h ave reta i ned the protective
s h e l l , but ot h ers (slugs) a re
na ked . Land for m s m ove a n d
feed b y b rows i n g , m u c h l i ke
aquatic for m s. T h ey have de
veloped lungs for brea t h i n g .

Skeleton o f Perm i a n a m p h i b i a n ,
Eryops. L e n g t h a b o u t 5 feet.

La nd Snail, Helix

1 19

Cycads

600 Psilophytes

terids
S p h e nophylls

g re e n a l g ae,
which now exist i n both the seas and i n fresh waters.
Like a n i m a ls, d ifferen t g roups of pla nts s how va ryi n g
deg rees o f a d a ptatio n t o l a n d life.

LAND

PLANTS p rob a b l y a rose fro m

BRYO PHYTES ( m osses a n d l iver

worts ) need w a ter in repro d u c
tion and f o r protection from
desiccation . T h e i r parti a l a d a p
tation to land life is a n a logous
t o t h a t of t h e a m p h i b i a n s .
S m a l l p l a n ts, w i t h leaves a n d
stems, they l a c k woody t i s s u e s
for s u p port a n d c i rc u l a t io n .

1 20

' '

w h i c h incl udes
the a lgae, f u n g i , a n d bacteria,
lack t h e roots, stems, leaves,
and va s c u l a r s u p port i n g and c i r
cu l a t i n g syste m typ ical of h ig h
e r p l a nts. T h ey a re e i t h e r u n i
cel l u la r or consist of loosely or
g a n ized g roups of c e l l s . li m i ted
to d a m p e n vi ro n m e n ts .

THALLOPHYTES,

A n g iosperms

G i n kgo

Corda ites

C o n ifers

VASCULAR PLANTS ( Tra c h e o p h ytes ) i n c l u de t h e m a

jority o f l ivi n g p l a nts. A l l have specia l i z ed vasc u l a r sys

te m s of con d ucti n g tissues that tra n s port water a n d
n utrie n ts fro m t h e s o i l t h rou g h t h e roots t o t h e other
pa rts of the p l a nt. T h i s syste m a l so provi d e s s u p po rt,
a l l owi n g so m e of these p l a n ts to g row to g reat sizes.
They als o h ave a n outer l ayer (cuticle) that p revents
d esiccation . The e a r l i est vascu l a r p l a nts were seed l ess
kinds, such a s those shown i n Devo n i a n forest b elow.
Devo n i a n f o r e s t sce ne. S h o w n a re : (1) a prim itive lycopod (P ro to
lepidode n dro n ), (2) tree fe r n (Eosperm atopte ris), a n d (3) s c o u r i n g
r u s h (Cala m op hyton).
g ro u n d .

Psi l opsids a re

low g rowi ng pla nts

in fore

PSILOPSI DS, wh ich include the

e a r l i e s t k n own vasc u l a r pla nts,

lack roots . Th e y h ave either
pri m itive l eaves or are leafless.
T h o u g h w ide s p rea d i n Devo n i a n
times, they re m a i n e d s mal l i n
s ize. O n ly two g e nera survive.

include the liv

i ng scou r i n g rus h e s a n d s i m i l a r
Paleozoic p l a nts that g rew to
40 feet ta l l . T h ey h ave roots
and long, seg m e n ted, ribbed,
c o ne be ari n g stems with circlets
of l e av es at the nodes.
S P H E N O PS I D S

S E E DL E S S VAS C U L A R PLANTS i n cl u d e psilopsids, lyco

pods, ferns, a n d sp henops i d s . The a d u l t p l a n t prod u ces

spores that d evelop i n to s m a l l speci a l i z e d l eafless
p l a nts ( g a m etop h ytes ) . T h ese l ater p r o d u ce g a m etes,
or sex ce l l s . Because sperm req u i re wa ter to re a c h t h e
eggs, these seedl ess pla nts a re restricted to d a m p
envi ro n m ents . Widespread i n t h e P a l eozoic, they de
clined a s seed-bearing p l a nts expa n d e d in Mesozoic.
i nclude the living
club mosses and g ia n t represen
tatives fro m t h e Pe n n sylva n i a n
coa l forests .
LYCOPODS

1 22

which sti l l survive in

la rge n u m bers, a re spore- bear
ing p l a n ts. Some foss i l a n d l iv
i n g for m s g rew to 50 feet ta l l .

FER N S,

SEED-BEARING PLANTS a re of two ba sic kinds: non

flowerin g a n d floweri n g . I n the non-flowe ri n g g roups
(gym nosper m s ) , t h e seed i s n ot p rotected ; it i s " n a ked"
-as i n pine cones. In fl oweri n g pla nts ( a n g iosperms),
the seeds a re p rotected. In both ki n d s , resista nt pol len
a n d e g g s a re p ro d u ced d i rectly from pa rent p l a nts.
Pollen fertil izes the egg, wh ich d evelops i n to a seed
wh ich is p rotected from d ryi n g . As a result, seed-bear
i n g pla nts h ave colonized a g reat variety of l a nd a reas
and a re the d o m i n a nt l ivin g g roup of pla nts.
GYM N O S PERMS i n cl u d e ( 1 ) e x
t i n ct seed ferns, perhaps a n ces
tral to other g ro u ps; ( 2 ) cycads

a n d their exti n ct relatives, which

were a b u n da n t i n the Mesozoic;
(3) exti nct cordaites, perha ps
a ncestra l to conife rs, (4) the
living ginkgos; and (5) the
widespread, a b u n d a nt con ifers.
FLOWER I N G
PLANTS
( a ngio
sperms) a re re pre sented today
by over 250,000 species. They
appeared in the Mesozoic a n d
ra p i d l y d is p l a ced t h e gym no
sperms, which we re then d o m i
na nt. Their flowe rs a re repro
d uctive structu res, m a ny of t h e m
speci a l ly developed t o a ttract
i n sects that ca rry the m a le pol
len to ferti lize t h e fe m a le
fl owers . Enclosure of the seed
i n a protective covering a l so
rep resents an adva nce over the
gym nospe r m s . Floweri n g pla nts
s how n u merous a d a ptations to
d iffere n t e n v i ro n m ents, ra n g i n g
fro m desert c a c t i to tropical
swa m p trees a n d f l owers.
C h a n g e s i n s o m e a n i m a l g ro u ps
a ppear related to c h a nges i n
vegeta ti o n ( p . 5 1 a n d 1 0 1 ).

Early
cycad

F lowe ring Pla nts

of the C retaceous

A M P H I B I A N S were t h e fi rst te r r e s tr i a l

v e r te b r a tes ,

b u t they a re o n ly p a rtly a d a pted to life o n l a n d . T h ey

n eed to retu rn to water to lay t h e i r eggs, a n d t h e i r
you n g d evelop i n water. Most k i n d s a re co n fi n e d to
d a m p envi ro n m ents as ad ults.
The o l d est a m p h i b i a n s, the i c h t hyosteg i d s fro m the
U p pe r D evo n i a n , a rose from the crosso pte ryg i a n lobe
fi n n e d fi sh , possi bly i n respo nse to popul ation pressu re
in the pool s where the latter l ived ( p . 9 5 ) . The stout
bony axis a n d m uscles of the fi n s a n d the p rese n ce of
l u n g s a d a pted lobefl ns idea l l y for m ig ration from sta g
n a nt a n d seaso nal p o n d s . Life o n the l a n d p rovided
u n l i m ited oxyge n suppl ies, the poss ib il ity of a d d itio n a l
food sou rces, escape fro m pred a tors, a nd t h e m e a n s of
reaching oth er bod ies of water.
LATE PALEOZO I C AMPH I B I A N S

showed g reat d ivers i ty. Their

adaptive rad iation onto land
was ra pid, a n d some for m s u n
derwent a secondary return to
: h e w a ter. Some l a byri n t h o
d o n ts were 1 5 f e e t i n l e n g t h .
A m p h i b i a n s d o m i n a ted t h e l a n d

f o r over 1 0 0 m i l l i o n years.
They d ec l i n e d in t h e early
Mesozoic, per h a p s a s a res u l t
o f competition w i t h t h e i r better
a d a pted reptili a n desce n d a n ts .
i n c l ude
a m phi bia n s
Livi n g
n ewts,
s a l a m a n d ers,
frog s ,
toa d s , a n d caeci l i a n s .

Late Pa l eozo i c coo l -for m i n g swa m p w i t h la byri n t h od o n t a m ph i b i a n s .

Early re ptiles d i ffered f ro m t h e i r a m p h i b i a n a n cestors i n o n l y m i

n or ways. They u n d e rwe n t ra p i d d iversification i n Perm i a n ti mes.

THE RISE OF T H E REPTI LES m a rked a new stage of

a d a ptation to l ife on l a n d . Repti les d evelop fro m a n

eg g with a tou g h outer coveri n g , provi d i n g a b u i lt- i n
food s u p p l y a n d a sea l e d , l i q u i d -fi l l e d c a p s u l e for t h e
deve l o p i n g e m b ryo . The i nfa nt repti les e m erge fro m
the eg g m o re o r less fu l l y for m e d . R e p ti l e s were t h u s
a b l e t o c o l o n i ze the l a n d a reas fa r removed fro m
strea m s a n d l a kes . Repti l i a n s k i n is scaly o r c o r n i fi e d ,
a p rotection a g a i nst d ryi n g u p ; the l i m b s a n d c i rc u l a
tory system s o f reptiles a re g e n e ra l ly s u perior to those
of a m p h ib i a n s . Reptiles u n d e rwe nt g re a t d ive rs ifica
tion i n Mesozoic t i m es, d o m i n ati n g l ife not only on
the land but a lso i n th e seas and i n t h e air. Th e i r de
c l in e, sti l l not fully u n d e rstood, was m a rked by t h e ex
p a n si o n of t h e i r d esce n d a n ts, the birds a n d m a m m a l s .
1 25

R h y ncoceph a l i a n s

Turtles
.-

1ity <

,,.,__ _ ,

',

.rd
Plesiosaurs

Ichthyosaurs

J U RA S S I C

AQUATIC R E PT I LE S we re a b u n

d a n t i n the Mesozoic, as rep

tiles m astered every major e n
v i ro n m e n t . Some were fi s h l i ke,
oth ers
rese m bled
the
later
sea l s , and s ti l l oth ers were ser
pentlike.

