O -f

* *

mmdmd-& mtR-wllst.
the i m w m h l s

*-

eta^^

Contents

XV

xvi

Port One General %laeonto!ogical Concepts

1 Principles of palaeontology 1.1 I t ~ t r o d ~ ~ c t m r ~ 1 2 Ocuurrcticc nf irlvcrtcbrate fossils 111 Pharlerozoic rocks Hard-part prercrv~tion 'loft-part prcscrvarion 1.3 llivlciotls of itivcrtcbratc p ~ l x o ~ ~ t o l ~ g 'T'axonorny The spcuc5 C O ~ I C C ~ ~ Noi~lenclarurcand ~ d c r ~ t i t i c a t ~ o n s s d specie5 of h T~sonomir~ 111er;rrzhy Usr of st$1t1stac;11n e t l i o d ~ ~

~l~cut>i~\l~~.gy P~I'I~o~c~~Io~ Functiorlnl rrlorpholoq, firowtl~ and form Strnt~gaphy L~tbostmtiqnphv Hioaratqrrphv Chronostra tlgrnphy H~lrl~cyrapli~ !$(\oh, trcatlsrs a t ~ d symposia I~idividual paprrs a n d other refere~lces

2 Evolution and the fossil record 2.1 Introductiori 1.2 Ilanvln. t l ~ c cpecit.5 and tlatural sclcctioil Inllcritanuc a11d the qource of varintiorl W1lt.r~ drwc variation cnmc from? Sig~iifica~~ce of allclcq Mutation 'iprr'id of ~ u ~ ~ t a t i n t l f thrnugh popiilat~otls I<otatluna t d (pecief f o r n ~ a r ~ o o I;ct~ztic.dnft: gene pool5 Molecular p c n c t ~ and evolution ~s Gc~ic rc.gul:ltlon d ~ i r i n s e v c l o p r ~ ~ e ~ l t d 2 3 F o v ~ rrcord dud ~rlodes f evolution l o

vili

Contents

Microcvc>lutir>i~ Allupatnc rpcci,~nnii klctcrorllrnny T c ~ t ~ m~crorvolut~o~ldr)'i pntturnc ng Ar~aly<ir carc h~qtoncc of Co-cvchutron Macrocvoli~tion KFXIP cclcctiori Clngriih c,f Illshcr r,lud ILrtc*\ o f ~ ~ v o l u t i o i ~ , ,~d,~ptive r,~di,~rionq c . x t ~ t ~ c t ~ o t ~ and 2 1 C c r t n p c t ~ t i oand I ~ Feffect\ .. ~~ 7.3 Snn11:m-y of pnlncoutologc,~l cvolutlon throv Rihliognphv B170kh. trc.ltl\cT ~ n r \ynlpow l I n d ~ v ~ d u p lp c n ~ n other rrfcreilc c\ a d

Part
"t

41'
4'

>
I

42
4 ,j

I .I

41
4-r

1 : 1 '
4

31

3l 3l

3:

-.

-1 4 4

3 Major events in the history of life 3.1 Introduction 3.2, Prok;it).ntcs nrtd cuk.~ryorc<
3.3 Enrlicct nirtazomi5 Ediacar.1 f;lun,l: t7vo vic~vpoiut\ T h e tmditroti.~lview Mcdusnids Pc~ln.~tul~cc..~nq A nnclids Fo-ilr nf unknolvn affinit~t.~ Vcndnzoan hypntlic'iir Sn~al~ h c l l y l fxsils. Prr-calnlinnn tr,icc Srls~ilc CJIISL'F thc Cninl~rinr~ nf 't.xplosion of lifc' 1'hycicochciliic;il facton Biclloyical hctors Rinlopcal c ~ i d c t ~ o n mrtazoan rcl~tionsliipq cc 3.4 MAJOT fcanircc o f thc Phancrozoic rrcord Dlrpcrsr fic.~rinno i invcstcbra tc lifc Cliat~gc:t~ q p r c i u ~ in divcr\ity a n d Iiahitat Psol,lcr~~ntic y PaZacozoic f x s i l r car1 M.~rinccvolutiotiaq ca'aun~s Clirrlatic ,ind ~ c a - l c ~ changes ~-l Eutinctinllq Pns~ihlc cauccs of r ~ ~ n ' ;~,stinctinns r E:.rthhound tncch.anisiri~ F.;rmtcrrcstn;ll rnt.ch,~ni~rns LJ~C <>i-tlc>vic~ati (Ash~jlliar~) cxtltlctlon cvctlt 1.di-e Dcvt mi311 (Frdcni,~tl-Famcr~nian) t~ction uxti event L.li-e Pcrmian e x t i n c t ~ n rcvcnt ~ Ldtr Tnas\ic {C~mi;~n-Norial]) cxtinctint~cvcn t

55
3 :

--

4
4

-3: 3:
34

-.

.;r
.?('

5'.
(611

1( l
($1 1

5 5

ft !

{rl
11:

0: 0:
(15

3
i

(.IF

h!:
(I"

I" b
-,

I
-1I

7 7

i. '

71

7.1 7.1 -.
1:

11

7:

71,
71, 77

r,

n
11

7' 7' 7'

Contents

i x

t Two Invertebrate Phyla

4.1 1)31!-lan1Porifcra: spntlgcs

83

4.2 Cla\\ificatinri
-1.3 C~.I\F ~ n i o ~ p o t i g r a D S p i ~ ~ idl ~ n ~ o s p ~ ) i i g c ~ ~r S~l~rospotlycs I:l~.~~tctids Strrunaroporoirls Spl~iiicrozt~ai~s 4.4 C:lais C;ilc;~rca ?... Cl<~i.; Hcsactincllidn 4.(1 Iri(.crtac qcdi.;: Arc-h;lcvc*yath;) Soft pan$,orf~~~iixatirlti e c o l c ) ~ ~ * and I)ictril,utio~r;~nd \tr;lti&mpllic uhe -1.7 (.;colo~~ual i~nportailcei sponges f 4.S Sprli~gcwcfs 5plculnr \ p o n y rwfi C:~lcarcou< hpc>ngc rracfi
l ~ l l ~ l i ~ ~ ~ . l p l ~ ~

trcatircs nt~d sytrlposia lrldiridual pnprrs and othcr rcL.rcncc~

Iintlks,

5 Cnidarians .;,I It~trod~~ctiotl .2 Mgor cl~arnctcristic-c~ i i d ~ ; I C C C 43f I111y113m Cnidilrin i ; C ( 7,; C1.1i.;Hyrlrc>zn;i Orrlcr Hychroirl:~ Orrlcr Hydrortx-~llina i . 4 Clxs Scyphoz0:1 i,i Claw S \ I I ~ ~ I ~ Z O ; I Sul~r.l,~ss Ccriantip.~thana Subclass C)ctc~rr)mllin Sr~l~clasr Zonntlt:~ri~: cornls Ortier l < i ~ y c > < ~ Ordur T ~ h u l a t , ~ OrJur S ~ l ~ r a c t i ~ l i a Cur.11 reefs C;uulogic,il 11sc~ vt~ral5 clf Cnr~~lq colorlics: thc lirnitr nf ztlnntharinn evolution ar
Mir~os ordcn I!iblingr.~ph?; Uotlk~, trcatiscs a ~ i d ryrriporia Ir1d1rid1tnl papcrs nrlA clthcr rcfcrcnuc~ Bryozoans h l lntrod~ict~nn h 2 T n o tuamplrc of l i v ~ n g hrl;c>zt,nn~ Rrl~r>rrl)nr~ km
,57l~!ttl??,?

Conknts

7 Brarhiopods 7.1 I ncrodrir-tlor~

7.2 M o r ~ I l o l ( i ~ ~ Subyln lunl 1x11ync \ ~ u t i c l l ~ L > r l l l c ~ h , l ( r p l ~ o l o ~ y t l ~ r c cgcrlcra of I - " r ~ \ ~ ~ v d t i\ot i~, , d ~ c l ~ ~ ~ i f i ~ a t of ;lrt1~1113tcd ~ K I C ~ I U ~ C ~ J S l 3tlJ 1011 ~ M.Gor L~nttirc.? br,lc I~it,purI~ r ~ o i p l ~ o l o ~ of Etidopu~~ct,itinil p w ~ ~ d r ~ p u n c t n t linm .u1~1 c thcllr '+,ti t,phvlurn 1 ~np~lifi>rrnca
!-it!y~i/<~

Orhcr I i i ~ ~ ~ l ~ f r r r n ~ ~ , ~ h~~bylir C:r.i~~iifi)l.t~~~~ lurn 7.3 C>lltogclly 7.4 Clafeitic.~tic>n 7.5 Evcrlutinii.lrv Iii5tr)i~ 7.h rc nlom .uirI d i c r r i h u u ~ n ~ E Lu l r > h ~ ~inch\ lclu.11 rpcclus t f 1:pit I I ~ I I . Ih r c ~ c I ~ ~ o p o d ~ ~ F:ii:ltrt:~un.ll hr,zt hiitpods !lt.rcllit>pod , ~ s ~ c r n ' t , l ~ 1 1 d ~ c '~o111111~rnltv' l n p \ c<o C)~rinl,ic~.ln p,il,lcc)coir~rr~uillties Srlui 1,lrl p.~lncoco~nillurl~tlc.c I_)crc~ni,~il l~r~cl-uopuclw i ~ l b l a ~ z > a\ l'crrrunn 1 c c f . I F ~ O C I tions ~I Mc\c>7rricb r . ~ c h ~ o p c d nrsuclatiul~s 7.7 Faun;] prrn irlcc\ 7,s S t r ~ t i ~ , r p l l ~ c d ~ L I 13lhliograpIiv Boo'k5. trc.lrirei ~ r \!nrpusrd ~ d I nciib itlu.11 p p C n ,>ndnthcr rr=fvrviiccq

mollusc^ r ~ t n ~ i . ~ r n c no n ly ~ l l i ~ ~ t ~ o t l tr 8.2 C:P.IWl hc.lrloll 8.3 S o n ~ c .~cpcct\ thcll t r ~ o r ~ h o 31ld~yo\%-th of lo Co~lcd <hc.ll r-norpllrrlogv S c p t ~ i cot' ~ i <livlJ ~ thc X . 1 l'rinc~l-r,~l t?r<cll group\ C:lnw I < i v a l ~ i ~
8.1

Contents

xi

203 31 0 300
7 -1 0

273 21 9 230
27 1
-u-

771 --371

224 224 226 726

Ld

37C)

2.10 238 23 1 2.55 2.56

2.1rr

237
Erl~inad~rnts

"I

111tl-odll~tl~ll 'h.2 Cl<~r<ificat~or~ ' j 3 Subylivlu~i~ Ir'cllinl,zn,~ Clnv Erh~unidc,i 4loryhnlogx~o l d I ~ f r ~ ~ ~ btjf'rhrcc gciit.r,b I rtc C~JFFI~~CZI~IO~I Cubclnsr I ' c r i r ~ l i u c r l i ~ ~ ~ n i J c a 5ubclas< C:idaroidc,i cul>clacq E ~ ~ ~ c l i ~ t iJIICIl d ~ . 1 1 1 o r p l i o ~ 0 ~ c h ; 1r1~ ~ t of C ~ I T C ~ ~ ~ Z I O I ~ C u the ~ ~~ a cn tvolu~io~l ccl~it~o~dc 111 Claf+Holuthuroitlc~ (:lnsr Z d r ~ u ~ ~ s t ~ r u ~ d c ~ 'i.1 Sl~ly>h\ j Astcruzt>a lu n Cribt.l.~rfA\terolJm 3ulll-lnfsSomastcrr>ldcn S t ~ l w l C)plllilr~iilcd ~~f 4t,1rtirl1bcdc '1 h~lyilivl~~t~~ (;ri11(wo~1 Cl,lrb Crinoidc;~ hlaiti qrouyr\ of criiioidc I',~larozo~c cnr.oi~ls h l c ~ o z c r ~ c rcc cn t i r ~ n c ~ i d.s : r t at> ~ icul:~tc.\ Er olo~l-~ ) i cl-it'toiti~ F o n l ~ ~ t t oni ii L r1no1~i~l Iimc~lorlc'c " (I 5t11~1111l'1ll~ll ! ~ l \ t O ~ l ) ~ l I~ I:l,l\(cc I)~plr)pc~r~w 1~hor~thifc.r.1: .utd crctnidc

xii

Contents Stnicntr~l chamctenqticr ~ T Qni~'tl1rCc C Clnw~ticntlon Ecolog Cl~v Rlnitnldca Rivcreir~; and functlon of h y d r o ~ p l r c ~ Clnrsiticat~onand c v o l u t i o ~ ~ hlastoids of E c r ~ l o g dixtribu tmn of b l n ~ t o i d ~ 2nd 0.7 Sr~hplivlurnMnn~nlozon, orhrnx~lw a l c i c h n r ~ n t c ~ c 0.3 E\~olutinn ' E ~ r l i c ~ ~t l l i n ~ d ~ n tihlc ~ r d i a t i ~ l l ~ ~~ and ~ n Evnlutinn of the tube fcct Why pcnwmcvi Cnnvcrgcnt cvolutinn J I I ~ i i t ~ m i ~ dfornms~ i ~at Bibliogaphv IJookt, trc.nrl\r< and <\+\.rnposia J~ldividual papcn aiid uther ~ ~ ' f e r r l n c ~ ' ~
10 Graptolites 1il.l \rructi~rc Ordcr Gr.lptnlotdca
S t l r t t ~ y r n pdtinlncrIl [~~~