1 26

R E PTI LES
had l i g h t ,
stro n g s kel eton a n d w i n g s , s u p
ported by a n e l o n g a ted fi n g er.
So m e were s m a l l ; oth ers had
2 0 -foot wing s p a n s . T h ey were
co n te m pora ries, b u t n o t a n ces
tors, of early bird s .
FLY I N G

lizards

Birds

Crocodiles

::::... Sauropods

d o m i nated l a n d
D I N O SA U R S
l ife for t h e 1 40 m i l l i o n years o f
the Mesozoic. Aris i n g from the
codont a n cestors, t h ey i n c luded
two g roups with d istinct h i p
structures : rept i l e - l i k e sauris
chians and b i rd l i ke ornithis
chians. Worldwide i n d i s tribu-

tion, t h ey were ada pted to

m a ny d iffere n t enviro n m e n ts.
They included herbivores a n d
carn ivores a n d a l s o t h e l a rg est,
most heavily a rmored l a n d a n i
m a l s t h a t h a ve ever l ived.
Reasons for their extin ction in
the late Mesozoic are obscure.

1 27

ADAPTIVE R A D I AT I O N of repti les i n to fo rms a d a pted

to l ife i n d ifferent envi ro n m ents was p a r a l l e l e d by

birds a n d m a m m a l s afte r the repti l es beca m e exti n ct.
Adaptive rad iation occurs i n the early h istory of m a ny
g roups, u s u a l ly fol lowed by m o re special ized a d a pta
tions to niches with i n the wirler e nviro n m e nts .
OCEA N S

FRESHWATER

EVO LUTI O N ARY C O N V E RG E N C E

i n form betwee n g e n etica l l y u n

re l a ted pe n g ui n s , d o l p h i n s , ich
thyos a urs and s h arks res ults
fro m a d a ptation to s i m i l a r e n
viro n m e n ta l co n d i t i o n s . I t i s
a l so pres e n t i n m a n y other

1 28

-<

LAN D

g ro ups. A l t h o u g h e a c h of t h e
mammalian
tetra pod
c l a s ses
repre s e n ts a n e w or d i s ti n ctive
ado ption to life in various e n
viron m e n ts , t h e t h ree " h i g h es t "
c l a s s e s h av e each s u ccess f u l l y
a d a pted to a l l e n v i ro n m e n t s .

C r o n e s, Rails,
a nd A l l ies

['"' "/1
4-

-'' '"''

Loo n s

- I't
.{;<!-

Grebes
Ralites

Pe r c h i n g B irds

Climbing Birds

o n d R o l l e rs

B I R DS a re ra re a s fossils, beca use of t h e i r fra g i l e

ske letons a n d because m a ny b i rd s l ive i n a re a s where
b u r i a l cond ition s that l e a d to p reservation a re u n co m
m on . B i rd s s h a re t h e e g g - layi n g c h a racteristics o f rep
tiles from wh ich they a rose, but the i m po rta nt, d isti n c
tive featu res i n t h e i r d iversification a n d su rviva l a re
t h e i r s u p e rb a d a ptation to flyi n g , t h e i r ca re of the
you n g , t h e i r fea t h e r cove r i n g , and their wa rm - blooded
n ess . The histo ry of so m e i m po rta n t g ro u p s of b i rds is
s hown a b ove .

1 29

EVOLUTION OF MAMMALS (T r i a s s i c to Rece n t) from

t h e m a m m a l - l i ke therapsid repti l e s ( p . 1 24) i s well doc

u m e nted i n the fossil record . S o m e fossi l s a re so

tran sitio n a l i n c h a racter between the two g roups that
there is doubt w h i c h they rep resent. Ma m ma l s a re
typ i ca l ly cove red with h a i r o r fur, h ave d ifferenti ated
teeth, a re wa r m - blooded, and h ave h i g hly d evel o p e d
senses. Nea rly a l l m a m m a l s give b i rt h t o t h e i r you n g
o n l y a fter a l o n g period o f protective e m b ryon i c d e
velo p m en t with i n the moth e r ' s body, a n d then the
mother fee 9 s the you n g m i l k secreted fro m her m a m
m a ry g l a n d s . These featu res a n d t h e h i g h l y d eveloped
b r a i n s of m ost m a m m a l s m ust h ave been of m a j o r i m
porta n ce i n t h e evol utio n a ry success o f t h e g ro u p .
Ma m m a l s ' reg ul ated b o d y tem pe rature e n a b l e them to
survive i n a m u c h g reater e nvi ro n m ental ran g e th a n
d i d the re pti l e s .
THE

OLDEST

MAMMALS

were

s h rew-sized c reatures, a few of

which reached about one foot
in length. They a re knon
from t h e i r tiny foss i l bones.
These m a m m a l s re m a i n ed i n
conspicuous
t h roughout
the
Mesozoic.
A n cestra l fos s i l m a m m a l

Ec h i d n a

MO N O TR E M E S , l i ke the echidna
(s piny a n teater) and platypus
(p. 53), lay eggs and secrete
sweat
milk
from
modified
g lands. T h e i r pri m itive repti l i a n
c h a racteristics s uggest t h a t t h e y
are a n a n c i e n t g roup, a n d they
a re l i m ited to t h e Austra l ia
area.

Pla typus

1 30

n u rs i n g you n g

SOUTH A M E R I CA

7, -- -rf

'" . ....\'1

, .

t.

NO RTH AM E R I CA

!'

Mars u p i a l Carnivore

%: .

r;r'
t:

Ca m e l - l i ke L i toptern

'(t:(
.. \

Horse - l i k e Litoptern

Toxod o n t

j
.

i1'
"'

H o m a l o d o t h ere

MARS UPIAL MAMMALS (Creta

ceous to Recent), s u c h as kan

garoos and opossu m s , g ive b i rt h
to very i m mature you ng that
a re s h e ltered and fed i n s i d e
t h e i r mother's pou c h . Ma rs u pials
were
widespread
in
South
A m e rica i n the Ce nozoic, s h ow
i n g many exa m p l e s of conver
g e n ce (p. 1 28 ) w i t h placental
m a m m a l s a s i l l u strated a bove .
T h e j o i n i n g of N o rth a n d
P LAC E N TAL MAMMALS i n c l ude

m ost l iving m a m m a l s . Because

t h e e m b ryo is nourished a n d
susta i ned b y the p l a c e n ta with i n
t h e m o t h e r ' s wo m b, t h e n ew-

South A m e rica by t h e e m er
g e n c e of the I st h m u s of Pa n a m a
i n t h e l a t e Cenozoic e n d e d t h e
isolation i n w h i c h t h e s e m a rsu
pials h a d developed. Com peti
tion with the better a d a pted
North A merica n placentals re
s u lted in the extinction of m ost
m a rs u p ials. Ma ny have s u rvived
in Austra l i a beca use of t h a t
con t i n e n t ' s c o n ti n ui n g i s o l a t i o n .
( After S i m p s o n )
born a re m ore m a t ure t h a n a re
m a rsupials, pres u m a b l y a n i m
po rta nt evol utionary adva ntage.
The o l d est ( Cretaceous ) were
s h re w l i k e i n sectivore s .

131

EARLY CENOZO I C MAMMALS s h owe d a period of ex

plos ive rad iation, re p l a c i n g the Mesozoic ru l i n g reptiles
i n a l most eve ry e nviro n m e nt. Reco n structio n a bove of
scene some 50 m il l io n years ago i n c l u d e d a ncestra l le
m u r Notharctos ( 1 ) , carn ivo res Oxyaen a ( 2 ) , Mesonyx
( 3 ) , hoofed m a m m a l s Palaeosyops (4), a tita no
there (5 ) , and the a m blyods Eobasileus (6), Uinta
therium (7), a n d Coryphodon ( 8 ) .
Phe n a codus

ARCHAl C HERBIVOROUS,
H O O F E D MAMMALS
i n cl u ded

Phen acodus, an adva nced con

d y l a r t h , with a l o n g ta i l , fi ve
toes, a n d a carn ivore l i k e s k u l l .
Co n te m pora ries were a m b l y
pods a n d u i n ta t h e re s ( a bove ) ,
w i t h teeth m o d i fi ed for chew
ing v e g e t a tio n . T h e clawed
toes of a n ces tra l fo r m s later
beca m e m o d i fi e d to h ooves.

EVOLUTION OF C A R N I VO R E S (After Colbert)

UJ
J z
UJ
u u

J
)
'
J
)

o
,....

CL

u
'

u w

5u u

L. 0
UJ

ARC H A I C C A R N I V O R O U S MAM
M A L S - th e creodo n ts - were

mostly s m a l l , s l e n d e r, long
ta i l ed creatures. T h ey deve l
o p e d c laws, s h a rp teet h , a n d
s u p p l e l i m bs. So m e rea c h e d the
s i ze of l io n s . Most c reodonts
beca m e e xt i nct in the Eoce n e .
F r o m wease l - l i ke m e m bers of
t h i s g ro u p t h e re s u bs e q u e ntly
developed t h e a n cestors of l iv
ing cats, dogs, a n d bears.

i n c l uded
LATER C A R N IVO R E S
a n ces tra l for m s of fl s s i pe d (s p l it
foote d) cats, dog s , hye n a s , a n d
weasels, a l l o f w h i c h a pp e a red
a t d iffere n t t i m e s . We b - footed
carn ivores (sea l s , wa lru ses) i n
vaded t h e ocea n s i n Miocene
t i mes. T h e c l osely re lated ceta
cea n s , i n c l u d i n g d o l p h i n s a n d
whales, a p peared i n t h e Eoce n e
a n d a re s u perbly a d a pted to
m a r i n e l ife.

1 33

M O D E R N M A M M A L S a rose i n Eocene a n d O l i g ocene

ti m es. P l eistocene recon struction s h ows g i a nt g round

sloth Megatherium ( 1 ) , b ison (2}, s a b e r-tooth cat ( 3 ) ,
horses (4), wool l y m a m m oth ( 5 ) , c a m e l - l i ke C a m elops
(6), g lyptodo n t (7}, a h u g e beaver Castoroides ( 8 ), a n d
g ro u n d sloth Mylodon ( 9 ) .
MODERN

H O O FE D

MAMMALS

( U n g u la tes ) i n c l u d e od d - toed
horses, tapirs , a n d rh i n os , a n d
eve n - toed, clove n - h oofed cat-

tie, pigs, ca m e l s , and d eer,

which d i s p l aced the earlier
o d d - toed u n g u lates. C h a n ges
i n veg etation h a d a n ef fect .
Cottle

Camels

.>.''