D~ploqr,~pftts It-pfotl~ern

Order Dcndroidc.3
Drt~du,pnlptlrs

Ijresen-at1011 ~ t stlid? of graptolite~ ~ d Ul t r ~ q t nhirc ;111d c h e ~ n l ~ t of gr~ptol~pcriclern~ ~c ry tc 10.2 C:lar~ific;~non 10.3 Rinlngical affiinl1t1t.q 10.4 Evnlunon Shape oi gnptnlitr thabdn~nmes Iymx~iiial in Lm'ptolo~dt cnti Thecnl qmicturc Cladia Stn~cturc rcriolitid~ of l(7.5 M o r ~ ~ graptolitcr Iivc? d d Pa~qivc dr~fting Autotnoh~l~~ U s c otlmodcls in Intcrprctlng thc mode of life of gaptoloids 10.6 Faunal province? 10.7 Stratignph~cal use E~blioqaphv Ltooks. treatises and svmposia Ind~vidualpapers and other references 11 Arthropods 1 1. I tntroductinn 2 1.2 Cla~5ificatioll and gcncral morphology D l v c m ~of-~ r t l ~ r o p o d ~ hpes Fcaturc~of arthropcld orp~nlsarion

Contents
[

xiii

I .7 'frilobit,~ C;cncr;ll r r ~ o r p l ~ u l o ~ ~ ~
i+1<c7~tr

(IOIIJI~~I!~~~~~

35 1 35 1 352
354 355 357 3 57

Ih=t,l~lcd~ ~ o r p h o l o p trilr>bitu< t of Cut1cIc Cc['hdZotl Glabclla Ccpltal~vsuturcq Hyposto~rlc

Eyt.5

357 359 300

3(>3

Ccphallc fringtc Erjrollrticnt and co~ptarlvc ~mictlircq Tllnras Pyg~Jl~l~rh Appcildagc? TrlIobltc track< anrl tra~ls Walkmg tnnvenrcntr in nrtlrrc-)pad? 13iffcrtiit kindr of tnlohirc trallf 1.1ft. attitudcc. h d b i t ~and c c n l o ~ y F,c dycl~ ,ind ontogcllv CPa+~rfi~a t~on E~olution C;elicral pattcrrl o f c v o l ~ ~ t ~ o ~ l Microcvulu tion Faunal yrovincrs Stranpaphic.~lL I ~ 1 I 4 Phvlurn Chclircr,itn Clav M c m ~ t c ~ t : ~ a t a Suhcla?~ Xtphost1r.1 Subcla5s E t ~ y p t c r i d ~ 1 1.5 L'l~ylurn Crust.xcc;i U~L~ho~rayhy Elookr, trcaciqcc dtld ryir~poria I~~divirlual pctpum .;~nd otllcr rcfcscnccs
12 Exceptional faunas; ichnology 12.1 Introduction f 2.1 Dtlrgcn Shdlc f;iunLl

365 3 60 3 07 307 370 372 372 374 377 3x0 3x2 3x2 3x3

380
RX7
3x8

3HX 388

392 39 7 400 400 4C)(1

Artluopodr Lobopotis CJtIlcr ~nvcrtc\,r.~tcc Slgnific~ncc. thc Htrrgc\r Shnlc h u n ~ r of Ecology G c o ~ r ~ p h i c dlrtnbz~tlon al

Diversity
I'cr;lstcrlcc

12.3Uppur Ca~nhrlartof coutl~urnSu-cdcn 17.1Hun~riickcchicfcrf,lnr~n 12.5 hfazon Crcck L ~ ~ i n a

12.6 Sr)l~>lloftm I~thogrnphic lirncctonc. Hnvann

xiv

Contents

12.7 I c h n o l n p Clnu~ifiratinno f trace cocrils Morphologic.11 m d prcccnratlonal rlascitication 13chnvto~~r.~l cI.~c~ifirntioi~ Phylngcilcric cla5sific ~tiola Urcq of ~ c l l n o l r , g C;crlinicrit.ly ctlvironlllent Crratigrapl~y Fossil l>chav~our Fiil~lio~~pli~~ Rook<. truatl'icc arid synlposi;i (e.rcrptiona1 fauna<) Inclividu,~lpapers .itrd othcs rrferenccr (cxccptioiinl Rook<,trc,lncc< ;ind qyrnposiCl(Iclluolo~y) Ind~vldu,~l p c r r p~ (ichrtology)
Systematic index

fmnnk)

1 hc f i r liqhcd I
11c\3- 0 1

h c t all1 tcxt for

General index

!'car 01 Cbnptc

tlOll 11;1

chnptc ,I zrra 4 t of t l ~ c i
for lot)
[""VC

'1lldtOll

Kcy rhc re> tlicy n l AF L

I~.l&.ll,.uc

.IHCC 11

C:olln
,1rlv1cc

shot1ld tdkcn r

~ I ~ C S CC

SP1cldo In1111 ( hcl p ~ illi1ilt'r

~~)r(o 1 w tl~,~~ fbr all

' l F c fin1 rllr~c. cditiclns o f tl~ictcxtbook wcru

pub- ~1r.c-11 a t every stage. H.id Cecll~a i~ndermkcn me 11or >(,,Ib l i F L ' a t O ~ { l r l ~ l 1002, <llK4 I tr11st tll<lt tllis t h r vast job of reburldulg o u r tench~tlpc o l l c c r ~ o n ~ !l ~i 1 IY nlic.. ~iccc<~imtc.d so malty ad\:arlcCs both i r ~ alld prepaxing new coilfie ~r~aterials,\vl-ould rlcvcr hy ,t .irl~Itl1r.o~. tvlll rct.~irlits tiunctioll as a course 11'1ve had the tline to cornpletc chic book bcforc t h t ,r filr F ~ L I C I I ' I I ~ T of' p d l a r o n t ~ l ~ ~ y thcir sccar~d dsadl~ne. fro111 ,lr t~~~\v,rrii\. 1 I~a\.c1n:ulc \ubstanti=~l h a i l ~ c stu c To Keisi~l Moral) of Unirenit). Collcgc. G~lwny ~ , i ~ w n 7, N ,~rrcl I?. ,arid thc Trncc Fo.iqilq rec- I extend 11. yratuii~lthanks for her two beautiful 1-3, 1) p i i r l t i n ~ thc Pnrt Title pngss. (or 112 t ~ > \ bwn trmsfcrrcd kotrl C I ~ ; ~ p t It rt che last o 1 wish to tl~3rlk 3150 nly editol-ial colleag~r<. .:1~tt.r. 11 tlic. ilthrr c11~1ptc.n: ,2 Iwvc bccn revised to Jane PEo\vman, \c.lio have p e r 1 jc.ltcr ivr Ic<';crcstcnt. 1 Il;uvc scdra\vn ; ~ l l o i half L)r latl Francis ~ r l d ~t rl:c i!l~~str;rtit~ri.: a ~ii~gi.rrlnrly i ~ , ~ ~occupation all posriblc asslstailcc. From Inn came the quggestlon, co iii~l wl~ich we l ~ a v c follo\vcd up, that the new c c i ~ r ~ o n !;lr loll? \vIotcr c\,c.iiinF, ;uld I hope that thcqc \\ill 1 3 ;'ruv~: o'l",>l~rt: 11rlp1ng<ttnlcnt.; tu ~ ~ n d e r s r a n d 111 the ~110~ild c c011~1cd 1%-ith J CII-KOM o f Fowl Iinagcr. This; has bccn urldertaken In corl,ii~i~ct~rr~l .HI Itclnl\ a i l ~ i c r n ~ i ~ r o o f fvlssil i~~vcstrbmres. t lo~~ Hiftory Murcum, Lorldon, and tcr Kc\r ~ Y C V . I ~ F i t ) brdcl ;I[ thrir first mcntion in with thc N a t u r ~ l appp.rr :!I:.t ; . ~ t . T l l ~ r ~ZIC) \puc~Sicirldux for thctn. hut Nvnnan McLeod and Paul Taylor of that ~~lstltntlon 15 I oxvt ,I g e n t deal. I am hkcwi~cpntefiil to L)av~d I ! I V \ . I ~ ~ C , I IT ~ l i c I I Gc~icr~il ITILL-x. Hicks and Eve Darntlth for thclr i r l r t ~ c u l n ~ ~ r copy4r !)l.iL3rc. 1 \\:isll to th,lnk ,111 n l y fncnds, coledit~ng pmofreadlrlg, rcqxctivcly. and IL, I!:IIL~< I I ~ L ? " l:~tnily~v!icjIIAIT y1~-11 ~ L I L - I I mist111~O n a wet Novcmbcr day, over 50 yca1-r ago. I \t,.rt . u ~ (:ji yi-~.p:~ri~q IIL-U. c ~ l ~ t i o n . * tllt~ L>cI.c~ Hrigps, r :o~:I! St,rrltton :~rld Ii;~chcl\T!t~nd p v t cxctllcnt take11 iiito a ttlureutrl ill my 113tivr N C W C ~ S ~ ~ P - I I ~ O ~ IVI~-;> . l r tl~c hrytitlil~gc>t'tlic p r o j ~ c r;IS to IIOW I T y n c , to cscape from the min. 'rhcrc. in a dusv &rr ,111Iil prr~t-chcd n-it11 thc ti-~urtlj cditiotl, and r havu casc wort two g s n t arnniniai teq (probably 7'i!r111irrs nri t l ( \ . ~ r t i~lo<t F their s u g ~ c ~ t i o n ~ l (I AS ;11\\7\1:1y~. g i q r r t t f t ~ l i j ) .and I wns given to itnderctat~dthat they , a rrlcrc. ~c~llcaquc~, tr>gctht,r u.itl1 IJ;~vid Harper, Pcrur had lrvud iirdtioris of y e d s J ~ O Thcv cvc~tctl hccihl~clLl~\i~. Iligly, Liz H.arpcr, Dick JctT'i.rics. r ~ a t ~ o t i S~~s;aii which has c n n t ~ n u e d ~11inE now, and I all1 still glad to cscapc to t l . 1 ~hills on a f r u h sutrlrncr mom]l L g t - m ~ : ( ' ( ~ 1 ~ A l ~ t l t . h ~ * ~ r h ~ ~ i ~ I Oorhcn Ilavc . L ~ F ; I ~ i~ I ~ o back I!I,\~<>L! 111i- ,9r ,111 <t;lKL%. Gl13<l\Ll 11;1vc 111:~lly Otllcrs, tL30 Irlg, to sudrch for fossils, 2nd tllen t hnng t h c ~ n tmok !1:1111~~1011~ 111~11tioll. my 0 1 ~ 1 1 C~lLtlltry ;~ild to thc laboratory for study. 1 Iiope t h ~ t h i ~ to 111 will be o f value to any 5tudcnt wislul~zto cxplorc ~ltrl>ail, 1 1 1 ~ IIIJ tI~.~tik\ duc t o all. :Irr oT ~ I \ ~ r > ~ ~ likc. .lhovC 311, r o rccord r r y hc;jstfclt coir~ctl~ing the ~ ~ C I I T I C S S A I I ~ C I I V C I T I ~ of a~jcierit lrl scirntific ri~.~:~h< trut rid st:ll\~nrrfricnd Ct-cilia Tnylor, lifc, a t ~ do f the rncthods avallablc for ~ t s r r > Irly 0 ,111 tlw \ u p ~ ~ o.und l~clpfl~l rt ~uq;crtinttr ~ h c 112s study. If ro, 1 will havc achlcvcd \\,hnt 1 srt orit 2 do. ,
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T h c Nnt~ir;llH15tury Mtr~rtirrlLc>nd(>n dnd Dr E.N.K. Clnrkcon havc uollahoratcd oil thc d c v c l o p m r ~ ~ t crf thl\ ~ n i p o r t a trlcw initlatlvc In pnla~.ontology ttcnc11111g. ~f :I l r l c r q J ; ~ ~O IiIl L'IL K O I I IC .I carrrbincci Itnage I~bmn;m d database, c o r l t ~ i i i ~ n g ~ < I recnrdr fur 1000 k r y fowl gciicra. E ~ c h rcroi-(1 iotltnln.; .I hct of i l l l d g ~ c ,;i11~1itlform~rion n stratrgrmphic rangc, hard-part rrunrmlog. o pnlacoorolnt;.v, p ~ ~ l ~ c o h ~ o ~ tcrc.o The~ ~maycshavc bren captured 1ivng thc Natural Hirtow ~ ~r ~pl~~, Muscunl'r l ~ i g rc\ol~itioi~ l~ I'AI.AEC>VlSION diptal lmnpng cvcte~n. atld the data arid It~iagtg rcude 111 tlic Gvrtlt~,Srrnt cint~barc rn;lilnpcr. Thi\ qwtctn ~ l l o w r u w r gc,at flcxlhillt): in di~playing the dat;l. For example, users cnn \imply broww frol~i rccnrd t o rcrorcl; pull lip fnsrllc frortr a taxorlonlic indru; o r sclcct and sort record< h gt'ogr3pl11~ I I ~ C . lift' ! U linhit, ~ t r ~ t l p n p h range. or hy ,I v.~ricty other cntcaln. l k ~ i i g c ic of cl~arts and p a l n c o h i a g c o ~ ~ ~ iniaps~+Ire ~~l ic pvrr \tl~crcappropnntc. and thc uwr ran print records. or cxport tlleill illto other ;~pplrcat~o~rs. Wltllin each rrf rhr Ir1,ljor gn>i1p rlicrc ,Ire a 11u111berof labcllcd irnngcc and dldgrarns to allow rhc 11cv1 to bccolnc fnm~li.~r ke!. rncvphnlo~icaltcrnls. n,ltll .'llitrvt;(;)scil< (111 UU-KC_S:\I tn.1). hc urcd as A I I a d ~ u n c t 111vfbrrrhratc to Prlfoeot~rth,yy, r a? A qtn~~dalorlc o prntluct. T a v n 1 1 3 been ~clcctrtl tlicir rclcv~t~ct. Earth qcicncc tcnching \vorldwidc. ~ ~ for to