P r i m i tive
Ruminant

(t .

1 34

f:f(v

;i...y;
\; C h evroto i n s
FAM I LY T R E E O F
EVEN-TOED U N G U LATES

P r i m a tes

I nsectivo re s

D e r m o pterans

,t!jf>.'

.>.
. t'

C h i ropterans

Pyrotheres

Adaptive rad i a ti o n of place n t a l m a m m a l s .

( After Col bert. )

Ra d ia t i o n of m a m m a l s i nto every e nvi ro n m e n t

is typified by t h ei r m a stery o f t h e a i r a n d t h e ocea n s
as wel l a s t h e l a n d . Bats a n d a n cestral w h ales both a p
pea red i n the ea rly Tertiary. On t h e l a n d , specia l i z e d
m a m m a l i a n g ro u p s d eveloped . Rodents a n d rabbits
a d a pted to a variety of food s a n d ways of l ife, i n cl u d
i n g bu rrowi n g . Prim ates, m a ny a d a pted t o l ife i n t h e
trees, a rose ea rly i n t h e Tertia ry. Elepha nts a n d e d e n
totes (sloths a n d a r m a d i l los) represent fu rther special
ization i n a d a ptati o n s .

135

EVI DE N CE OF EVO LUT I O N of o n e species i n to a n

o t h e r over geo l o g i c t i m e i s p rovi d e d by m a n y m a m

m a l i a n g rou p s . Two typica l exa m p l es a re g iven h e re .
TITA N OTHERES were a g rou p
o f large Te rtiary m a m m a l s . Their
evo l u t i o n a ry
deve l o p m e n t
is
shown i n h i storical seq u e n ce

below. Other for m s a l so e x

i sted, a n d t h e i r evo l u t i o n i s a l so
we l l documented. (After Os
born.)

B rontotherium

Ill
c
Ill

platyceras

4;

_g

_,.,

*"

Brontotherium
leidyi

cr.;.;
.

--, .

Ill
c
Ill
v

Dolichorh inus

w
4;

hyognathus

a.
a.
::>

'\

Ma nteoceras
ma nteoceros

Ill
-o
-o
-

Ill
c
Ill
v

0
::E w

{'/-.;
.

\ . . ..
.

1. ;
t:::;t >

Ill
c
Ill

Lambdotherium
popagilum

,.,"),<

Mesatirhi us
petergon1

;;f

....

Eotitanops

Eotitanops

prin ceps

gregoryi

EVO LUTION O F PROBOSCI D EANS, G R EATLY S I M PL I FI ED.

( A FTER OS BO RN . )

E le p ha s

Pleist.- Rec .

Stegodon

P l i o .- Pieist.

P l io .

Mammut

M i o .- P i i o .

Gomphotherium
Moe ritherium

Eo c . - O i ig .

Palaeomastodon

Olig.

Mio.-Piio.

1 37

G E O G RAPH I C D I ST R I B U T I O N of livi n g m a m m a l s re

fl ects the pattern of i n terco n nection betwee n conti

n e nts during the geologic past.
E U ROPE, ASIA,
AMERICA were

AND

N ORTH

co n n e cted for
m u c h of Cen ozoic t i m e , a l low
ing m ig ra tion a n d e x p l a i n i n g
m a n y si m i l a ri ties o f t h e i r pre s
e n t fa u n a s . T h e d i fferen ces t h a t
do e x i s t reflect differi n g c l i
m a ti c e n v i ro n m e n ts a n d rece n t
develo p m e n t o f desert a n d

m o u n ta i n ba rriers to m i g ra t io n .
The fa u n a s o f South A merica,
Austra l ia, a n d Africa south of
the Sahara a re q u ite disti n ct.
These conti n e n ts have been
separated fro m o n e a nother
throu g hout t h e Cenozoi c. North
Africa n m a m m a l s a re m ore s i m
i l a r t o those o f Europe.

Reindeer

Bison

W i l d Horse
PlAEARCTIC

Hedgehog

.. .

...

O R I ENTAl

I n d ia n E l e p h a n t

Flying
Phalanger

1, -t-

..

..

Marco Polo
Sheep

Koa l a
AU STRA l i A N

K a n g a roo

I S O LATION of South America

a n d Australia p rod uced very
d ifferent ma m ma l i a n fau nas, i n
wh ich marsupials were a t fi rst
the dom i n a nt for ms. They re
main abundant in Austra l ia be
cause of its conti n u i n g isola
tion. I n tercon nection of North
and South America i n the late
Tertiary led
to
com petition
and exti nction of m a n y South
America n p l a ce n ta ls.
Convergent evolution i n ex
ternal form
between
North

American placental m a m mals

and fossil South Am erica n mar
supials (p. 1 3 1 ) de monstrates
the i nfluence of natural selec
tion in ada ptation to s i m ilar
modes of l ife.
Geog ra phic distribution of
other a n i m a l g roups does not
n ecessa rily s h ow s a m e bou n d
aries a s m a m m a ls . Pla n ts a n d
m a ri n e i n vertebrates, f o r e x
a m ple, have q uite differe n t dis
persal m e a n s , a n d therefore
different d istribution pa ttern s .

Mou n ta i n Goat
M u s k Ox

N EA RCTIC

Caribon

Porc u p i n e

Pro n g h o r n A ntelope

K i n k a j ou
Howler Monkey

Capybara

1 39

Arboreol
i n sectivores

Relation s h ips between the m a i n g ro u ps of pri m a tes.

bert. )

PRIMATES a re t h e m a m m a l ia n

( After Co l

o r d e r to w h i c h l e
m u rs, ta rsiers, m o n keys, a pes, a n d m a n be lon g . They
ten d to be rather ra re a s fossils, l a rgely because of
their c h a r a cteristi c a l l y a rborea l h a bits . Most prim ates
show two fu n d a m ental a d a ptations to t h e i r tree-dwe l l
i n g existen ce : stereoscopic v i s i o n a n d h a n d s c a p a b l e
of g ra s p i n g . These two features, present i n a l l but the
m ost p r i m itive m e m bers, a l l ow the p ri m ates to j u d g e
d i stan ces accu rately a n d t o swi n g fro m b ra n ch to
bra n c h . They were a lso i m porta nt, together with his
l a rge brai n , i n t h e d evelo p m e n t of g ro u n d - dwel l i n g
m a n , a l lowi n g h i m t o develop i n crea s i n g s ki l l s i n m a k
i n g a n d u s i n g tools.

1 40

PR O S I M I A N S (pre-monkeys) i n

clude living l e m u rs, aye-ayes,

bushbabies, a n d the more mon
key- l i ke tarsioids. They a rose in
the Pa leoce n e , probably from
arborea l i n sectivores, a n d be
came d ivers ified d uring the
early Tert i a ry. T h ey d e c l i n e d
i n n u m be rs d u r i n g l a t e Tertiary
t i m e s, proba b l y beca u s e of
com petition from their des c e n
da nts, t h e a n t h ropoids. Prosi m
i a n s st i l l s u rvive i n s u c h places
a s Madag asca r and Southeast
A s i a . Prosi m ia n s have less w e l l
developed b i noc u l a r v i sion a n d
g rasp i n g l i m b s t h a n ot her pri
mates.

Notharctos, an Eocene prosi m

ian, a bout 1 8 i n ches h i g h .

i n c l u de mon
keys, a pes, a n d men. They de
veloped i n t h e O l i g oce n e a n d
Miocene from pri m i tive p ros i m
i a n a n cestors . O l d World m o n
keys s h ow funda m e ntal diffe r
e n ces fro m those of t h e New
World. Flat-nosed, pre h e n s i le-

ta i l e d South A m erica n form s,

such a s m a r mosets, ca p u c h i n s ,
a n d spider m o n keys , see m t o
be more pri m i tive. O l d World
and New World m o n keys a rose
i n depen d e n t l y fro m prosi m i a n s .
Their s i m i la rities a re t h e res u l t
of converg e n t evo l ution .

ANTHROPO I D S

P L E I ST O C E N E

Ne w World
Monkeys

;\

,_____

Australopithecus-Homo

o n key

. .

;,

PI i o p i t h e c u s

O reop

"'!!!_ _ _ _ __j----

___ ________

- -

ec

- - - - -

l_o ;;;pith; us

APES

"

It

R a m a pithecus
MEN

EVOLUTION OF P R I MATES
(After McAlester)

f.l
,.,....

#,

.:

\u'!

!.

Gibbon

1)., .
.

C h i m p a n zee

...
Gori l l a

, Pon g i d a e

Hylobati d a e

(Apes)

RECENT

PLEI STOCE N E

Oreopithecidae

PLIOCENE

'
'
'
... , - - - - - - -- - - - ---

M I OC E N E

'
'
'

'
I

'
'. - - - - - - - - - - ''- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - '

('
l \

\.,. .,.
--

'''
: .;

..
..

Pliopithecus

Oreopithecus

Dryopithec us

HOMINOIDS-a pes a nd men-a re cl a ssified together

i n a s i n g l e superfa m i ly, Hominoidea. They show fewer

d ifferences fro m o n e another t ha n d o the O l d World
fro m the New World mon keys, which a re i n sepa rate
superfa mi l i e s . The h istory of the h o m i n o i d s a bove
shows their possi b l e evol utio n a ry relatio n s h i p s as re
vea led by s k u l l s a n d by d ental patterns.
1 42

.,.
-,, ,,,
,
..
.
.,
I>
.,
.,
.