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Concepts

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Amixeras cf. hartmanni (Oppel], an assembla e o immature ammonites from the Lower Jurassic of Black Ven, Charmwh, England F [ h e r Sinemurion). These specimens were pro&bly ca~strophicall~ buried, since he soft parts must hare been in when + e ,, died prwenting sediment penetrofing the chambers. Po~nf~ng Roisin Moron; original specimen in the James Mitchell Museum, by University Callege, Galway, Ireland.

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of nur u~lrlerstandl~lg t l ~ c11istc)i-y o f lifr-, of pcrl1.1p~ rno5t significant of all C V C I I ~ F ID(>^ ~ I : I C C tlic about 343 Ma ago nt about the Gcprlnlliq ot' t h c ~ ~ I I I .(l l ~ l l t )~i11111ot1 , vc.lrc . i g ~h c Earth carrlr inti) C:nmbrlan Pcriod, For nt tlliq sta.yc tlrcrc \\:IF A q ~ c i t .II~:,yrrllh~lllt fi>rriijnqf r r ~ i .I lionrlcnr~ng ~ dlrc of den pmllfcration oidifl?sent kiilds nf lllarlttc ii~\vcrsl I I L lr.,. ~ l l t r t,iiid g;i\. I 111~11 <lowly ri>t;lteci rouild \ t r b m t c ~ . 13un11g rh~scritical period r l ~ cpriiicipnl I, 51r1i 1 , I r y r p~rt~c.lc<, ~ ~ l ~ i t f0~111ed ~~r~vcrrchrnte~ 3 wurc c5tabl1rht.d, and thcy thrn 0s ~ ~ t i ~ ~ ~ ~ lquups Il.nt~aI?II, ntbl~l.ir dirt, .qttcd nr tlle~t.roll~dedthcy drvcrq~fiedand cxpa~ldcd.Somt. of tl~c'iuorgL~nisrnc I ~:c.rt.,l, rrttu;rll\ ionnlng rhc pl~ncrg c\ ncqnircd hard rhells and wcrc cap~hlt. buns focof Ot' ,111 rllt. itil~cplnl~ctqIn thc Sal.~r Syktrnl onlv slllzrd, and only becausc of this can t h c r r hc any 1. rth. ,J< lj: ,IF k kll0~77,~ ~ p p O r t. S v d n c c d I l k , d clinncc of ~itldentnild~tlg 11lrto1-y of-invertcbratu thc .I,l$.i;h ,ir ;hc tmic i ~Lvntlng ~nlicli lntcirqt ha< f Ilk. ':]I ~ t l t * r . ~ hv l thc diccav~.n of orgrlr-~c n ~ t c r i a l tn ~ T h c stmtitied ~cdlli~ciltary racks laid drrlril ~lixcc 11 luyirtm'5 rntcllitc Eurrip;~. ir. ho\vcvcs. a qtr~h- t h e enrly C a ~ n b n a i ~ , It n11d h i l t u p througl~clut thc t l i t tli i t ]ifr ~ c l t l I:artli I ~ c g , ~ n c.irly ~ t ~ d c c d ,nbllc>leof P!lanrrozo~c tltrle. nrc dictinpirchccl 1s). .I \.cry 1rh111tlic t i n t 30";I O F tlw ~ J , I I T C ~ * F l~i~ltm-y, Tl~ert. r i c h l ~ c r i t a ~ c the fossil rcunmnq of thu invcstcof c r ~ * r i ~ , l l{n>\ < \ I T T I ~ ~ O .~ ~ I I I \ I I I(<l > ~ c t c rIjI ~ ~ L , I L 'h111rE~rntef that rvolvcd thro~~g!r succcssivc Ii~qtnncnl ,r~.:.tl ,rlc,rc', or L-ynt1ol7acrcn.1)t ? rock? 341 It 1 ,Ma pcnndq; their ~ t l i d vI$ the dornail~of in\rcrtchr;~rc i .1 ,rl l~ir. prcwnl,~l~lv onpnntrd hciorc cl1t.n. pal.wnntnlo~?; nlid rhe qul?iect of thi\ book. Tt.c\: rrnit~lc form< r, !iir hcc111 ~o hnvc dom~rl.~ted f itcrlth c , thc 11c~ut2ll01) Mn. nnd evolut~nll Lr t h . ~ t tllrlt, \v,~c vcn slozv. Nr\~enlieless, tllc rr\cib,ictc.n,~ atld phato~y~ltlrcric , ~ c t e r ~ a b werc I [ ,r~u!rlcr~tnl ch.111y111~ 111 thc cnvirniitllcllt. fur they #:I: jltf LK!grn i i ~ t o .Itrnosphcrc t11.1t war previ,irl LI~vtjiil ~ t \ L I dint nr~imallife eventually , IIIIC y~lv.~hl~. hlly \vll~.~r qnlilc. of thc cnrly Irvitly lbeaii<s o t ' t h i ~ l r t l ? llarl rr,,lrhr.d n high Icl~rl nt' phy51oluqlcal diiri F o ~ v iiivcrtcbratcs occur it1 m a n y kinds ofredimenl y r q l ~ l r t ~ t i i - 1 -u r q n t i ~ ~ . l t ~ o i ~ (,117ci Inort part~c~iPnslv c.lry rock depo~itcd 111 the senr driritlg the ( I I L ~ I +r\u,~l rcprociuctioi~nnpr,'~tcd) w.lr the rsjte Ph:rnt.roroic. The?? In,ly be very ablitldnt~t in (1 r'rollrt~r>rl,ln,cl~,ingc,.~ct-clcratrd, and with it all lirnc<tt>r~e\. FIIJICS., ~ it~rc)nern r ~ d~-nucf~tot~cs o n l but 11 I I I I ~ L ' ~ . of IIC'W ~ c ) ~ + I \ > II ~ 'I~~~I C F I upened up to rlic WIIOIC nrr not cor.larno11 m s a i i d ~ t a ~ ~ e s . c ivlv:~~. Thi5 \v.ls n n t 1ir1t11coiiip~rativclv ~ t e Scdimcnt.iry ironcrorlc< iilny t~avc rich fo\sil l~ir. l 11 :~rl~1~~gc,~l t i d tl~crt* hissnn. a ,we 110 tbrcil ~nltnals rern;~ins. Occnsionally they nre f i ~ ~ m dFonw conme 111 IIO>L\III L ~ Y ~ L ~ I I I I P t)ldcr t11aii s h o ~ ~ t MA. mcks F U C ~ F L~cy%vnc-ker I I>~\ 7tbO A nr~dcvcn coirglo1nerntr5. A:<,:!,* ' to 2 . 1 ~ . tllt*cr ,II-C all I I I V L = ~ C C ~ ~ ; I ~ C nrdmal? T h c ~ t n t co f presrn.,ition of fos~~lf v,lnr\ ~ r r a t l y , IhIL.!~, h,~c.Lhrlnc~.1 of thctr~arc i t i . i n ~ ~tlicrt' I F dcpcnding 011 t h r ~ t n i r t l i r c A 1 s: a11d cornpn<itiott of thc
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oripn;ll 4irll, tllr nature ,irlA grJin ~ i ? c f t h c c i i d i ~ ~ -tlnnr durinp rhcw crncioli periods \vun~lJprol~:~l~lv o nr Irilut,~tiur~ or] i n s s u d ~ n ~ r ~ ~clicr~~iial tlir t , co~lrllttorif ~ttl ~ c . ti111-r of br tr.in~yort.c~d d c ~ t r t y c d .1not21~r thc n d c q n n q of thc fnwrl tr.rt>rii. ~ t ' d i ~ l ~ r n t .~~i tt d ~ l S~~ I ~ W C ~ I I p r I ~~ c r \ ot'diat11c l CI~ ~lf~ O n thc rtth~-rII,~TIO. w ~ r n c . rn.lrltic i n ~ c r t r b r ~ l t c c g ~ n c s i s (clrcii~lial .rilkl plivsir.11 I-hnilpei) t3k11lg foiii~d ttl ccrr.ltn rc~cbtypes 11,ivc. i ~ r c r ipr~-\c.n~t-il pl,icr 111 the rock ~ t t r -d c p o ~ ~ t r n n . r dctsil, \u t F ~ ~t 1% porrit Thr qtuilv o f tlir procrcscs I e b ~ d r ~ l shrriliznt~nn nhniidnt~tly.in{[ in cxcluiu~tc tc, rhcis b i u l o p li-r~~-rit l l c ~ r 1, kno\vri JF tfipllonnrny. I n n l w t s:wc only t l ~ c hlc tn )nF~.r inuch ~ h o u t U -CV U h ~ t dpart\ uf t t w ~ l,~rltrn,~F~r c , ~ pr~-rcn,c.d,n ~ ~ d rci-n.~inr. M,IIIVof tlir ~ ) ~ ' ~ ~ - ~ T C~ ' ~CC ~ II ccwnc I ~ Tnr clr uilty t l ~ c l ~ c1 0 b r f0~~1117td. , ra1plrll'rlirl,~IIF 13(1n11,111y .I prr- from 11n~r.ctc~nc.c k i - ~ ~ r ~ rctlirncllt\ 1vit11 a high (Fig. I . I) ttlc o n g ~ n , ~ ] rerlurcltr. rhc cofi-llodrrd r l c r ~ ~ in n rllc EILIIJJ. r.11cnrcous c-(>ntr.nt.I r l ~ h c w ~ t~ rhcll\ nlnv h c sct.iinctl in 1 1 1 ~ . f i w ~ m t c I J I T ~ rhurc irlrni, \ ~ 1 t t i tlnu or$anli slirll<. d ~ d Trot calc,l~-couc i i n n n ~ l runJli.e d~.~gc+rlc.\i\ hcr~cc l~ ,111d hnvc lcfr littlc with rclntivcly littlc altcrntioei, dcpcnciing Ilprln thc n-itliirr t o r nrt cvitllr~lst. ~ rllc.11r f c\l<tr.nic othcr than rcc.:rrilq chcn.ricnl c n ~ ~ d i t i o r ~ < die wriirnciit , ~ thc tlrnc . ~ .~ficr. of t h r ~ ncrivltv 111 tlrc f i l r r l l o(tr,li.c fo~<ik. r What \te..c 0f-~2~].m$inor1n d A < c d ~ t r ~ coftcii concirt\ (h'roriipui~cntc nt cjcnvcd ran scc In t h r rock< i c t l ~ c s c h r ~ a n.lrrn\v hntd onlv cnvirnnmcnrs, ,111cl ivheri dl1 o l tllcqr. i n .I nlir~lr\pr.ctn~mnf the o r ~ ~ t i i < i that \ ~ ~ c r cfrntn v n r i i , ~ ~ ~ n\ o ric.,~d 'irdlonce Ilviiig; nnlv r.c.nJI-.irclv have thcrc h{-crl f i r z l i l r l ii~cludingdcc,~ving r g ' a ~ ~ i s ~ r ~ \ , c11cIlq ~ 1 1 d t). tllc. hcd~ cullrditung ~l111c ,IEI i)r tlic w$r-l-rnrlicd clr- ~ r ~ r i i t i pornclcs, :lrc ;hrt>wn tc>g~-thcr c!~eiilical 111~11tr1t1 t l ~ c t;nun,r tvr-ll. 'l'llr.\c ~ r rrninc.n~cly ? cig- halamcc i r uilrtnblc. T h r \cdirncrrt 1i1111 bc in r t w l n ~ cal cquil~briuni(>lily.iftcr dlngcnctic pl1!.cict7clrciilii ~ i f t u . i r ~itb r pn l,~contc\lr*,q. Il.~vc y T h c olclc<t qurh f ~ i i j I~ , ~ Llse I'l-cc~rrthnan n=c. c;ll altcr-,~tioi~c t:tkcn pl.~cc.Tlic\c t r ~ . ~~iivolvr \ ciF .ind tlic ~~i-o\\~tli 11cw ~ n i i i t . r ~ l ~ of totnc 01 . . old, .and 1% (7tir nnlx rccord o f anitnal sccry~talli7,1t1nti i1 M >TI lifc b c f n r c t l ~ c C:niiil~ri,in.4ii.citIlcr ~ l i c h 'xvindow.l.'i \ (authigencsis) ns \vc11 d'i C C I T I C H ~ . I ~ ~ T ,iricI ~ 0 1 1 1 kiio\vn rn Middle Ca~nPin,in i-(ir-k5 holn Rrrticli pactlon of tllc rock ilithification), and rl11nn# . ~ n c o C01trinhi.1. \\.11~-rc In nr(ditlot~ tn hic nonnnlly onc nf tlitrr proccwcc tllc fowrl~I I I , I ~ hc. ,~ltvrril r cxprctt-d trilc~bitc~ ~ ' ~ . I c ~ I I o ~ c ~ C ~ F 1s .I Lmeat dcctroycd. Sliullc zhnr nrc nriqin;~ll\ of' c.ilcitc prr.it,d tlicrc Figure I tr f~ l railer o f <ot't-h~>~l~r.cl dii~i-chtlFcd ~nlnt;ll\ - scrvc hcst; ar;l~nnitc n It';$ ~ t d 3 1 n n ~ ot' C . I ~ C I I I I I I and she11 was ~r I IivirtG orghrnsrnk (c.g. spun yes. \vcrrirlq. jdl\-ti\h, ~mnllshrimp-like crr.1- c n r b o n , ~ ~c r c t c dn cc.rt,~in ~nally of ~ i ~~l turer . ~ n ~ l .ulitri:~l< of qtiitc ~ i l ~ k n r ) ~ v n nFtii1itic.c - coral<) and i~ ~ f t t ' r l~ ~ n ~ ' r t 3 l lt43z calc~trrllir~iig genetic or ,I\V:IY inlr~plctclr.. <hellpre whizli ,Ire the o l l k tracc of n rllvcrrc Kilt~iid \vhirh dlngciic~i< d i ~ o l v c d core, (h) Cnlcarcnn~ 5kclcton~ prcqcnbcd in rrlorrn SAIIL?Y or ~vnul(l~ i t l l r r \ \ ' ~L~ tc .L J L ~ I ~ C ~ 1iii!i~101vi1(C'linpter 11). {I) iron.ib ~ Therr ,Ire ~niitlar'windo~\~'i' crtlicr luz-rlc m rhc s i l rcdimcnt(: r11ay clis\olvc .~Ttcrt l ~ crcdilncnt 11.n at liardcned nr during n~catlici-i~1g t l ~ crock lorig ol' gc@loqc,~l i . > l ~ l l 1 1 ~11ketviw lll~lll2lTl~rill~. ~ 1. casts) T h r tils\ll rrrr~rtF ,i\ .i gu~Ic. thc c v o l u t i ~ n 1.r. rn of nftcr its indurntion. Moulds (oitcn inlcc.~ll~d the ord 111Ri' JIICICIII I I ~ L . . ~tlqlit'stit>n~tL-l~ liiiitted, patchy and o f thc eurc.rnnl ~ n intCt-tlalcurt:+cch\c)f thc f i > ~ ~ i l . -+-- I . t11c i ~ l c o n ~ p l r tU\u,~lltc>nlu rllc hard-<lirllcrl tlcmcnt< hc Icft. .liirl if thc wdinlcnt is tint c.t~c>uph~ dctail~ r. chow rridy b~ V C ~~O C \ S ( ~ r l c ~ J C ' ~ ~ ~ f C ~~ J. I O o C 111 tlic l>ior.~c i ~ , , ~ i -fti i l r ~ itrncc inqctlc) nsc pre~cn,cd. t l i e ~ c ( arc clcccribcrl in ccctirrn and thC h i v l , ~ c s r r ~ ~ b l .prcrciit in t h t rnck nlny tht* rturly of cucli ~-rlould\ yc< If llavc tlccrl ~ T J I I F ~ O I . ~ C C C ~ I I ~C ! I E ~ J I ) L - C . nbradcd. dnm- 7.2. with rcGrct~cc rt> h r ~ ~ l ~ i c ~ p r > c l \.I. fowl LOI cncloccs ,111 c ~ r i ~ i n . ~ 11ollou~FFBCL-, ,i'r For il~sc.~ncr lIy a ~ r and rnl1xt.d \I it11 eleinellcc nf n t l ~ c r d fizln~r. Even (hit k-~hcI1c.d ,111j1n.1litycrc ~ r i ~ n , ~preqeiit 111 .I hchvccn the p,Iir nf khclls tlf .I bivalve ur br,~tlilnll\ pod. tliir qpnrr may citlicr hc lcfi rrr~pr).or Lwcomc f>und, t1lc.y rnay not br p r c ~ r ~ \ ' r 111 C . I I I L ~ ~ ~ 6 1 d: ~ I nle11t- I T ] \ V ~ I I L -t~l r ~ - ~ ~ - c ~ i lv ~ i t c r ~ ~ . t i n ~ rtci~li~-, tjllcd w r h ~tclvnicnt.In thC 1.1twr C R C C . ~ ~ d i l ~ l c ~ i t re for core t f prcrmn,t.d. ~ ~ ~ l ict>rriccn ~ i illtdct W ~ I C tI1~1 ~ ich t I I l F t d t I C C ' . C " I ~ C . ~ ~ C O I511~ll~ 1 ~ 1 ~ F W I > \vltkl~ll.I fc\V IC 11 ~ ~ V ~ rock i c crnckcd ilpcil. Thiu hcnrh upon ~ t c l ~ r t ; ~,I:) t , c ~ I C J ~ S L l c f i ~ rt~ r \e~lliilri~t r . ( v > ~ p , \ ~ t ~ d rnrk. h tilrn intetnal mould of thc Lwil shcll. whctcas cxcerhinct. ihc. ica floor is !lot al\vny+a rcgroll r d ct-mt~r~~tous scdirnetlt dcpour~on. rllanv a p p a r t n t l ~ corznliu- nal moulds art- left i n thc caviry from \1,311cli lr slrcll, nr ut15 r e d i r n r r ~ t . ~ ~ \rqucucc< c o ~ ~ t a ~rluinrrmir came. In rx-c cirruitistnncr< the r-orc nr t l ~ c ti both. tn;rv he rcplaccd hv an cntirclv d~ffcrcntmln51n,tU-scalr Ihrsa ks ( d i a s t ~ m s ) rcprcrcn ting yerlodq r lc of V J I I I I I O \ \ I ~ ~ ~i i ~ cro\loli. Any ~ 1 1 ~ 1011 ~ t h r SC.I rrnl, JF h;~ppeiiq i l f o ~ ~ i prc\r-rvrd In iroii<tnnc\. I f L d 1
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Occurrence of inyedebrote !ossils in Phanerozoic rocks