Ora n g u t a n

Homo

Austra lopithecus
Hominidae

(Men)

R a m a p ithecus
.
I
I
I
.
I
I
I

- - - - - - - - - - - -'

- - - - - - - - - - - - - - - - - - - - -

.
I
J

R a m a p ithecus

LIVI N G APES i n c l u de the ch i m

panzee a n d gori l l a , w hi ch a re
c h iefly g rou n d - l i v i n g forms, a n d
t h e g i b bon a n d t h e ora n g uta n,
which a re bea uti f u l ly ada pted
to a rboreal l ife. All lack the
typical ta i l of mon keys. Al l , e x
cept perha ps the g i bbons, seem
to h ave arisen from g e nera l -

Austra lopithecus

iz:ed a pes that were wides pread

in t h e Old World in M iocene
and P l ioce ne times. Dryopithe
cus (Proconsu/J, w h i c h i n c l u ded
severa l forms most probab l y of
apelike proporti o n s , may a l s o
h ave given rise to t h e a n cestors
of m a n . ( I l l u s tra tio n s a d a p ted
from m a n y a u t h o rs . )

1 43

THE FAMILY O F MAN w h ich s p a n s t h e l a st 2 m i l l i o n

yea rs, i n c l u d e s t h ree g e n e ra . Two of t h e m a re n o w ex

t i n ct. Because of the ra rity of pri m ate fossils a n d per
haps a l so because of the i nten s e i n terest in the o ri g i n
o f m a n , there i s so m e d i s a g re e m e n t con ce r n i n g t h e
deta i l ed relatio n s h i ps o f particu l a r species wit h i n t h e
broad patte r n o f evo l uti o n a ry develo p m e n t . N ew d is
coveries a re sti l l bei n g m a d e, a n d the p rovisio n a l ac
cou n t g iven here m ay well req u i re l ater m o d ification .
RAMAPITHECUS, a

sti l l poorly
known
h o m i n id,
has
bee n
fou n d i n late Miocene a n d Pl io
cene rocks of I n dia a n d Africa.
The pa.ttern of its teeth s hows
a rather s mooth s e m i c i rcular
outli ne, which is far more si m i
lar to that of livi ng m a n t h a n
to the quadrate pattern o f the

apes (p. 1 4 2). Little is k nown

.
about other pa rts of t h e s kele
ton ; a n d because of this, the
reco n struction shown below is
very te n ta tive. But the tooth
pattern i s so " h u m a nistic" t h a t
i t seems p robable t h a t Ramapith
ecus
wa s close ly re lated to
modern m a n .

(Southern
ape) i s a lso regarded a s closely
re lated to modern m a n , prob
a b ly d i rectly a n ces tra l to t h e
genus Homo t o which w e as
sig n o u r. own species, Homo
AUSTRALO P J TH E C U S

sapiens.

A u s t ra lo p i t h ec i n e s , o n c e w i d e
sp read i n Afri ca, a re now
t h o u g h t t o i n c l u d e two s p e c i e s ( p .
1 4 3 ) . T h e y were g ro u n d d w e l l e r s
a b o u t 4 f e e t ta l l . A l re a d y , how
ever, t h ey had a n u p r i g h t pos
t u re . Bones f o u n d w i t h t h e i r re
m a i n s s u g g est t h a t t h ey were
ca r n i vores, but t h i s i s not cer
ta i n . In d e n ta l pattern a n d i n
g e n e r a l s k u l l fo r m , t h e y were

very m a n l i ke, desp ite their rat h

e r p rotru d i n g jaws a n d brow
ridges. T h e i r bra i n capac ity
(about 600 cc) was only half
that of modern man.
There is sti l l s o m e dou bt
w het her crudel y c h i pped stone
tool s associated with fossi l de
posits were made and u sed by
a us tra lopi t h ec i n e s or by t h e i r
desce n da nts a n d u l t i m ate con
te m poraries,
Homo
erectus.
Austra lopithecines beca m e e x
tinct a bout h a l f a m i l l io n years
ago.
Rece nt
d is cove ries
in
Ke nya suggest t h a t early forms
m ay date back a s far a s 2.6
m i l l io n yea rs.

1 45

W H AT I S M A N ? T h i s question is s u rp ri s i n g l y d iff icu l t

t o a n sw e r w h e n a p p l ied t o fos s i l s . I t see m s b ette r to
restrict t h e term " m a n " to o u r own s p e c i e s , Homo s a
piens, a n d to re g a rd o t h er c l o s el y r e l a ted fo r m s a s p re
h u m a n , t h o u g h so m e of t h e s e m a n l i k e c reatu res d i d
s h a re t h e h u m a n c h a racte r i stic o f too l m a n ufa c t u r e .
Mod e r n m a n a p pea red a bo u t 5 0 0 , 0 0 0 yea rs a g o .
HOMO ERECTUS i s known fro m
fos s i l s fou n d i n t h e Ple istoce n e
sed i m e nts ra n g i ng fro m about
750,000 to 200,000 years in
age. Though sometimes de
scribed by other names ( m ost
co m m o n ly Pithecanthropus}, i n
d ividuals of the species a re
k n own from Java , C h i n a , Af
rica, and Asia. H . erectus was
an erect, g rou n d - dwe l l i n g i n d i
vidual w h o fash ioned various

tool s a n d was a pparently a

h u nter. Ma n l i ke i n stru cture a n d
i n a p peara nce, H . eredus h a d
a b ra i n capac ity of 900- 1 1 00
cc, i n termed iate between that of
A ustrafopithecus
and
modern
man. H. erectus was a contem
pora ry a n d perhaps a com peti
tor of l a te r a u s tralopithec i n es
from w hose e a r l i e r m e m bers
"he" evolved. H. e re cf us used
fire and led a com m u nal life.

MODERN MAN, Homo sapiens,

seems to have a risen from H .
erectus. For a l m ost 200,000
years, t h e two s pecies were
con te m poraries. Modern man is
cha racterized by less cons piCu
o u s brows a n d j a w s t h a n t h e
earlier h o m i n i d s a n d h a d a
m uc h larger b ra i n ( av. capa
city about 1 3 5 0 cc ) .
Severa l races were i n vo lved
in the fos s i l h istory of m a n . Ne

g a rded as a race of o u r own

species, l ive d t h roughout Eu
rope , the Med iterra nean area,
and parts of Asia Minor from
about 1 1 0,000 to 35,000 years
ago, a period that i n c l u ded
t h ree e pisodes of g l aciation.
Nea nderthal m e n l ived in caves
a n d were s k i l lf u l too l m a kers
a n d h u nters. T h ey were n ot t h e
s t u p i d brutes t h ey a r e often
pictu red as being.
MAN replaced
Nea n dert h a l Ma n i n Europe
about 35,000 yea rs ago, prob
a bly m i g rati n g fro m the Mid
d l e East. Physically s i m i la r to
modern m a n , Cro- M a g n o n m a n
m a n u fa c t u red
s u perior
tools
and prod u ced m a sterpieces of
art and scu l p t u re ( p . 1 4 9 ) .

CRO-MA G N O N

a n dertha l m a n , l o n g regarded
a s a d isti nct species, was a race
of h eavy- browed , m uscular in
d ividuals. The later C ro-Magnon
race had facial features that
more c l osely rese m bled those
of modern m a n . Alth o u g h these
d ifferences are rea l , t h ey seem
a n a logous o n ly to those of l iv
ing
human
races,
betwee n
wh ich i n terbreeding freq u e ntly
ta kes place.

T H E EVOLU T I O N O F TOOLS. We a po n s, societies, a n d

c u l t u res a ri s e fro m a n d reflect m a n ' s p hysic a l a n d

m en t a l evo l utio n . B i n o cu l a r vision , m a n u a l d exterity,
a n d i n crea si n g m e ntal capa city were p a ra l leled by t h e
i n crea si n g perfectio n o f m a n ' s work a s a craftsm a n .
T H E O LD E S T TOOLS were prob
ably used b u t not made, con
sisting of stones a n d bou lders
conve n i e n tly shaped by natu re.
La ter too ls were crudely c h i pped
and s h a ped axes and scra pers .
These were gradually s u pple
m e nted and replaced by d e l i
cately c h i p ped b l a d e s a n d ar
rowheads of a va riety of mate
r i a l s , i n c l u d i n g bone.

About 1 0,000 years a g o, t h e

early c u l t u re of c h ipped i m
p l e m e n ts, t h e Pa l eo l i t h ic, g a ve
way in E u rope to the Neol i t h ic,
w h i c h was m a r ked by f i n ely
g rou n d and pol i s h e d too l s a n d
wea pons. About 5 ,000 yea rs
ago, man f i rst learned to fa s h io n
i m p l e m e n ts of m e ta l . Th e "sto n e
a ge" sti l l pers i sts a m o n g s o m e
l iv i n g peo p l e s .
SCRAPER
Mousterian

HAND AXES
Acheulian

SPOKESHAVE
A u ri g n a c i a n

BONE A N D A N T L E R W E A PONS
Magdelinian

W E A PON H EADS
So l u trean

M A M M O T H C A RV I N G
Magdelinian

C U LTURAL EVOLUTI O N of m a n

i s g l i m psed i n c ave p a i n t i n g s
a n d carv i n g s t h a t d a t e t o about
2 8 ,0 0 0 yea rs ago. Both a re
c h iefly depicti o n s of h u n t i n g
a n d ferti l i ty. T h e y m ay h a ve
had " m a g i ca l " s i g n i fi ca n ce .

Ma n 's a n c i e n t belief in s u r
vival a fter death is s h own by
N ea n d e rt h a l a n d C ro-Ma g n o n
s ke l e to n s b u ri e d i n feta l o r
s l e e p i n g pos i t i o n s , w i t h i m p l e
m e n ts a n d to k e n s to be u s e d
i n the new life.

149

EVO L U T I O N O F H U MAN SOCI E T I E S a rose fro m m a n ' s

g rowing a d a ptation to h is e nviro n m e nt. Such l a n d
m a r ks a s t h e d iscovery a n d u s e o f fi r e b y Homo erec
tus a n d the d eve l o p m e n t of crop cultivation , a n i m a l
h us b a n d ry, a n d pottery b y Neolithic m e n prod uced
ra d i cal c h a n g es in the patter n s of h u m a n life. Man ,
ori g i n a l ly a n o m a d i c h u nter a n d herbivore, cou l d then
constru ct dwe l l i n g s a n d gather into g roups who estab
lished settl e m e nts.
The need for com m u n i cation fostered the d evelop
m en t of in creasi n g ly sop h i sticated l a n g u a g e . The g row
i n g size of co m m u n ities n ecessitated the d ivision of the
various ta sks involved i n s u rviva l and the creation of
some fo rm of govern ment. We know l ittl e of t h e de
tailed d evelo p m e n t of a n y of th ese, for writi n g was
not i nvented until a bout 5,000 yea rs a g o . Early re
cord ed history i s very patc hy, bei n g i n fi n itely m o re
co m pl ete for some pa rts of the world, such a s Egypt,
t h a n for others . Th e earli est societies a n d thei r u nwrit
ten l a n g u a ges, u n l i ke the i m p lements of ea rly m a n , l eft
no records i n t h e sto n e .