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CUP F I possible rocerses OF foss~lrzat~on a bivalve shell ( a ) original shell, buried In mud [leh) or carbonate (right);(bJthe of hl *or m t ~ ta gun& in o m b a n a s d i m n ? m d r o s preserved ~ntoct h man o o small cmtollired rtch; (c) shdl *rig~, na r ~ ~ ~ ~ ~ o g o nrerytnILzd to m i c i t e rhlrh dertmyr the fine ~twcture, original ~ I C I C &dl retainJsurroundangc dim now ~ t e , (dl i - P C core 0 SIIICU, 1 (el u s~lrco growing o n the ovtslde 01 the shell, (f) tectonrc d~stortion o shell preserved in mudstone; (g) rim of i e I ~ r ~ s e r wIn lmud w ~ t h r orrginol shell moterral leached away, lew~n an external ond on ~nternol mould, svrround~ng rnwdsl~ne a ore I o coicoreous concret~on ; ! growrng round the shell and Inside I~f tie orlglnol cavity was empty), wlh p k h e s of pyrite rn ploces; I fonttane re,>locernentof core ond port of shell

q?.icr~ tlw ~ h c I l i~nprryintcdtv~i-I1 111 ,lrcm I ~ I I ~ c ~ I1i-I ~t~, 5.11d to hc perminetalized, I* IIIIL tilt. ITOM t h of ~ C Cuid.tl7' ~ ~ i i n ~ r ;;I[i l <t l l ~ I -\!l~ijrcoc rht +hell 1s replacement. Corcs ninv ~ . ~ t ~ : l r i i t - \ of p ~ r i t c .~;r.ll-tollt~\ IN.1rc 0ft~11prcrrr:!l Ilkc rhrr, . ~ n ~ r o r h ~ c 1 1 . 1 ~ 1 1 1 rcleawd rIcr.~\ ~ i\~lrl~:.:ri \nIpl~~de. n.llt. 11 r~-.~ctccl witll frrrous I I I I I L I i i t c i t I t i ~ I I t I i o I f 1 I \ r t l . l r ~ ,111 i ~ n ~ l t ~ r alc~tc i = d clicll T h r ~ linppencd 11nc
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rllc ulica 1ncre;lqer ~rlarkctil!.. nnd thc s~hca G O s c l c , ~ c ~ d r r ~ v e throng11 thc rock nild precipltatc ill l L\~IICTCVC~ r l l t pI I i q lowcr. The zii5idc of 3 qt.,~ urchrn cieu,ryirig undcr dl6icrcnt c a n d ~ t ~ o rwould trap juqt i\ cuch an intern21 ~mcrocnvironrnc~it. witl?in u ~ h t c h thr- 51llcn could prccipitntc nr a gel. Tuch siliccou< corvq rctnin cvccllcrit tc..~tur~*q, prc\crv1ng the ~ntcrnnl ~ ~ l n r p h n l o gof t11c sIlull. i3n chc ntllcr I l a ~ ~ r~licn y d, 111.1~ r p l a c ~ r C,IICIIC very t & ~ n ;IF rhcll ovcr tllc w r hcc of d fosil ns a rvsult of wrnc crrr~lplcus~irKitc
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1~~1.1iiJ 1 i \ ~ c 1 ~ 1 r r 1 1 ~ ~ d ~ i i o f cnlcalc- tIc~.~rlcrl TIICY 111 o ' it~~it cvprefsiot~c)f the ~ u r - ~ o fctllr: few[ and, u ~rlud,tlong \v~th 111.111v y v n g c c , w h l c l ~ c c t c t e d m c c . t h t y ciin hc freed frorn the sock by d t ~ \ o l v ~ t ~ g ~ I!IL~IIL*\ r l f htoqet11~FIIIZJ :I ~l\t'lc'toi~. alk.~liiic tlic I~tr~cqtor~ct h hydrochloric ;rcld, 111d1vidu;ll In w
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Principles of palaeontology
tr~lolxrt.~ bral.hiopcd< ~ r c LIIO\I Froill ~nntc.ri,rl rciilaln\ u i r n m c part, 1i~unlly ~ h c l of r k c l c t n ~ 0 1, .j .lnd . rl n l ~ slit-I1 a\ rill?, ~liicc-l~ving urganlsm: (2) trace fossils, whir-h ,lrr A rclar~vclv ~ I T I L - O ~ I ~ I I but I IC ~ T L ~ I I I F I T C1 n o ~ 1 ~ ~ track<. tr,~ils.llurrcrws or uthvr cvldci~c-c i t h c .irr1\.I~) * I E n p r c ~ ~ ' r ~ ~ ~i~ t 'phosphatizatio~l. C(rin~~tirnc+~ c it? ot' n r i a n i r u d (71' FOTIIICI. tliilet - C ~ T T I C ~ I I ~t~hLc-rF; ~ t oi1 tl cxtcrrl.11 <kc-lctc~ii. :F~C'CI.II~V thin r>r~,:,lr~t~--d~clltdthc oiilv q ~ i d c thc fi>riilcr prr.;cilcc nf'.iofinf arc. to nnilnnE~,Inn) I3c r ~ p l a ~ c d ovcrgrolvn bv .i thin bvdicd anirrlals 111 .I pnrt1rul.1~ i ~ r ~ ~ r n n ~ n(31 ~ t : or c ct s h c c ~ p1lo~pli.t ~ LOT thc- I,> trcr t n ~ vrciii tbrcc :it> chemical fossils, r c l l c ~nf b~clgcil~c 47f t , nrganit coinor1~111.111~ o \ p l ~ ~ ~\licll. 111 ~ l i cf n r ~ qtttidmt~n poi~ndswhich IIIJV b r drrccrcd gcnr.f~ciiiir.~llyi ) pli tit~ ~ ~ r t thc csrcmnl iO17ii r r f t h C I,~11jy1, ~ ~ C C I S C I Y r c p l ~ c ~ ~ t c d . r(7vks. tllc Altcriintivcly J F E I I > T ~ ~ . I ~ I L hllit~yof tlic i i i ~ ~ r i ot' r , At t l ~ r l~ Ileart OF in\-crtcbntc pnl:~cniitc>Io~~ ct.1nil4 thr \lir.ll r t ~ ; ~ form .I r-orr, yrc-king o u t r~ltcrn,~ltaxonomy: thc clarrlfic~tionof f I > > ~ i 1 v .111rI rnodcrn stnicturr5 1t1 rcmnrknldr* ~ % ~ t . SiL.I LprCsrw.lti~i1F \ A ~ I I ~ I ~~~l i tCvorrlcrrd and 11.1tuml p(111pi11p. T ~ I C W i l '~~ I proL.ahl\. .~rsoc~.rrcd wrth Il,ir.tcn.~l .~~.rivlty dirrctlv gn)~ip111gs, u ~ o j ~ ~ taxa, i i ~ ~ r bc ilnrncd .~nc l a% i \t nftcr t h c dcnrli nt'rlic- ,1111111.11. M . I I ~ c ~ l ~ Canil-ri,aii cirraugc-din n 11wr~rcll:alcyqtcm 111 xvhirh t h r ~ rr.l.~\ ~ l l r fos~115 lin\,c brrn yrt-\c,n~cd b!. p \ ~ c ~ \ p h : r t i ? ; ~ t ito ~ ~ l ~ l i ~ p s rn:~declcnr. a n d ;IS t:lr ,I< pclrsihl~,murt io drr (Cl~aptcrs ~ n t2). hut much largrr fu<\~lq 3 d illay Ilc hc cccn I l r evnln~ttonnry pcrspcctn0c prt.wr\*cd nlco, 67s r.r,~ll~plc c-ru<tJccans with a Evolution theory ir rnrnpout~dcrlo f vanolis dlsHr~nr~p~r~tr 111fil1111g i i d ~ v t h. ~ l t t k t C 1 ~ C ~ C ' ~ I C . I ~ Ccipliiit.< - I-TU~C' blolov, curnpnrnrivc ,Itlatoirlv. ~ ' appczlda~e~ 1r)t:lct cmbwchogy, gi-11ct1t-s, ~ n dt ~ n p ~ i l . ~lt~ n ~ o g- INI: i ol~ * Forr11.i .lrc o i t r i ~f i ~ . ~ i iId~ concretiuns: c ~ l c , ~ r ~ - i h C ) I I ~ Vtlic p a l n c o i ~ t n l o ~ c L~rpccth ~ nlln\iru thr I lt .~l t t ous o r silirCouc i i ~ n ~ c c -hrn~c.rl .~rtruildtlic f o ~ r ~ lprrdictione of evolutionary ccirticc to hc t.c\tc.d r .shorrlv .iftcr ~ r sd r it11 ,~ni-l1711n.1lI'ri~i~.rctlot~~ fLs111 ,bgniiict the b a c k ~ r o u i ~of gcoln~ic.iltirnc. pcrinlr7 d u ~ l d c r C ' C ' T ~ ; ~ I I FC I I I I ~ I ~ I C I o111\, \vE~vrc .r d r l ~ r , l t c tlir tract'tlg of t.~.nl\,rnq ~ i i c , ~nnd < II~ I ~ c 11111~t~;lt~'i~ C FOII cllcnllc,il h,il,lncccclit< \lcn\.r~cii \ V A ~ C T~ t l r d i the. l of the patrcrnc of cvolurt'oi~ rhnt .irtunllv 11:lt.c mcnt, br. prncr.c<c*.H vcr ~ ~ ih lI I t! ~jrrd~,rst~rcsd. urcumed. T h c ratcr at which at'li~11;lls 11.1vc. r.volvcil I L I Y ~ varicd through t ~ ~ r ~ c , mnrt .rnt'm,il t\'pcbut {species) 11,1\'r 113A a geolcrq,lr,~llit% of ordv .i 1111111oiivearq. Sornc nt^tliece rvnlvcrE rapirll~', t ~:I\c l ~rlqvrp 1 . 2 ~ 111 \.cry r:ltc clrcilni*;t,inccc\oft-brsciicd ursanlsrlls thc ,~nullo~iites. othcrs v c y slowly (CEI,I~~L-T A 2). rnli bL- ppl-cwn-cd Ir f i ) ~ < i I h ,2nd t11c.c~provitic clthrl-roclr F L Z C S ~ C C ~ O of liiJrlrlr ~edii11~11t~ 11p OVCT II lluilt \ V I ~ P~ ~ i ~ n h t . l ~ i C.V Ih IC I ~ I C t h d t ' v ~ r ~ lof i~lt'tal ~ ~ r ot ~ ty ".". rrlany n ~ i l l m unf- ycan mny rht.rcf;,rc li,lvc \cvcral ~ zontls 11\+111g , ~ pafiiculnr rc.nnd\: this tc di\cusccd 111 t klT' t -I ~CISFII P C C ~ C ' Snccurrii~g a partic~il~lr S in ccc111~>11cc.r h cn Chnptrr 1'. LC.rt~"ll I qpccicq corltillcd to iliir parr of thc ~ucccc5ionnnlr and rcprcscrltirlg tlir timc ~ v l i c ~ l spccicq \V,IF th,>t Irvir~g.!-l-lurcm lirr the oldcct 2nd 111mi gcncsal .1ppI1carioil of n~vcrtrl-rr.~te pnlacon mlrrh? : biostratigraphy. Uslrly the qcqucllrc o f fr~\nl K ~ l i i l . t ~tllc . geolo9cal cnhuil~llh a i been c I I ~ ~ F C J C ~ .I ht'nrq lip 1i7tn I~~vcn-tchr~~tc. ~ c n r ~ tI \r ~ l r ~ ~ ~ 5 t i l ~ ~ JS d of m.lior gcc3ulapcal t i ~ n eurlttr {prnndc). chit-li ot p~l non~i.i!/v~ l~ subdivi~inno f qr'olop, . I < i t iq t ~ i r l ~ n Enrtl~\cit.ncc \vhicl, i~ furthcr d~vidcdnltc3 ,I Iiirr,ll-thv nf ~ n ~ n l l t h ~ it< w n t c c t , ~ ~ ~ l i ~ . ~ lti i .i > t I. s .~lct) rci.11 ~c unlts. Tlrr \\ 11rrlc b ~ < fur tl115 li~rtoncnl t r It l 111 t)ca cSlroiinlop .I bt'ologt'c~l u h l c r t I x ~ tlilc rh,lt h,i\ thc uiirqur pcr~ , t i~ dnvumentcd qcqucncc r ~h5cll.; 111 rhr rtjck~. tflc f G p c C t i v ~o f p . v j l i y i ~ . ~ l t111l~. W1tl1111 tllr ~ l t ~ r l ~ ; lof Uuc difTcrriit h n d s of (05~11q 11.ivc diftcrrnt <tr.lnin it'lvurtcl3r.1 p.~l,~crtohrgv t hr-r~. art. .I r ~ ~ ~ r n loe r p a ~ ~ l u c n l tt. ln il v;~liter.. ~ r ~ d CCrtJlIl p,lrtx nf thr $coEogir.~l ititcrrc1,itcti topic^ (Fig. 1 .1), ot' \ v l i i ~ ! i IlavC .I rccvrd are nlorc. closclv wl~divlded tf1.1n n ~ l ~ c r . 211 hcnring 0 1 1 the othcrc .md ~ r h ~ c ~ l c u l link L I P w ~ t l l Suiiic 'ah\olutr' agcf b a ~ c dnil r-ndiotnctnc dating cjtl~er qcicncr~. Ilnvc b c c t ~FLYL-J p.~rt~rular ,rt pclints to this rcl;lttrtc Tl~rrctnnin sLitc.<unrr of i i . ; \ ~ l + t l 1 . 1 ~hc JIFTIIEsc,,llc. ~ n d rl~csc prrwidr J fiatricwurk t;>r ~ i r ~ d c r p ~ i h r r l (:1 ) body fossils. ir, ntlwr ~vordkthc actual s w i i ~ l i n ~ g c o l n g ~ record in tcr1in of rr.11 lriirlr tlic ~l

iI~urn

I ? The wrlovs sukdiriplines of palaeontology

l Y 5 h . is concerned 1 ~ 1 t h rclatioi~ship~ fosd thr of aniln~lsto therr r~ivironmt.~-!t.both as ~ ~ i d ~ v i d n a l s {autccology) nrld i11 the faunal co~nmuuitics In r,~.:, tl~: c~~:c.nlayc,~l timcrcnlr hn~cd l>iostmtipa- ivhicll thcy occur ~ i a t l ~ r d l y : lnttcr i c ~oinctirilcc on tllc 1"" I \ ~ ~ ~ l ~ ~ t qhl . ~ 'j ~+Rc~ d ~ C > Z C ~C ~ J I I ~ I C I Iknon.r~a5 synecology. rI C IC ~~. ;ilrl~ou:l+<trat~qnphl- the basrc nE tlic pnnlnr). 1 s Sincu thc <oft pans of furfil anininlr are not nnrE;~i\l>l[llr rlt' peochronolwp;y. J s r ~ ~ a lhtwt- n f l ~ ~ i a lprc%crvc.d, lv but only thelr hard FIICUS, therc arc 7111~ ~-TI;~IT-~LIcJI ~ t ~ l h i 3 hmnng ou wlint liiay he ~~1.1tivcIyC Wm-~ys n wllich thcir biology and Ilft d3 ~ ~ i ~rrr.~ol gcochronotnetry'. L1v counting daily Railits car] br ~ i ~ ~ d e r s t o o d . Studicc ill functional :tslirfE~ nrlp 111 cxtil-rct corals and hivalvcr. infornumorphology, howcvcr, which dcal with the Inter: r ~ ! n l1,1\ tlct.:~ n l v ~ i ~ ~lr,c l : ~ r ~ i ~ g t l l ~ ~ ~;ipon nurnher of pretation of the b ~ o l o wof fo<~ilizcd skclctons or i n > 111 III~ IITII~IT 111n11th yc:~r111 : ~ n i - i r ~ ~ti l ~ c s .h m l c ti1rc.s in tenns of thcir original f i ~ ~ l c t i o n , ,111d ti I~avc 1111~ 1h.1~ Il~.lpcd (*oltfin~i to ~topliyritalcstrnlntcs rill hccn ruccc<rtirllv~ t t c m p t r d with marly kliidq of fcrq' I II ~ I O I ~ ~ I ~01'tllc E,~rtli'~ L: rot'it~oil(Chapter 5 ) . sils, rcqtnutcd in scope though thcrc ci-dcavours may ~ I I I ,~rr.~riyr,~pltic,lI ~ nppl~catinrl< p.~locontnlng nccrsqarily be. Ichnology i? thc sti~dy tract- COGaf of 11 lr, , ~ l w . r I r r ~ impnrt,~~lt. iiiorc h i n l o d ~ ~ ldh: thc t s n c k ~ , surface trails, burrai~brand b o r i n p ~ w <el rhr 1 3 ~ ~ jc l01 p:tlat*nntolo~~ t rvcrc rclativcly ncgltctcd nladc by once-living anirnals and prcscrved i n ~ c d l I I 1 i 1 r~'ce1111v. Palaeoccologv. lncilts. Thic topic ha5 proved valuable botli in ~ -111 ~ l rlln+ tEcvrlapcd pnrtict~I,~rl\. sfracc the c ~ r l v i ~ r ~ d c r ~ t n n d ithe bcIrai4c)ur of t l ~ canilnals th,lt iig
I
I ,

I,tron r i pcnnds o f ~ n i l l i o l ~ ~ c m ) r;~thcr oty th;ln I pur.-I\frr.l,~tn.c binlc. 1his Ir o i ~ l y pnnihlc a t ,;*FIIII ilor~zoi~~. \ ~ ~ v rand for p r ~ c t ~ c a l Iio r, pur15

livcrl whcn the. sr-rl~mr*nr hr-lny rlr.pnr1rc.d and 111 \\,a\ Tlicrc word? cliould ~n.ikcunnrcly clc.lr tlic ti111Inteq~rctlrly thr. c-ontr.lilpor 11lc*o11c cn\ironIncnt. darncnt.11 trltllrc o f mstlnomv. For, n k ha7 rrt-rcn hcct Finnlly. it I \ only h t t l ~ r~nrcp-nt~on tflsoiinmlc wid, to idei~tifv incs~lc o ~ c i t ir ~ t i n t rtcp. ,ilv ni a l thc' Ci~t.1rln loi.~l Cllinn~thnr rlir ~ l t > h distnl>~~rinn indccd thc kcv. to tilidin? out itirtlicr ir~fonii.ittr~t: ~l nf mnri 12c 11l~~'m'b~.: 111 r ~ l l ~ y fKlI I I + C cnii hc cluct- :ahnut i t . Stwi~clcI,~s~ltic,i ,tnd ~ I D M C ~ ~ C ~ . I C II iIr T C trs ti011 dntcrl CIICII c r u r i ~ c q t > K palaeobiogeography (nr nt r31c rnot nf ,111 b ~ n l o p c n l.it~d p:~l:~co~itnlnpii palaeozoogleograpby in thc c.1~~- nnirnnlr) rnrl li~nrk; w ~ r h n u t rhcm iin cnhcrcl~t nnd orrl~.rc~ of Zit* r ~ r c t l 111 r.nnlilnrt1on nrrth ~ c + c ~ y l ~ \ - ~ i c a lIn rint.1 y t c r n c)< dot.1 ctnrJgc and rctr~cv~il pn~.;ihl, 1.c ~rnrIcnt,~n(Iiny tll~. ti>rni~-r . ~ l a t ~ FChlRUIIh n t d T.~x.ionnmy, or systrmatics .I.; ~t ~ \rc winctltlr~, iiiovciiiciitc c~~rnntiiicnt.il m.~\\c\. k ~ ~ o w ii s ,tlic \rrc.ncc o f ~ l , l r q i h ~ . ~ r ~ onrgnlllrim i or i i All nf rhtrr aqiccr$ ~ r t ' p : ~ l : i c o ~ ~ t carc nii~rclrc- ' It n the oldesr of dl1 thc h o l o q c a l diwipliiicr. >l ~ 1.1rcd. rid :III ,~rlv.uiccit1 nnc iii.1~h : l \ ~.I hc.>nng thc pnlKl~!t'> i~titlinrdLlv Cni-1 t;ti\tn\' I ~ ~ ? n , i r r ~ . iipnn nnv ~ > t I l ~ T r . ~ w.I p : ~ r t ~ r u lctutly i r ~fiii~c- ( 1 7 17-1 77s)in htq pioncrnny S)~,ft'~ri~t =h .~r 1 rrrrrrr?.rtc, ~ r . tlnt~:~I n ~ c > q ? h n l n ~ t11.1y ytvr irit;,im.itinr nil y chll in uqc tr,dny. though ~ ~ ? ; i t lilr.r~diticrl In8' v p~lacc>~=cologv ~ ( I F E I ~ IYC~ITIC* ~ C ' C C I F M C ~ to tnw311- curcndcrl. ,1111Y o n ~ p \~1.11.L i k ~ :a\ ivi\c. rccchllr r t - t i n r t ~ l c r i~lt ~ l t.iuonotnlc pr.1~-clcc li,ivc cr1.1blcd ~ h c dcvr.lnprncnr nf .i . The species concept rrluc: 11 nirlrc, prrbils<. ctmttg;~ph\. Thc fi~nd~~rncnr,ll n f t.lxolrtln1v r\ t11c qpcclr# unit Chcr~iic,ll r.r>n~piv~~itlc IvI~lnq,?c,~l rlf ongill can Anim.11 ~ p c c ~ c(s g. Sy\vcctcr-Uradlry, I'l.5f1) Jrl c 1101~ hrb rccovcrrd frclm ,rnc~c-n lrkr ,lnli f ~ m n t rl~c q r o u p a f ~r~dividualr rhar gcncr~llylook ltkc cat1 h,lrir of biornolecular palaeontology. Such fnisil ntlicr .lnd can intcrbrccd to product ~tT<lfrl-'r~~~g ofrl~r rnolc.iulr.r rnn) Ii~.lp t o r l i , ~ g n o c\VIIICII rlrgillsrns FJlllC kind. Thcv cannot 11itc.rbrcrd with nrhr ~ thcv CoriiC frcm~,ilici thcir hrc.il;rl~n~i~ p,r;rllt\~,ry~ mav qp:pccnc\. Sii~cc i q rcpl-nd~1ct1r.crrol:~tion nlnnc rl~,, ir $;I\. soincthiny c : ~ h o i ~ t cnv~n,nincnr. Mvlolrcular thc. dcfirlcq rpccicr ~t ir oil111 rrally po\~lI-rlc dirtunyu~rl, to pl~vl(,goncti~ s I>,~\s:d ripmi pra %tchvn ~ - q l r ~ n r i I1-lln\. s lg clnrrlv rel.lrcd <pc.c.rcr ~f thrir brccdiug h,\i71r\ Jr \hi>w hov~K I ~t \ v ~ ) r r~101-r ~ ~ 1 , 1 t c~~ l I I I < I I 1$1 ~ v r ktiowii. I3t' ,111 thc ricscnhcd 'cpcclcs' ot' liv~iig o O ~J I ,JW rlivcrqrd horn ,L ~-l-rmrni~~i ailcr-cttlr, nud to ~ornc ~nnl<, ~ n ~ v ~ vonly ~ 1 7 i l l 1 r n rivtli .Ire 'good', I. l cr, ; cstcnr thc .~v.~~l.~l\ltrcChni~luc\111 i -. ~ p p l ~ c d rhc propcrly drfiiicd, c y c c l c ~ l ~ i f i ~ r ~ i i . ~ilpnu~ tlit c. x to tioi r r ~ t1 r I T ~ I I ~ ~ L T I ~ C ) I ~ ~ _ ~ ~ C - ~ I - ~ rcpmducnv~.pl-cfrrci~cc< f t l ~ cotlicr tivc-q~url~< ~CIC~T~~~~~;II~~ n I . trclln?clur\ can hc ~iw.tlrt) IIV~CL t prot~.il>c n polv- .11I n.ltlii-.~llyocrurnng . nirnal pop~tlanniiric p r r n r ~ d I ~ n ~ ~ * h , i 111 d ~~r i ~hcllc ri h l htrr. thl- rhc tnntncnt. c.t,ly docui~lcntcd. ~ I i ~ l l younycl- t l ~ , 2 Ma laavc prnvc.rl :um.cnal.Ic t43 lr ~r~ Tlic rlifl;.rci~r~ntionnf i n o q limn2 3nc-I .rll fn\\l a n , ~ l y \ iTli~.rc.lrc .11~o ~. pm~tll\tny E C V C I C I ~ ~ ~ C I I,115~) ywcicq thcrcforc Ilns to hc based lipr>ll ot11r.r- .In, C ~S 111 palact71-~ocIicrn~\tn-~ r l r f ory,11111. ~ ~ ~ ) i - t ~ ~ tcchnirally Icw v,~lid . ~ i i i ~ t y rntcria. which arc .~pplat-.~hlr> thr. f i ~ i 1 to rr.r.~td. t h a t l q h COf tlle5c hy fir tth- inmt ~mpnrtnnr.r.spccinll\ 11 t11l.s~ bcvc~ntlrh~, .>rc ut-npr nt-tl~rr l,nok. p a l , ~ c o n t n l o p . I < morphology. rhc kncncc r i h n , slnc-c Innst nnrur.11 spccir-F tcnd to be cntr porr-d r.rf ~ n d i \ n d ~ ~ of l c , ~ sitn~lsr cnouvh rstrni Taxonomy ;zypc;lmncc to bt- ~ d r n r i t l ~ b l c o!"rhc vnmc klnr nr K h + h n p l ~ r l i ~S Pg C ~ C S of l i w i i ~ ~ nC niiimnls by moq-llo lnpcnl c r i t c r i ~nlnile I< ilot \vlthottt h.17nrdr. c q i r c~alEy\vl~crct11c ~peclcsin qucalon .Ire ri~ni1.1~ all1 cl{xt.ly I-el,i ted. S~~pplclneil ' ininnnn tjcln. a 11 I tan 1:1 rhc ni1.1lyur of spcc1c.;-~pccific protcitlr, may he hclp 111 some carer whcse thctc 15 good rc,lqnn h t t s he <ought (e.g. for dlrt.;ice-c;tq+i~~g o rilr~ct';). Fr thc rcsr qniiic dcp-ec of whlectiuitv t r ~ tnsnnoiilv II I to bc arccptcd, tl1011~1i tI11s cnn bc n ~ i n ~ r n ~ z c d cnourh ~norphcllcrSicalcntcna arc u ~ c d .
1-0
L m