1 50

TH E M EAN I N G OF EVOL UTI O N

Even before D a rwin publ ished The Origin o f Spe
cies, s o m e rel i g ious l e a d e rs atta c ked t h e concept of
evo l ution beca use t h ey t h o u g h t it t h re a t e n e d t h e i r
view p o i n t; o t h e rs h a ve e m b ra ced it a s a n ew i n si g h t
i n to t h e wor k o f G o d i n t h e created worl d . Evo l utio n
a ry t h eorists h ave c l a i m e d evo l u t i o n a s a j u stifica tion
for m i l ita nt politi c a l tactics; others h ave endorsed it as
i l l ustrati n g t h e i n evita b i l ity of h a rmonious political de
velo p m ent. So m e eco n o m ists h ave clai m ed it as an ar
g u m en t for la issez-faire eco n o m i c policies, while a few
scientists h ave u sed it a s a basis for a n ew code of
ethics. So m e po p u l a r writers, atta c k i n g trad ition a l re
ligious belief, h ave a d o pted evol utio n i s m as a n ew re
l i g i o n . Se l d o m h a s a scientific theory so q u i ckly beco m e
a l l t h i n g s t o a l l m e n . Seldom h a s a n atu ral pro cess
been so ca relessly used as a n expository basis for the
whole pattern of h u m a n life .
Contem porary
ca rtoon
and
verse i n d icates Victorian i n ter
est i n the m ea n i n g of evo l u tion

A m I Satyr o r M a n ?
P r a y te l l m e w h o c o n ,
A n d sett l e m y p l ace i n t h e s c a l e ,
A m o n i n a pe' s s h a pe ,
An a n t h ropoid a p e ,
Or m o n ke y d e p r i ved o f h i s ta i l ?
T h e Ves t i g es ta u g h t ,
That all came from naught
By "deve l o p m en t , " so c a l l e d ,
' ' prog ress i v e ; "
T h a t i nsects a n d w o r m s
Ass u m e h i g h e r f o r m s
By m od i f i c a t i o n e x cess i v e ,
T h e n D A RW I N set f o rt h ,
I n a book of m u c h w o r t h
T h e i m p o rta nce of " N a t u re ' s s e l e c t i o

M O N K E Y A N A.

V i ctor i a n ca rtoon d e p icts a pu zzled Darw i n a n d h i s a n cestors.

The I m p l i cations of Evo lution

The proces s of evo l u tion i s a
fact. N u m e ro u s l i n e s of evi
dence i n d i cate t h e desce n t of
new s pecies by m odification of
a n cestral form s over extended
perio d s . A l t h o u g h t h e m ech a
n i s m i s s ti l l t h eore t i c a l , t h e re
is very s tro n g evidence t h a t
n a t u ra l s e l ecti o n , g e n e t i c v a r i
a t i o n , a n d i s o l a tion a re t h e
c h ief co m po n e n ts . ( p. 1 0 2 )
Evol u t io n , l i k e a n y other nat
u ra l process o r scientific t h e
ory, i s t heolog ica l ly n e u tra l . I t
d e scri bes m e c h a n i s m s, b u t n ot
m ea n i n g . I t i s based u po n t h e
recog n i ti o n of order b u t i n
corporates n o concl u s io n co n
cern i n g t h e o ri g i n o f t h a t or
der a s either p u rposef u l or
p u rpos e l e s s .
A l t h o u g h evo l u ti o n i n vo lves
t h e i n terpret a t i o n of n a t u ra l
eve n ts b y n a t u ra l processes, i t

1 52

n e i t h e r a s s u m e s n o r provides
parti c u l a r co n c l u s i o n s co ncern
i n g t h e u l t i m a te s o u rces or t h e
s i g n i fi c a n ce of m a teria l s , events
o r processes.
Evo l u t i o n p rov ides no ob
concern i n g
co n c l u s i o n s
vious
p o l i t i c a l o r eco n o m i c syst e m s .
Evo l u t i o n n o m o r e s u pports evo
l u t io n a ry pol i t i cs (whatever t h e y
m i g h t b e ) t h a n d o e s t h e Seco n d
L a w o f T h e rmodyna m i cs s u p
port political d i sorder or eco
n o m i c c haos .
Evo l u ti o n offers no b a s i s for
ethics. It i s n o t s e l f -ev i d e n t t h a t
s u rviva l i s t h e h i g h e s t good
a n d that any mea n s of i t s a t
ta i n m e n t i s v i rt u o u s . T.H . H u x
l e y w rote " T h e e t h ica l pro g
res s of society depen d s , n o t o n
i m i ta ti n g t h e co m i c proces s , s ti l l
less i n r u n n i n g away from i t,
b u t i n co m ba t i n g i t . "

fo r man,
h e n ce is a s i g n ifi c a n t contri bution to h u m a n u n der
sta n d i n g . Reco g n iti o n of the i m m ensity of the spa n of
geolog i c ti m e , t h e aweso m e scale of cos m i c d i m e n
s i o n s a n d p ro cesses i n volved i n t h e l o n g period o f p re
o rg a n i c evo l ution, a n d the p l a ce of m a n h i m self wit h i n
the e n d l ess d iversity o f t h e tee m i n g life on t h e fra i l
su rfa ce o f o u r p l a n et-a l l these hel p t o e n l i g hten a n d
susta i n m a n a s he faces the c h a l l e n g e, the d i l e m m a ,
a n d t h e mystery o f h i s h u m a n con d ition .
EVOLUTION

PROV I D ES

PERS PECTIVE

Ma n k i n d , t h e p ro d u c t of o r g a n i c e vo l u t i o n , i s n o w
te c h n i c a l l y e q u i p p e d w i t h p o w e r , i f n o t t li e wi l l , to c o n
t r o l t h e f u t u r e d e ve l o p m e n t of l i fe o n e a rt h . P s y c h o so c i a l
evo l u t i o n

h a s n o w d i s p l a c e d t h e o l d e r p ro c e s s e s of

o rg a n i c evo l u ti o n i n h u m a n co m m u n i t i e s . K n o w l e d g e ,
t r a d i ti o n s , va l u e s , a n d s k i l l s a r e n o w t ra n s m i tt e d fro m
o n e g e n e r a t i o n to a n o t h e r t h ro u g h b o o k s a n d te a c h i n g
i n st i t u t i o n s

rath er

than

being

learned

a n ew

"fro m

s c r a t c h " by e a c h n ew i n d i v i d u a l .

Psyco-Social

E a c h seg m e n t

Evolution

represents
a p p ro x i m a t e l y
5 0 m i l l io n
years

of
Earth
Appearance of oldest known
foss i l s

1 53

THE FUTUR E EVOLUTION OF MAN, of other species,

a n d perhaps of the whole i ntricate ecosystem of which

we are a p a rt now sta n d s i n jeopa rdy. The pol l ution of
the atmosph ere on which o u r existe n c e depends h a s
now reached a c r i s i s of m a j o r p roportions in most i n
d ustri a lized a reas o f t h e worl d . Rapidly dwi n d l i n g re
serves of s uch essential co m m od ities as petro l e u m a n d
ma ny m etals th reaten t h e future not o n ly of i n d ustria l
pro d u ction but a l so of technologically b a s e d society.
A conti n u i n g explosion of h u m a n population, espe
cia l ly i n the less i n d ustri a l i zed areas of the worl d ,
raises the awesome possi bil ity of wid es p read fa m i n e
a n d sta rvatio n within the n ext 3 0 yea rs.
Man a l ready possesses the tech n i cal power to solve
these th ree m a j o r prob l e m s : poll ution, dwi n d l i n g m i n
era l resou rces, a n d overpopu lation . Whether h e has
the wisdo m , t h e wil l , and the e n e rgy to solve them re
m a i n s to be seen . It is i ro n i c that the future of the age
lon g p rocess of o rg a n i c evo l ution m ay n ow depend on
the conscious c h o ice of m a n, a p roduct of that process.
The d a n g e r, the c h a l l e nge, a n d t h e choice i nvolve m a n
kind i n a com m o n peril a n d a com m o n hope.
Evo l ution p rovi des n o easy a n swers to m a n ' s long
search for m e a n i n g a n d no i n sta nt sol utions to m a n 's
most p ress i n g p ro b l e m s . It i s rather for m a n h i m self
now to p rovi d e the i n put-i n the recog n ition of an
eth i c beyo n d th a t of s u rviva l , of a p urpose b eyo n d
that o f g a i n , a n d o f a v i s i o n o f l ife beyo n d that of
m e c h a n i s m a n d p rocess. On such col lective com m it
ments of m e n a n d of n ations depend the s u rvival of
m a n k i n d a n d the future cou rse of evo l ution .

1 54

Po l l ution of a t m o s p h ere in i n d ustri a l areas is a worldwid e prob

l e m . World fa m i n e poses an i n creasi n g ly s evere t h reat a s bur
ge o n i n g population co m petes for l i m i ted res o u rces. World popu
lation projections suggest g loba l population of 2 5 bi l l i o n by 2 0 70 .
2 5 b i l l i o n p e o p l e b y 2 0 70

World population q u a d rupled

by 2044

1 6 1:
a.
0
..c:

12
10 5
:::
14 U
Q;

V)

Asia : q u a d rupled by 2040

Wor l d : d o u b l e d by 2 0 0 7 ;
po s s i b l e fa m i n e
Asia : doubled by 2 0 0 5

MORE

I N FORMAT I O N

T h e fo l l o w i n g l i s t of books i s o n l y a n i n trod u c t i o n to t h e vo l u m i n o u s l i te ra t u re
on evo l u t i o n . Many m u se u m s a l so prov ide d i sp l a y s , ta l k s , a n d l i terat u re .
H i stori c a l
D a rw i n , C h a r l e s , T h e O r i g i n o f Spe cies, O x f o r d U n ivers i ty Press, N . Y . , 1 9 5 6 .
( 6 t h ed . , 1 8 7 2 re p r i n ted : T h e W o r l d C l a s s i c s . )

Gre e n e ,

J . C . , D a r w i n a n d t h e M odern World V i e w , Mentor Books, N . Y . , 1 9 6 3 .

A . , D a r w i n a n d t h e S e a g l e , H a rper a n d Row , N . Y . 1 9 6 9 .

Moore h e a d ,

T h e Proce s s o f E v o l u t i o n
D e B e e r , G . , A t las of E v o l u t i o n , N e l son a n d S o n s , l o n d o n , 1 9 64 .
Mayr, E . , A n im a l Spec ies and Evolution , Oxford U n ivers i ty Press, N . Y . 1 9 6 3 .
Moore , R . , Evolution , T i m e - L i fe, I n c . , N . Y . , 1 9 6 2 .
S a v a ge ,

J . M . , E v o l u t i o n , H o l t , R i n e h a r t o n d W i n s to n , N . Y . , 1 9 6 3 .

Sheppard, P . M . , N a t u r a / S e le c t i o n a n d Heredity , H u tch i n so n , l o n d o n , 1 9 5 8 .

S i m p s o n , G . G . , T h e Major Features o f E v o l u t i o n , Co l u m b i a U n ivers i ty Press,

N . Y . , 1 95 3 .
Smith,

J . M . , T h e Th eory o f E v o l u t i o n , Pe ng u i n B o o k s , H a r m o n d s w o r t h , 1 9 5 8 .

Tax, S . , E v o l u t ion after D a r w i n , U n ivers i ty o f Ch i c a g o Press, C h icag o , 1 9 6 0 .

T h e Course o f E vo l ut i o n
C o l bert, E . H . , E v o l u t ion o f t h e Vertebrates, J o h n W i l e y , N . Y . , 1 95 5 .
Rhode s ,

F . H . T. , T h e E v o l u t ion o f l ife , Peng u i n Books, B a l t i m ore, 1 9 7 4 .

F . R . S h affe r, Foss ils, a G u ide to Pre h istor ic

Rhode s , F . H . T . , H . S . Z i m , a n d
L if e , G o l d e n Press, N . Y . , 1 9 6 3 .

The E vo l ut i o n of M a n

T . , M a n k i n d E v o l v i n g , Y a l e U n iversity P r e s s , N e w Have n , 1 9 6 2 .
F. C l ark, Ear ly M a n , T i m e - l i fe B o o k s , N . Y . , 1 9 6 5 .
LeGros C l ark, W. E . , H istory of t h e Primates, B r i t i s h M u se u m of N a t u r a l H i s

Dobzh a n s k y ,
Howe l l ,

tory, l o n d o n , 1 954; T h e Foss i l Ev idence of H u m a n E v o l u t ion , U n i v e r s i ty of

Ch i c a g o Pre s s , C h i c a g o , 1 96 4 .
O a k l e y , K . P., M a n t h e Tool-Maker, B r i t i s h M u se u m af N a t u ra l H i story, l o n
d o n , 1 96 3 .
T h e Me a n i n g o f E v o l ut i o n
Barbour,
Lack, D . ,

I . G . , Issues in Science and R e l i g i o n , S C M l td . , l o n do n , 1 9 6 6 .

E v o l u t ionary Theory a n d Christian B e fief; th e Un reso lved Con f l i c t ,

Meth u e n a n d Co . , l td . , l o n d o n , 1 9 5 7 .
S i m p s o n , G . G . , T h e M e a n i n g of E v o l u t ion , M e n t o r B o o k s , N . Y . , 1 9 5 1 .
Te i l h ard de Chard i n ,
1 95 9 .

P. , T h e P h e n o m enon of M a n , H a rper a n d B r o t h e r s , N . Y . ,

PHOTO C R E D I TS : p . 6 - 7 , To u r g o P h o t o Serv i ce; p . 1 0 , Bett m o n n A r c h ives; p . 1 2 ,

D ra w i n g by Sa m u e l l a w re n ce , p h o t o g r a p h repri n ted by perm i s s i o n of George
Rai n b i rd , l td . , Bett m a n n A rch ives; p . 1 4 , Bettm a n n A rc h i ve s ; p . 1 5 , C u l v e r Pic
t u res; p. 1 6 , George R a i n b i rd , l td . , p . 1 8 , Reyno l d H. C h ose; p . 1 9 , Rey n o l d
H . C h o se; p . 2 0 , Bett m a n n Arc h ives; p . 2 2 , C u l ver P i c t u res; p . 2 3 , Bet t m o n n A r
c h ives; p . 2 4 , U n i ted N o t i o n s ; p . 2 6 , Bettm a n n A rc h i ves; p . 2 7 , Bell m a n A rc h i ves;
p . 2 8 , Victur A . M c K u s i c k ; p . 4 6 , T h e American M u se u m of N a t u ra l H i story;
p . 4 7 , T h e A m e r i ca n M u s e u m of N a t u r a l H i s to ry; p . 7 2 , Photo Resea r c h e rs ;
p . 1 1 0 , E . S . Borg h o o r n , ( H a rv a rd U n i v e rs i t y ) ; p . 1 5 1 , P u n c h M a g a z i n e 1 8 6 1 ;
p. 1 5 2 , from Ev o l u t i o n a ry T h eory of C h r i s t i a n Be l i ef, p . 1 54 , U n i ted N o t i o n s ;
p . 1 5 5 , U n i ted N o t i o n s .

1 56

I N DEX
A c a n t h od i a n , 1 1 5
Acorn worms, 1 1 4
Ada m , 6
Ada p ta t i o n s , 2 5 , 3 3 , 4 2 ,
54, 79, 8 1 ' 88
Agnatha, 1 1 4, 1 1 6
A l g a e , 5, 1 1 0 , 1 2 0
A l lele, 6 1 , 66
A m b l y pod , 1 3 2
A m i no acids, 70, 7 1
A m m o n i a , 1 05 , 1 06
A m m o n i te , 1 09
A m p h i b i a n , 1 1 7 , 1 1 9 , 1 24
A n a l o g o u s s t r u c t u re s , 3 9
An c o n s h ee p , 7 4
A n e m i a , s i c k l e - ce l l , 9 2
A n g i os pe r m s , 1 2 1 , 1 2 2 ,
1 23
A n i m a l s , n u m be r o f , 5
o l dest, 1 1 0
A n ky l osa u rs , 1 2 7
A n t e a t e r , s p i n y , 53 , 1 3 0
A n t h ro p o i d s , 1 4 1
A n t i a rc h , 1 1 4
Ape, 1 40 , 1 4 1 , 1 4 2 , 1 4 3 ,
1 44
A ra u c a r i a , 5 3
A r c h a eoptery x , 5 2 , 9 4 ,
1 29
A r i s t o t l e , 7, 1 0
A r m ad i l l o , 1 3 5
A rs i n a t h e r e s , 1 3 5
Art h r od i re , 1 1 5
A rt h ropod s , 5, 1 1 2 , 1 1 8 ,
1 19
A rt i f a c t s , 1 09
A r t i o d a c ty l s , 1 3 5
Astra p o t h e re s , 1 3 5
A t m o s p h ere, 1 0 7
ATP, 40
A u s t ra l i a , 1 1 0 , 1 3 8 , 1 3 9
A us t r a l op i t h e c u s , 1 4 3 ,
1 45, 1 46
A y e - a ye , 1 4 1
Bacte r i a , 1 1 0 , 1 2 0
Bat, 1 35
B e a g l e , voya g e o f , 1 6 ,
1 7, 1 9
Bea r , 8 6 , 87
Beaver, 1 3 4
Beri n g S t r a i t s , 77

B e r k e n d e r , L C . , 1 07
" B i g b a n g " t h eory, 1 04
B i rd s , 1 1 9 , 1 27 , 1 2 8 , 1 2 9
B irke n i a , 1 1 4
Bison, 1 34
B i ston b e t u la r i a , 47
Blending, 61
B l ood p i g m e n t s , 40
Bove r i , T . , 28
B ra c h i opod s , 1 1 1 , 1 1 2 , 1 1 3
B reed i n g , s e l e c t i v e , 4 4
B rya n , W i l l i a m J e n n i n g s ,
30
B r y o p h ytes, 1 2 0
B ryozoa n , 1 1 2 , 1 1 3
Bush boby, 1 4 1
Coec i l i o n , 1 1 7 , 1 2 4
Co l o m o i t e s , 1 20
Co l o m oph y t o n , 1 2 1
Ca m b r i a n , 1 07, 1 1 1 , 1 1 2 ,
1 13
Ca m e l , 1 2 6 , 1 3 1 , 1 3 4
Ca m e l ops, 1 3 4
Cope V e r d e I s l a n d , 4 3
Capuch i n , 1 4 1
C a r b o n , 3 5 , 97, 1 0 5
Ca r b o n d i o x i d e , 1 05 , 1 06
Ca r bo n i fe r o u s , 1 1 8
Carcho rodon , 1 1 5
C a r n i v o re s , 1 3 3 , 1 3 5
Carti l a g i n o u s f i s h , 1 1 5 ,
116
Castoroides, 1 34
Cot, 1 33 , 1 34
C o to s t r o p h i s i s , 1 4
Cott l e , 1 3 4
Cove bea r , 8 6
Ce l l d i v i s i o n , 5 8
s t r u c t u re , 3 2
Ce n oz o i c , 1 1 6 , 1 2 9 , 1 3 1 ,
1 3 2, 1 34, 1 3 8
Cep h a l op o d s , 1 1 2 , 1 1 3
Cerotops i o n s , 1 27
Cetacea n s , 1 3 3 , 1 3 5
C h o l i co t h e r e s , 1 3 1
C h a m e l eo n , 4 2
C h i asmata, 73
C h i m pa n zee , 1 4 2 , 1 4 3
Ch i ro pte ro n s , 1 3 5
C h oo n i c h t h ye s , 1 1 7
C h o n d r i c h t hyes, 1 1 5