Divisions of invertebrate poloeonto!ogy

4omenclature and identification of fossil species conlparing the cpccicc point by point. S n t r ~ c i thc n lit. fnrn~al ~ o r n e i i ~ I . ~ of any ~pccic<,i n n g o r sprrlrs m3y prnvt to bc ~dcntical u~it11 nlrc.tdv i ~urc l 11, taxonomst< Knllu\v the biolcrpc-nl fi'ftrrli of dcscri\>cd spccics, o r rhow only rnlnnr \.ari.~tinn<>Fa n.lcu5, \\,l~crcbvt-nc11 cpcclrs is drfined hy n ~ ~ klntl that would hc c ~ p c c t c di r l a lclcnl v,~ri,~nr n w t h i i l r h fame ~ ~ C C O tC c r . ~ ~ h C \pccic.\ i r ~th? CIUIIJ ncc: tllc generic nnA specific (or trivial) ndint's. rt.l.~tcd. m.iy hc ~ C W snd if rcl a fill1 rcchnical drqunption . csnnlplc, all cars. 1.11-scand < i i i a l l . illu5trationc rnlict hc. prcp;lrcd for cnch ncni I onc pnrnci11,ir ~ ~ t w i p heen ~ I . I C C L ~ ln t1w ~ v ~ r h 11.1~ should frc pzihli\l~cdin a pdncontoaoa pt.!i.r. C)t the. IfLlnnuc .;pcClc.< 14t'/;i thc spc- ~pccicc,~rliirl~ of Iogic~Ejournal rlr ~ ~ i o r ~ o y s n p l i . dcrcnptintl 15 This i nniiicr I: rrltri.i, I-. !r.o nntF I . . pnrdl~r tbrni.illy . to tlic Jumertrt- r.lt. lit>il .4iid leopard, rcq-cc- h.ncd upnn type specimens, which arc n1wav.i *. 111 fulF t;lronotnic t ~ o m c ~ ~ c l a ~ i i rn~lthnr'l, rherc,~kcrkept 111 n InuGcmm o r rc~catch~nctitutc.. the c ~ ' .ltld the dLltco f pllL7l1~,ltl0tl 31.C glVrl1 R ~ T t C l ~ U ~ u a l l yniic of thew. tl1c holotype, is. ~ c l c r t c dnr l ~ <, e,p. I:t,lrc irlrtrx ( L i t ~ n a c u177s) (rcc hclow thc refereilcc spccimcn and fully 11luc;tratcd;coin: ~ p,lmn\lc dctnil 111.1y bc ,~ddcdfrmn o t l ~ c - r p c c i ~ ~ ~ c n r s ~rthcl- i c c u ~ ~ i n n ) . d tElc pnlncnt~tolo? I[ c n n nc-vcr hc kiia\;ll for c,~llcd paratypes. Thew arc v a n o w othcr kind\ c, L L I I J ~ \vIlcthcr n poplil.lt~orl ~ \r.trh n pnrhriilar mor- o f n p c spccimt-ns; fix c x . ~ ~ n p l,Eneotype mnt. bc hccn Inst.. o r ivhci1 ;I ~ h n t wnq rcprndtictivr-Iy tsolntrd o r not. Hr.:~cc r,rcctcd ~vhcrrJ 11nl01ypc h n ~ n ~ 'hr ~ C ~ I I I ~ I C > ~ I c>i<prcirc 111 pnl.lentrmln=, a< In rnosr s17c~1rs h n j : rcdt-ccnhed In fullcr and 1ttorc 11pIlnnr: spec1ilreil<. inuqt b-. b,~cednIml-r<rciitircly on ro-dntc tcmis whcn 110 type spccirncn ha.; pzcviouqlv ,~~~i~-phologc,il cntei-I,I %4nrrt>vrr.nnly rlie hard t-ccrl dcngnntcd. A i~tw gtnur wtll ht' d c f ~ p ~ t as d c gcn, new. 17): it\ ~pinrof' t11c t'mcll ,1i)tn13l Art= p r e w ~ c d . 3114 much 11rr.tl1l cl:~tn1 ~q ~;1111511t(1 1 A c.~rt'fi\I~ Y J ~ T I I I I . ~ ~ I C I T I nritlior. t l i ~ q follnwing thc gcncnu rlntnc, a nvw and I!lx~rment,~tic~n.]I1 tllc an.~trrniiral~ C A ~ L I ofC tlit' sptcic\ ns sp. nov, niid n ncw suhgccics .I< FCP. 1101.. nf ~ C Thr ncur qpccies must hc narncd and allocatccl t o 1 ~ 1has to be the maill giidc 111 c.;tnhlichir~gthnt 1 I~IILI P C C I C FI \ rlifl>rtwt irnrn ,I rclnted qpccic~. ~ In rnrc nn cs~'iting genuc, or if rhcrc I$ 1 1 0 dcscsil~cdKcnus to \vhtch i t pcrtnin~ thcn a ncrv gcnuq muqt ~ 1 s hc t~ ---: ran bu ~lipplci~~r.~ztcrl cornp,rwon nf thiq b\- .b l:herr~i~tn. thc \hell. .I< Ilac provcd e<pec~.~llyerected. 'To apprcciatc the mctliod Ict ur con~idcr of the follom-ingh~storicaltalc. In the early ninceccnth LII In rhv crcctmn nf h y h c r t , ~ r i l n n r ? ~ ~ c rntecr With~n anv ~ntrrbrecdint: pnp~il.~nnll tlicrc i q ccntliry hr~chiopods\vcrc poorlv kiinnn .111d fc\v ~ilrqultr. a s~.7wd05 r~~cwpllolugi-~~l v,ln:+tlon. dirtinct gcncra had hccn crcctcd. Oiie of thcsc w.lr ; I~ro;lElt-r r d l r t h r r r iiiav 11t. hnt11 g r o g ~ h i r : ~ ltlic living Trrrhmrrrln. nnmcd by 0 . F . Miillcr in 1 c ~rratlgaphtcvan.jnoiic. . I I I ~111 thecc tnllrt bc 177h. W h c n E.F. von Srl~lothcinl fint srudicrl , fnmlr From North Gcmiany in 1870 11c it~llvdncutncntccl I < thc rpcrlrs. 1; rc? h r 1dr.11ly D e i ~ o n i m ' I-rll~hed. u c h srud~cqnLn7 be vcrv vynificant. In r r c ~ ~ p i ~that come of thr shclfr wcrr b r ~ F ed chlopod<, , ~ n d de~crihcdonc o f ttlc mnvt abunhe Lutinii~~ ~ln~iri~ pL tihop. rlnr~t &mi\ a$ thc nclv qpccic.5 T e ~ ~ ~ b r .itlti-it~rrlt~rr:.r. nhr~~~ A ' l l ~ 1 1d ~ ~ . ~ C O I I ~ O ) O C Jtll'l7l)~tlll~ dl5tll1If ~ F I t0 Hv EX.70, however, r n ~ ~ c h nlorc waq knolvn dbout , : ~ I I F ~ rhe qpc.clrf 111.I 11cwYv iliccoucrcd Ounn. s;ly of trmchinpods, a n d G. Fiqchcr dc W d l d l ~ ~P i~~O~~l C I S C ~ t FEII hrai111~1~0ds. iv IIC IIJ\ ti1 iep;lrAtc. rlir tndi<)IT n so t r:du,~l h \ 1 1 ~out l r l t i l gmtipq of n~clrphnlop~c~ljY ntw penus tior thiq qpccic~. t h ~ it bccdinc cor111ri i ~ E ) ~ ~ . i d' ~i Ei I~ Vl ~' . 1 1 1 ~ to, t . ~ k 1 1 C ~ ~ ~ ~ cxn~nple, rcctly desi~ma tcd Chonf8re.r j l l l r i ! l t i l n t i t r (Schlothcin~j. I V h a p hc clqht such potjps, r..~cl~t s t ~ t ~ p i s h e c l cl hv Thiq ir thc 'tvpc rpec~er'of C I ~ i u ~ t ~,1t e ~ . C ~ ~ - ~ ~ I ~ L V nrrlculnr cct o f C ~ ~ I S , I C Solllr of tllt'w ~ C J I I hiluro-13cvom3n genu< of thc Cl;tn Stmphomcnat.~. I ~ ~ . ~ ~ Nore thnr whcrc a FPCCIPF W A Sunpnally drscril7t.d r 1 clcarly rll<tlnct h 1 1 o m nnothrr; 111 d i e m c . 01 l lll\t~ncr~c>ir 1.r l onsidrr.ihlv lew, incrcn5ilig urldcr a dlffcrcnt gcnrric tlnmc, thc o n p i ~ n nutl~or's in,ih nrk tit gt,~ E I ~ ~ > J ? c ' ~ ~TIlT \ ~ '. fp-nups nrc naiiic i q quoted In p.~rc*nthcwr. In 1917 F.R. trr v ~t ~ I V I ~ I L ~ I ~ J I~ ~~ r i c l c l r r c ~ ~ C C I P S .\ \ ' l l i < I ~ 1111lqt th~11 Cowpcr Rccd, t k c i ~w o r k i ~ l gnn Ordn\+Fc~an I S~ ! :~nd of ~dclmtirrl.This I F doll r bv r-oiw~ilr~ p.rl~ennto- Silurian hrach~apodr thc Girvan clrcrricr. Srotl:find. IIC L ' : ~ i r il o r i o g r o r~ ~~ ~ ~ cC O I ~ ~ ~ I tkrnilrd~ recogived r n m y ncw spccics. C)nc of ~ ~ C F had ~ ~ ~ . ;I ~ P; T U I ~ lu~r(.~l ~ c s ~ p t ~ ~ i~~ lc u < t r ~ t ~ ofnprcv~ously simrlantie~in r ~ ~ o r p h a f o to Cllorrr~r~~. it \ I ~ J ~ dc a o d~ l rr s w hut :trlhc~l I~r.~clunpctd a ~ a n a ~ slrlulnr ayc, nnrl s~itllcienrlyrlifl2refit to 7nc rcgardcd a 6 A <pccir.c uf n I of

~ I

ncnr ~l~llgcnnis;l i ~ \1s nrnrtr-n U/rt~~icrr~ r (Eorl1trtrrrr.r) Thcrc nlay hc vnriuuc ~ t ~ b d ~ ~ rof ftil~ ~ ~ r i ln l t > ~ i r : t lf c IL)17. WIIC'II111 I q I X thc tasnnornic g ~ r n r s ~ s u p c d ~ t ~ ~ i lwth ~ r d c r ~ ctc.. ,mrl I c.g. i . o. . i\ Il1c.h tllc, p r n h l e ~ ~ ofs C/rtvt~,tt'.r ~ 211d un11l;lr f;\rrnh wrrr tnln grnnpc thcrc if pvun n cnrc for rrccrinc pl~ylnycn addressd b\ tJ.T. I n t l c ~ , the nrlv S~rpcrtirnily 'rupcrphyl~', Tlicrc drr only . ~ b n l .3n phyla 111 tlic ~t 114.I1~ 1 C'\Cl I'lt.c.t,~~~~bonrt~~ct-~ \!.ns crcctcti to . ~ ~ r o l n m t ? . d a t arlitn~lkingdom. and nnlv ahont n do7cn of tlirrc. ~. Illc*,lnt rllc r d r ~ ~ i !,iiic( Iilnny nthcr rc1:lrcd hmcliiopodr. At a ,~ c.9. Mollurca and Rmrliinpod,~.Icivr- nny fn.~~11 Ill( I1 . I l l i I Inter st.~qeI;tlOr~~rt,!vi \vns c l c ~ . ~ t c d t l ~ c to mnk ut' a runlains. E !I'TITI\ I n Full gcrius. I n tllc moxt rtcrrrt trc.itlncnt, l>.A,T. E tnxoriunlv h i ~ h c LIXJ Arc t i r ~ ~ , ~ lil~ h n p ~ i ~ h r r rt!llr\C* LTc n r d v l H a r p e ~dcwnbcd n Earsc inuna frmn thr Glrvan hy thcir suffix (i.c. -ca, -3. ~ t r . ) . ,171 c-xamplc, thc AF oi(lcr o t d~qtnc-tof Scnrl,mJ. 2nd ~ ~ r ~ t c h t~ hc recop-rlizcd folluwitlg docurncritq rlit= clns~~fic~tion tlic l i ~n of' ?I l ~ ~ l ~ c hvo w b < p c c ~ cof I:'. i l r l r ~ i ~ r r E r t ' t o n ~ c \ v l ~ ditfcrcnr r a. at CSrdnvics,~n hracl~iopnrl Fnriroli~ta ndru.lrlr pi?rf/lc , Lc !ding r lt n s r q nnd d ~ s t ~ n p l ~ r h chv rllliinr rlltFc;.rc~~cc+i cl In rcfcrrcd tn carlicr, accnrrling to 3 tnsnnnrnlc qchcn~r fi l \ ~ ~ t ~ ~ > n r o r p h n l o ~ Thccc. 111 I-larl?cr's ( 1 08") ~ ~ l l o ~ ~ u p a 111 hwhich thc aurhor of thc taxnn : ~ n drlic ycnr nt . p , 11~1,: lilt :I 11-1 ,ire \i7n trc11 ~ i o d ~ ~ ~ t~ it d r~c ~ r t t t ~~~ t t c di 1 17 (dcc- pnbl~cntiotiarc quc)ted. ~ - < ~ K ~ d l~~q~c: clar ig7ahnc R r e d ' ~o n p ~ l n lma teri,il). and Eoilra?~rtrs lv\[ri1ct11 r~r!~lrr~#i Rccd 1') 17 ynrr~lr~qsp I ~ C ~ V$libs~qucnt I'hylutn Br~cliic>poctl . I3urrlcril 1806 1'1 I L L - rrtucl ~11tllnn rT refrr to rlic I.~rter.111 full. as Et~clivx~r~tcsSubphylum Khvr~clioncllifonncaWilliam~ , I / . 1l 1 ct 1 !t-~:k c ~ l ~ f , ~ ~ l r / ~ * r ~ ~ l r r. Hatprr I ')ti". or 111 d l ~ h r ~ v i ~ t ~ac i ,rmiilt forrr~~ 19ofi I'~'t0). 1: d t , i I ~ , p ~ i ~ ii ; ! : . 17 Class Stropho~ncnat,~ Willinm~ #,I. I Ylh cat !.or cc.rni Where. d u e tc9 ~ndrtYrrt-nt prcscn..~tiorl.o r lack of CJrdcr Strop11r)nicnid:i a p i k 19.31 , t ~ ~ clrlgcc rl nn up-tir-darc 111onopap1i~~- J ~pccieccanriot S t ~ b o r ~ iStropi~onicnidir~.~ l 'U-1 b.lcc, rr apik 1 111~1 CI<I# ot. hc i d r ~ i t ~ f i c dV I ~ ~ I ccrt.t6nh. it rn.7~ clcs~q~areci Si~pt.rtI~rnilx- c t t a n ~ b o n i t a c c ~ hc J, Pl Joncc 1 Q F l : 1 I1 I l . l S ~ 1 ~ , 1 n t T (rclntcd tn) nr i f (rnav br. i u n ~ p n r r d with). an Famil!- Soowcrbyclhdnc Opik 1 ) 7 4.0 I I rLY<lr cslfring .;pccit.v ( c q. .\ l~lritlpnptrts cf. t~o!~!~vnnrrr). Si~bfi~nuly Soxvcshycllinac (3pik 1030 1 l'.~>TIL>ltlK Whcrc thc f r w i l be ~ d r n t ~ f t c d bclotrgnl: ro a Crerius E ~ ~ r i r o n c r ~ ~ 29 17 a.c ILccd ~prlIly L J ki~own TL-IIII~. hut L-.I~~I-ILX7r .~sirbl~rd .L spru~cs I to (as S p c c ~ r s ~ d v r t ~ a i Rccd 1 9 1 7 l l l l \ rc.llf~ ma\* l ~ c tlir c n w nhi.rc prr-<t.rvntion 1% pilor or ~f Subspccic\ <crrlrrli< H ~ r p c r 108') i lr,lv that onlv a fr6~g-n;-nrcnt prcscn~ccl. ilic ~l~fTi';iu ( p l r ~ r i ~ l 1s sp. l ~ ~ ~ . ~ ~, ~ p ) ~ lt ~ c - dr . ? . (,'8~lt~it.til~ If the cprcimt.11. In n 111 tlic abovc scctlon wc Ilnt,c WCII thc d ~ v l n n i ~ ~ ~ l-tllstcr , 15 ( ~p.). o morv rvtrrilic c,i<c..c.ln onlv terit.~t~vt.ly rrferrcd thc taxonomic liicrarchy, hut in dcfinlng thc5r be * ~3t1ldbt. I to .I ~UII". n n c wnl~lciIVTYCC e . ~ ? K I I ~ O [ ~SF.IIf y - o u p i n ~ . ~I O ]low do ta?ronorni\ts nctunrl y yo ,111outrtl Nr~merica 0 1 1 1 ~ i = h ~ ~ C C ~ C Y C ~ ~ ~ I O ~ L ~ I i ~c 211~ ~ c t i p t h o n,IS < ' l Thc bacrc principle tq that niorp11ologic;rl ~ t arhcr l e i <4?11lc'rnr ~ d Et~pl~trfaritv;r,~ 7/)rr/l,ilkr1irj rrr~plitbr uccd. ~imilari tie< rcflcct phvloscnct~c ntfinit); fiomol- I T l r t ' t t Tlncc All t.>snnnrn~c ~vtrrk .ruth as tlriq Irlilht foUow a clgy). Tlik is always so trnlun, for othcr rcnwnr. t 1t>\v l>cqt t pnrncuI.~r of n~les, cct whrch h a v t bccn woskcd out s11r ulnrity rcrults fror~luonvcrgcnt cvdution . Hut I n I l ? > V flcrcl t t by a SCIYCC of 111~ P ~ I S ~ CC,IIIIT~~\SIOIIF I O ~ I ~ ~ and arc a5sccsiilp 'rin~ilan tics' hot\* doc\ nnc d c c ~ d c~~pcln 111111-c i ~ l i p d o c ~ r n ~ c u t cin hll 111 the oyrnllly paKcs of cac11 which cliaracccn arc in~portantiHow h o u l d tllci d ~~l)tC1l~1 11lL-2 ~-olun,~c rhc -/'t.,-nrr ,111 I t : r,t'rtc*/rr~~rc n k a n ~ a l q y(a bc c h o ~ e n to miuiinize s u b j c c t ~ v iand prc>duc~ r~rrn~rric~ of P ~ t contlrlulng wnec ni vol~uiirs p ~ ~ b h s h c hv the natural order h ~ o u p c ? T h c ~ic n o ~inivcrf.~ll\ l'he tllirr d Gcolng~c~l Socicw iri A~iirnrn). accepted rlrttllod of E ~ c i l ~ t n t i n g thccc cndc ,111ri tnnc. empl t.lxonolni~tshavc uwd diffcrcnt m c t l i o d ~ . rtccnt ~ r i . ~ n i ! > q t , 11, In Taxonomic hierarchy ycnn th ruc f l ~ ~ r p l y cnntr,xsting ~ ~ l i o o I 11avc :Ill. distribut < Althntr~h 13 tnsonanuc catcgorlrq ahove tl~c 3 spccj~*s c n ~ c r ~ thrcc arc rht. schnoh of ( I ) c.vnlutio~ian !>Ills i t i\ d: lcvcl .II r to samr c s t r n t art~ficlal 2nd ~~~hjecltivc., taxunonly, ( 2 ) t ~ u n ~ c r i ctaxonomy and (3) cladlqn~ FTS tcmatic al ~ d e , ~ l they shanlcl .IS f ~ ns ~ O F F I ~reflect evoluly r ~ C Ullril the 197115 rnoft palncorirolopsts, crprcl,~ll\ l . l t ~ ~ 7 l ~ l n o n a y relation\hrp.c. tl~osc wurkins with fowil liiatcrinl bric h tlicy h ~ v c , i p p I ~ ~vtgr ii similar speclrq ,Ire ~;.roupcd III genera (rinp~bdr rallcctcd in thc firld, wcrc cvolutinnnry tasonn- rr~dCracrnf genus). ycncr"in families. t;lrlul~cs 111 orders, miqts, 1n crccting ; hicr,~rthicnlclnwificntiou, circh t1i~v.c' 1vct.c I o r d c r ~ rlasscs artd clnrsrc I I F tilt Iargrst C11~1siut1 m ~i ilasqical tasonornlsts lircd a tr,~d~tinnnl very flex- ,hftlllt, 1 1 p and ~ thc an~mnl klngdn:~~: phyla ( a ~ y ~ l a r phylum). ihlc ~ i m i h i ~ ? a t i u n trlrcnn. Fint thcrc i~ morpho- i.i olvcd. c l , ~ of
r7dt~1ril Itct'd
6 [