C h o n d r i te s , c a r b o n a ce o u s ,
1 05
C h ordata , 1 1 4
C h o rd a t e s , 5
C h r o m a t i n , 67
Ch r o m o s o m e s , 2 8 , 2 9 , 5 6 ,
57, 58, 59, 63, 65,
66, 67, 73, 74
C lodose loch e , 1 1 5
C l o m s , 4 9 , 1 09 , 1 1 9
Classificat i o n , 36
Climati us, 1 1 5
C l u b m o s s , 1 20 , 1 2 1
C o c k r oa c h , 1 1 8
Cocos I s l a n d , 83
C o l o r b l i n d n e s s , 66
C o m m u n i t i e s , 35
C o n dy l o rt h , 1 3 2 , 1 3 5
C o n i fe r s , 5 , 1 2 1 , 1 2 3
C o n v e rg e n ce , 1 2 8
C o ra l s , 1 1 2 , 1 1 3
C o rd a i te s , 1 2 1 , 1 2 3
Coryph odon , 1 3 2
C o t y losaurs, 1 2 6
C reod e n t , 1 3 3
Cretaceo u s , 5 0 , 1 2 3 , 1 2 9 ,
1 3 1 ' 1 32
Crocod i l e s , 1 2 7
C r o - M o g n o n , 1 47 , 1 49
C r o s s i n g - ov e r , 7 3
Crosso pte ryg i a n , 9 4 , 9 5 ,
1 1 6 , 1 1 7, 1 2 4
C r u stacea n , 1 1 9
C u l t u re , e v o l u t i o n o f ,
1 49
Cuvier, Georges, 1 4
C y c a d s , 1 20 , 1 2 3
C y n o g n a t h us, 5 3
D a r w i n , C h a r l es , 1 5 , 1 6 ,
1 7, 1 8 , 1 9, 20, 2 1 ,
22, 23, 24, 25, 28,
42, 43, 44, 80, 82,
91 ' 1 51
D a r w i n , E ra s m u s , 1 2 , 1 3
Deer, 1 34
Demes, 4 5 , 76
Dem ocrit o n , 7
D e r m optero n s , 1 3 5
Devon i a n , 5 5 , 1 1 2 , 1 1 4 ,
1 1 5, 1 1 6, 1 1 7, 1 1 8,
1 1 9, 1 2 1 , 1 22 , 1 24

1 57

de V ri e s , 27
D io t r y m o , 1 2 9

D i c y n od o n t s , 1 2 6
D i n os a u rs , 37, 50, 1 09 ,
1 26
D i p l oid, 57
D ip loverte bron , 95
D i s t r i b u t i o n , g e o g ra p h i c ,
43
D N A , 5 6 , 6 7 , 6 8 , 6 9 , 70,
74
Dobz h o n sky, 8 1
Dog , 1 2 3
Do l p h i n , 1 2 8 , 1 3 3
Dreponospis, 1 1 4
Drepo n i d i d s , 8 8 , 8 9
D r os op h i l a , 2 8 , 2 9 , 8 1
Dryop i t h e c u s , 1 4 2 , 1 43
D u n k loste us, 1 1 5
E a r t h , 9 6 , 9 7 , 1 04 , 1 0 5 ,
1 07
E c h i d n a , 5 3 , 1 30
E c h i n ode r m , 1 1 2 , 1 1 4
Ecosyste m , 3 5
Edentotes, 1 3 5
E g g s , d ev e l o p m e n t of, 57
Elephants, 1 35
E m bryo n i c deve l o p m e n t ,
38
Eobos i l e u s , 1 3 2
Eocene, 1 3 3 , 1 4 1
Eospe m o top t e r i s , 1 2 1
Eryops, 1 1 9
Esch e r i c h ia c o l i , 4 6
Essa y on Pop u la t i o n , 2 0
E u rypte r i d s , 1 1 2 , 1 1 3
E u s t h e n op t e ro n , 95
Evo l u ti o n , 2 3
a n d f u t u re of m a n , 1 5 4
c o u rse of, 1 05
i nd i c a t i o n s of, 3 0
patte r n s o f , 1 0 2
perspec t i v e prov ided
by, 1 53
p r e - o rg a n i c , 1 04
processes of, 5 6 - 1 OJ
role, 1 00
s y n t h e s i s t h eory of, 2 8
Fe r n , 5 , 1 20, 1 2 2
t ree, 1 2 1
Fiji, 43
F i n c h e s , Ga l a p a g o s , 1 9 ,
82, 83, 88

1 58

F i s h , 1 1 4, 1 1 5 , 1 1 6, 1 1 7
F i s s i pe d s , 1 3 3
F l ower i n g p l a n t s , 5, 1 2 2 ,
1 23, 1 32
F o ra m i n i fe r a , 4 8
Fossi I s , 48- 5 3 , 86
living, 53
o l de s t , 1 1 0
rec o r d , 5 4
types of, 1 08 , 1 09
F rog , 1 24
F u c us, 1 20
F u n g i , 5, 1 20
G a l a pa g o s I s l a n d s , 1 8 ,
1 9, 43, 82
G a m etes, 5 7 , 63 , 7 5
G e n e s , 5 6 , 60, 6 3 , 6 6 ,
67, 72, 74
Genes i s , Book of, 6
G e n e t i c d r i f t , 75
G e n o type, 6 1 , 65, 7 3
Geo l o g i c t i m e s c a l e ,
98-99
Gi bbon, 1 42 , 1 43
G i n kg o , 5 3 , 1 2 1 , 1 2 3
G l ossople r i d s , 1 20
G lyptoda n , 1 7, 1 3 4
G o r i l l a , 1 4 2 , 1 43
G rad u a l i s t , 1 4
G y m n osperm s , 1 2 3
H a p l o i d , 57
H a rd y - We i n berg
p r i n c i p l e , 65
H a wa i i , 8 8
He l i u m , 97, 1 0 5
Helix, 1 1 9
Hemop h i l ia , 66
Henderson I s land, 43
H e r b i vores, 1 3 3
H e reford c a tt l e , 74
H e spe rorn i s , 1 2 9
Heterozyg o u s , 6 1
H o m o l d o t h ere, 1 3 1
H o m i n oi d s , 1 4 2
H o m o e r e c t u s , 1 4 5 , 1 46 ,
1 47 , 1 50
sap iens, 1 45 , 1 4 6 , 1 47
H o m o l o g o u s s t r u c t u res,
39
H o m ozyg o u s , 6 1
H o r s e s , 5 1 , 1 00 , 1 0 1 ,
1 3 1 , 1 34
H u tt o n , J a m e s , 1 5

H u x ley, T . H . , 2 3 , 1 5 2
H y d rog e n , 3 5 , 1 04 , 1 0 5 ,
1 06
H y d rosp here, 1 0 7
Hyena, 1 33
Hyro c o i d s , 1 3 3
H yrocot h e r i u m , 5 1
l ch t h yorn i s , 1 2 9

I c h thyosaur, 1 26, 1 28
l c h t h yostego, 9 5

l c h t h yosteg i d s , 9 5 , 1 24
I cterus, 3 6

I g uana, 1 8
I n g ro m , Ver n o n , 9 3
I n heritonce, 56
lows of, 2 6 , 6 2
mechan i s m , 27, 66
patte r n s o f , 6 0
p r o b a b i l i t i es , 6 3 , 6 4
varia bi l ity, 72
I n sectiv ores, 1 3 5 , 1 4 1
I n s ects, 1 1 8
I s l a n d s p ec i e s , 4 3
I solation, 1 39
genetic, 76
geog rap h i c , 1 9 , 7 6
J e l l yf i s h , 1 1 0 , 1 1 2 , 1 1 3
J u p i te r , 1 05
J u ra s s i c , 5 2
K a n g a ro o , 1 3 1
Kett l e we l l , H . B . D . , 8 5
K u r te n , B . , 8 6
L a byri n t h od o n t , 9 5 , 1 1 7 ,
1 24
L o c k , D a v i d , 83
Logom orphs, 1 3 5
Lam arck, 1 2 , 1 3
L oncelets, 1 1 4
L ea d , 97
l eo d -.t h or i u m , 9 7
L e m u r, 1 40 , 1 4 1
L e p i d od e n d r o n s , 1 20
lepos p o n d y l e s , 1 1 7
L i fe , a b u n d a nce of, 4 , 5
adapta t i o n s , 3 3
b i oc h e m i c a l
s i m i l o r i ti e s , 40
ce l l s t r u c t u re , 3 2
c l assification o f , 1 0
c o n t i n u i ty of, 3 1 , 44
dev e l o p m en t o f , 6

d i ve rs i ty of, 5, 1 2
e v o l u t i on o f , 1 07
g rowt h , 3 2
h i story, 1 4
i n terdepe n d e n c e , 3 5
m e t a bo l i s m , 3 2
n a t u re of, 3 4
on land, 1 1 8
o r i g i n of, 8 , 1 8 , 1 06
protop l a s m , 3 2
reprod u c t i o n , 3 2
rese m b l o n ce , 3 6 , 3 8
sero l og i c a l s i m i l a r i t i e s ,
41
s i m i l a r i t i es o f , 3 6 , 3 8
u n ity o f , 3 2
variation of, 2 4 , 44,
4 5 , 7 2 , 7 3 , 75
l i g h t , s pectros c o p i c o n o l y s i s of, 1 05
L i n g u la , 1 00
l i n n o e u s , 1 0, 1 1
l i n n o e o n S o c i ety, 2 2
l i ptopte r n , 1 3 1
l i verwort, 1 20
l izards, 1 27
l o b ef i n , 9 5 , 1 1 7 , 1 24
l o be l i a , 8 8 , 8 9
l u ngfish, 1 1 6, 1 1 7
lycopod , 1 2 1 , 1 2 2
Lye l l , C h a r l es , 1 5 , 1 6
lyre b i rd , s u p e r b , 9 1
Ma l a r i a , 9 2 , 9 3
Mo l t h u s , R o b e r t , 2 0 , 2 1
Mam m a l s , 1 1 9 , 1 2 8 , 1 30 ,
1 3 1 ' 1 3 2 , 1 3 3 , 1 34 ,
1 3 5 , 1 3 6 , 1 37 , 1 3 8 ,
1 3 9 , 1 40 , 1 4 1 , 1 4 2 ,
1 43
Ma m m ot h , woo l l y , 1 0 8 ,
1 34
Ma n , 1 40 , 1 4 1 , 1 4 2 , 1 44 ,
1 4 5 , 1 46 , 1 47 , 1 4 8 ,
1 49 , 1 50 , 1 54
Marm oset, 1 4 1
Marsh a l l , l . C . , 1 07
Mars u p i a l s , 1 3 1 , 1 3 9
Medus ion , 1 1 0
Mego t h e r i u m , 1 3 4
Meios i s , 5 9 , 7 3
Me l a n i s m , 4 7 , 8 5
Mend e l , G re g o r , 2 6 , 2 7 ,
2 8 , 2 9 ' 5 6 , 60, 6 6
Meson y x , 1 3 2

Mesoz o i c , 1 1 6 , 1 1 8 , 1 2 2 ,
1 2 3 , 1 24 , 1 2 5 , 1 2 6 ,
1 27 , 1 30 , 1 3 2
Meta b o l i s m , 3 3
Meteo r i t e s , 1 0 5
Met h a n e , 1 0 5 , 1 06
Michelangelo, 6
M i g ra t i o n , 77
M i l l e r , Sta n l ey, 1 06
M i m i c ry , 90
M i ocene, 1 3 3 , 1 4 1 , 1 43 ,
1 44
" M i s s i n g l i n k s , " 5 2 , 94
Mitosis, 58
Mo l l u s k s , 5
M o n key, 1 40 , 1 4 1 , 1 4 2
M o n o t r e m e s , 5 3 , 1 30
M o rg a n , T. H . , 2 8
M o rp h o l og y , 3 8
Mososo u rs , 1 2 7
Mos s , 5 , 1 20
Mot h , peppered , 4 7 , 8 4 ,
85
M u ta t i o n , 2 7 , 2 8 , 4 6 , 7 2 ,
7 4 , 7 9 , 1 03
Myolino, 4 9
M y lodo n , 1 3 4

Oposs u m , 1 3 1
O ra n g u ta n , 1 43
O rc h i d , s l i pper, 9 0
O r d ov i c i a n , 1 1 2 , 1 1 3
Oreop i t h e c u s , 1 4 2
Orioles, 36
O r n i t h o pods , 1 27
Oste i c h tyes, 1 1 6
Osteolepis, 1 1 7
Ostrocod e r m , 1 1 4
Overpop u l a t i o n , 2 4 , 2 5
Oxyoe n o , 1 3 2
Oxyg e n , 3 5 , 1 05
O z o n e , 1 07
Poch ycep h o lo pec tora lis ,

1 02
Pa l eocene, 1 4 1
Pa l eo l i t h i c c u l tu re , 1 48
Pa l eozo i c , 1 1 3 , 1 1 4 , 1 1 6 ,
1 2 2 , 1 24
Pontodonts, 1 3 5
Porus, 4 5
Paste u r , 9
Peng u i n , 1 1 9 , 1 2 8
Pe n n s y l va n i a n , 1 2 2
Perissoda cty l a , 1 3 5
Perm i a n , 5 3 , 1 1 4 , 1 2 5
N a t u r a l s e l ec t i o n , 2 0 , 2 1 , Pet r i f i ed Forest N o t i o n a l
Pork, 1 23
2 5 , 78, 79, 80, 84,
P h e n o codus, 1 3 2
9 2 , 9 4 , 1 00
P h e n otype, 6 1 , 6 5
N e a n d e rt h a l m a n , 1 47,
P h o l i d oto , 1 3 5
1 49
P h os p h o ro u s , 1 0 5
N eo l i t h i c c u l tu re, 1 4 8 ,
P h otosyn t h es i s , 1 0 6
1 50
Phyla, 5
N e pt u n e , 1 0 5
P i g , 1 34
N e w G u i ne a , 4 3
P i n n i pe d s , 1 3 3
N e w t , 1 24
P i t h ecanth rop u s , 1 46
Newto n , 2 3
P l o ce n t o l s , 1 3 1 , 1 3 5
N i trog e n , 1 05
P l acode r m , 1 1 5 , 1 1 6
N oa h ' s F l oo d , 1 5
P l o c od o n t , 1 26
N o m e n c l a t u re, b i n o m i a l ,
P l a n t s , 1 06 , 1 20
11
n u m be r of s p e c i e s , 5
Noth orctos , 1 3 2 , 1 4 1
o l dest, 1 1 0
N o t h o s o u rs , 1 26
vasc u l ar, 1 2 1
N ot oc h o rd , 1 1 -4
P lasmod i u m , 9 3
N ot o u n g u l otes, 1 3 5
P l ato, 7
P l a ty p u s , 5 3 , 1 30
Obelia, 1 1 1
P l e s i osa u rs , 1 2 6
Olenellus, 1 1 1
P l i oc e n e , 1 43 , 1 44
O l i g oc e n e , 1 4 1
P l i op i t h e c u s , 1 -4 2
On th e Ori g i n of Species,
P n e u m ococcus, 6 7
1 6 , 2 2 , 2 3 , 52, 80,
1 51
Po l y p l o i d y , 74
Ophrys, 90
Porpoise, 1 1 9

1 59

Potassi u m - a rg o n , 9 7
Preca m br i a n , 98, 1 1 0 ,
1 1 1 ' 1 20

P r i m ates, 1 3 5 , 1.40 , 1 4 1 ,

1 4 2 , 1 4 3 , 1 44 , 1 4 5 ,

1 46 , 1 47 ' 1 4 8 , 1 4 9
P r i n c ip les of G e o l o g y ,
1 5, 1 6

Proba b i l i t i e s , 6 3 , 64
Probosc i d e a n s , 1 3 5 , 1 3 7
Procon s u l , 1 4 3

Prosi m ia n s , 1 4 1
Prote i n s , 6 7 , 7 0 , 1 0 7
Proto lep idodendro n , 1 2 1

Protop l a s m , 3 2
Protoz o a n s , 5
Ps i l a p h y t e s , 1 2 0
Psi lopsid, 1 2 1 , 1 2 2
Ptera sp i s , 1 1 4

Pterosa u r s , 1 2 7
Pyrotheres, 1 3 5

Ra b b i t , 1 3 5
Races, 45, 76
Rad i a t i o n , a d a p t i v e , 1 2 8 ,
1 35

Rad i oa c t i v e e l e m e n t s , 9 6 ,
97

Rad i u m , 9 7
R a m ap i t h e c u s , 1 4 3 , 1 4 4

Ray , 1 1 5
Ray, Joh n , 1 0
Rece n t , 1 3 0 , 1 3 1
Reco m b i n a t i o n , g e n e t i c ,
72, 73

Red i , Fra ncesco, 9

Rep rod u c t i o n , sex u a l , 3 3 ,
5 9 , 7 3 , 1 07

Rept i l e , 1 1 7 , 1 1 9 , 1 2 5 ,
1 26

R h i n oceros , 1 3 1 , 1 3 4
R h y n coce p h a l i a n s , 1 2 6
R N A , 67, 70, 7 1
Roden ts, 1 3 5
R u b id i u m -stront i u m , 97
R u s h , sco u r i n g , 1 2 1

Sa l a m a n d e r , 1 2 4
Sa t u r n , 1 0 5
S a u ropods , 1 2 7
Sca l e i n sects , 4 6

1 60

S c o rp i o n , 1 1 8
Sea l , 1 3 3
Sea sq u i rt s , 1 1 4
S eed f e r n s , 1 2 0 , 1 2 3
Selecti on , sexua l , 9 1
S e x , determ i n a t i o n o f , 6 6
Shark, 1 1 5 , 1 28
Sickleb i l ls, 88, 89
S i c k l e - ce l l g e n e , 9 2 , 9 3
Sil urian, 1 1 2, 1 1 4, 1 1 8,
1 19

T h a l l o p h yte s , 1 2 0
T h ecodo n t, 1 2 7
Therapsid, 1 30
T h e r iodont, 53, 1 2 6
T h e ropod s , 1 2 7
T i m e sca l e , g eo l og i c ,
98-99

T i t , g reat, 4 5
T i ta n o t h eres, 1 3 2 , 1 3 6
To ad , 1 2 4
Too l s , 1 4 8
To r to i se s , 1 8 , 1 9

S i re n i a n s , 1 3 5
S ka t e s , 1 1 5
Toxodo n , 1 7
S l oth, 1 34 , 1 3 5
T r a c h e o p h yte s , 1 2 1
Snai l , 1 1 2, 1 1 9
Trans m u t a t i o n of S p e c i e s ,
S n a ke s , 1 2 7
20
Societies, h u m a n , 1 50
Tree h o p pe r , 9 0
Solomon I s lands, 4 3 , 1 02 Triassic, 53 , 1 30
S o u t h e r n - a pe , 1 4 5
Tr i c e rat ops, 5 0
Spassky, 8 1
T r i l o b i te , 1 0 8 , 1 0 9 , 1 1 1 ,
S pec i e s , c h a n g e s i n , 4 6
1 1 2
foss i l , 4 8
T u b u l identes, 1 35
island, 43
T u rt l e , 1 1 9 , 1 2 6
new, 1 03
n u m be r of, 5 5 , 1 0 3
U n g u l a te s , 1 3 3 , 1 3 4
S p e r m , deve l o p m e n t of, 57 U n i f o r m i t a r i a n i s m , 1 5
S p h e n ops i d , 1 2 2
U n i ntatheri u m , 1 3 2, 1 3 6
Spider, 1 1 8
U ra n i u m , 9 7
S p i d e r m o n key, 1 4 1
U ra n u s , 1 0 5
S p i n y a n teater, 5 3 , 1 3 0
U rey, H a ro l d , 1 0 6
Sponge, 1 1 2
U rod e l es , 1 1 7
S p o n t a n e o u s g e n e ra t i o n , 9 Ursus arctos, 8 6
Spr i g g i n a , 1 1 0
spe l a e u s , 8 6
S t a rf i s h , 1 1 3
Stegosa u r s , 1 27
V a sc u l a r p l a n t s , 1 2 1
S te reospo n d y l a , 1 1 7
Ve rte b ra tes , 5 4 , 1 1 9
Stone age, 1 48
" c l asses o f , 5 5
S t ro m a to l i te s , 1 1 0
oldest, 1 1 4
S u b s pec i e s , 4 5
V e s ti g i a l s t r u c t u re s , 3 9
S u l f u r, 1 0 5
S u rv i v a l of t h e f i ttes t , 7 8
W a l l a ce, A l f red R u s se l ,
S u tt o n , Wa l te r , 2 8
1 5 , 2 0 , 2 2 , 43

Ta p i n ocep h a l i d s , 1 2 6
Ta p i r , 4 3 , 1 3 4
Tars i e r s , 1 40 , 1 4 1
Tarsioids, 1 4 1
T a x od o n t , 1 3 1
Taxonomy, 36
T e r t i a r y , 1 00 , 1 3 4 , 1 3 5 ,

Wa l r u s , 1 3 3
Wease l , 1 3 3
Wh a l e , 1 1 9 , 1 33 , 1 35
W h istler, g o lden , 1 0 2
Wolf, 1 3 1
Wood pec k e r s , 4 2
Worm , 1 1 0 , 1 1 2
W r i g h t , Sewa l l , 7 5

1 36, 1 39, 1 4 1
Tex t u laria , 4 8

Zygotes, 57, 65