t a t

02

Divisions of invertebrate paloeontology

11

logical (nr phenetic) resenlblance, thc c s t c n t to Kir thr 11105t etTcctivr tri~th~>cl ~ C C I O I I F ~ T Itin? for IC phylogeny. S l ~ ~ i t l r 1994) c ~ ~ l ~ n of nrladist~c ( '~ r n ~ ~ r r 1 ~ i ( . I l ~IIC ,71111~1;115 ~ L Y C ~ I ~ J ~ C ,~rlotlicr Sccnnd, ,111~ ~ylageacticrelationship? .Irr ~lr,iigwith phc- nlcthodology i ~ c rn L-c>mprrhci~sivc that mJv hrlef rrc r ~ ~ r c r n h ~ .corlc~dc.rc~I ii~c~ ~~nport:iiit, thlq is commc.nts ,trc ~ ~ v hcrc. By c u rhc 1v.1~(2s h r .I\ c,~rl bc d~.tcnnincd) in Hcnirip war of rllc aysinlnn that rccciicy of coluI l r 11 ,~nlnl.ilq nrr a c t ~ ~ . ~rclntcd to C A C othvr, 1.c. Inon o r ~ g i coulcd b c ~ hc choliln 11). rlir sl~nrrd llv ~ i~ t poctrnnv nf rcccncv o f c c m i t n o t ~~3rigi11. \vIii~hCIF w ~ snil of cvolu tiom? novc-lnc~ clr 'drrlvcil r urw yr.rrlcr intn ~ ~ ~ l l ~ i t i i~~ IrS O I I~ \ ~Thr . ~11.1rnttcr;~. . ~ . i O ~I I ~ Thus in clorcll; rel,brrd g r o ~ i p ~ \vould wc srr~i~r ' i l l ~ ~ c ~ ~ 111 1C7 ~ C k r ~ c o r d, ~ n dgcnOV i0 rot ' ~ l ~ a r cderived c'l~ar,~ctcrr' d (synapomorphics) ~~pllicnl di<trihutlon niag play an inlp\~rt;\nt pdrt in tvhiub would d i s t i n q ~ i ~th ~ i x grnup fi-0111 othcrs. l 4:!.1ditij: rdanonshipf 7'Ills pr.ittic,~l ;rppronch tn Ficnr~ip's csr~tral ct>ncupt war that in a11j7 g o n p lTrlnnmv,whi~-h took . ~ l l h c t n n into ronq~dcr.~tinn. char~cters,lrc ci thcr 'prim1 tivc' (symplesiomor1.11 for n Fni19 tlnlc I ~ P T ~ backhcli~c the nf p a l ~ ~ o r ~ r o phic) or 'dcrivcd'. TIILIY vcrtchsatcr hnvc lurk211 ! lac,rf cP:i<~it;catinn, anti ir 5t111 C C I I I F I C ~ ~ *tnChc rhc bone<; rlls pus<cssion of ;I bnckhnnc t~ n pnirltrtve ~ '~ .r,t n~crhnd strntnphenrtir pnl~ctr~~tr>logi~tr. chnmctrr [or all vertt.bmtus ~ ~ i rl not, o f COLITCT, h~ whn d :.'Ir.c much ci~~p1in~i.i t t ! t l r 111 st.t.kiiip .lnuu.;toron indic~tive of any c l o ~ t rcl,ition\hip bctivcrrl a n y i cndniir rclarinn~liiy~ : (Hcnn;, I QS4; Gir~gcrir-h. gmup o f vcrtcbrate.;. What is a pninitivc chnractrr l l'h1) fnr all vertebrates i r ~ 3 C O I I ~ Co dcnved c h a r . ~ c t ~ r f ~i Itlr toillr t n x ~ ~ t n n i i \I~owcvt'r, ~ i r r ~ c n . i i n h ecot~ipdrrd to i~~vcrtrhratcc u~l13potnorph-y nnd t~, tJrc ~ I! %t1h,jr~tlvit7: l ~ l arc~ ~v c l nlnicxt m c r c . ~ p ~ t ~ l c in any c v n ~ p l e s i n m o ~ hthilr dclinc.btc thc rclntlvr xt.~t~is v IJ rliil~r<iliration P C I I ~ C ~ be particularly acute o f p ; i r t ~ u l a rcharnctcrc wirh rcqpcct tn .I fprc~fic C t ~ ) ,!,irucal r,lxonnmv. n< dtd zllr J ~ r ~ n t ~ t i nf tllc piwL~lrm. on< ,$rl rccord In tcnnq of pre<rnratinn.Thc nurricricnl 1-lcnnig endcnvo~ircd to providc ,Irl olyectivc .oi,nini<t<rncd tn cscnpc Trpm t l ~ i hprahlcnl by rncthodcrlom for dctcnnining rcccLncvof corninno nlig fur rlt~.it~riI?erI pllcrlct~crcsrr~lblancc ac tlic nnk4n In rvlated tnxa, bascrl upan pritnitrve .lnd I\ rralirtic h~iidc- 11.41 11~~11r 1 3 u p i n g . W A F tIici~ rlrrivcd ~Ilarnctcrs.S i ~ c hr c l : ~ s ~ o n ~ h i y ~ to ~ It ,Ire csprcwrd rv that BC cnc11igl1 ~ 1 1 . 1 r . i ~ \vcrC r n c n ~ ~ ~ r c d , J cladogram (Fig. I .3), I r ? n . h ~ c hd t c l l o t o ~ n o ~ ~ t~~s in ir~tificdand cnniputcd . ~ n d rrprc.\ct~cccl the I I ~ C hlallrhing points Arc arr.+iigcd in r ~ c ~ r c d bv hiernrrlucq. r.lartrr staticnc\. thc d~ctanc-csbct\~ccii clu~tcse Hrre tasa A and I3 ~l1~1i-c uniquc cornmor ,I IIIJ bc uced n\ ; i i i c a ~ i ~ ru f t h c ~ rdiKrrcnccr. nnccstor. They nrc +did tu bt. sister groups. Thr): I e ~rn~erical taxonomy 11.ic bern fuilrld very uqrf~ll hnth ~ h a r c n evoluliorlary novclq clr qyi+i.iynrnnrn nlmc inrt.incec, but < ~ r h l t . c t r cannot bc clirniv~~ phy. nor possessed by taxor1 C. Now C i q the w t c r * ~ ~inct. opcmtor l ~ i su clloosc (~l~l?jcctivclv)group o f tllr conlbincd taxa A and R, 2nd D Ir rlir d the t ljc~tto nnnlyc the ulraqurcmcnt., iiinrlc, dnd slrtcr %qorlp E c o t n b i ~ ~ c d A. R and C:. In pel; o tdXa nwrl to ' ~ i t - ~ ~ h t ' glr'irlg c-t'rt;lir~ l ~ a r ~ ~ c t cfclrn~ing cladistic aii,alysi<, chcrcfnrr, A tax-otlorniqt thenl, c n a >rc irnpartnncc (ag.1111 s ~ ~ L ~ J L ' c ~ L Y ~ I rlrclcr to in ~ ) assurncq that d;vl~otomou~ splitting had occurred t,lln rneanlnghl rccultr. Hcncc the olljrctivity nf In cnch lincngc 2nd c o ~ r ~ p i l cnn (tinwc~gh:hccd) r ~ncrical taunnorny 1% less th.m i t r n ~ g l ,~ t p r ~ r . ip TIIL,tllrrd ~ c h n o r ? l l e ~ l rlporl phyloqciict~c criten;~ I I I I I ~ . cniph,~s~zing icaturcr rliarcci hy c\rganisms that miftct, in narure, a h l c r a r c h ~ ~ l pattc-rr~, cvidcnt 111 d~~trihtitinii U I I R ~ I I C C C ~ h ~ l r tnrnonpt o r p n of ~ .3 Is I F ~ ~ C ) \ V I IA< dadism or phylagenetic rntematics - n SCIFQO~ t'o~~ndrri tllc C'T C M ~ R I ~ hy sfl>rnolopstWilli Hcilrug ( I W C I ) ,i~ldWJ+ \ O C ~ ,rlllrd v r ~ m r o u ~ lto pnlncoutc~lnp(c.g. Eldrrdgc v 111 Cmcmft. l Qt;1)). 111 t l ~ c t.drlv d a y of cl.~diitlcc srr. wcrc iiinnv d n u h t e r ~(ir~clitding,q I havc tn a m m thc tpresrtlt ni~thor). ~~ . Lj~lt, thc nrcthod itrclf <IF , ~lrcd, ~ . E ~ ~ F ~1I1C S iaonlr t u bc rccop1?7c~il! h y C ~5 A r Figure r 3 A dadogram (for s~plr3notion text) see

12

principles of palaeontology
character datn tnktns. T h e r31urc c l ~ n r ~ c andr ~ opporcd by k d l r y , 'E .'r%_S). tlic succcssjvc ay-pear;lrrrt t~ cannot bc denied ar an eqrencharacter stat<-T tl~cre JTC availnl.lc the larger the o f nxa in ~tratigrapliy databnsc, and large dLirabaqcc arc. ~ifrcn routtncly tnl qource of d,7t41, hnwcver itnprrfect tlic I ~ W I ~ ~ ~ O C C F S JC ~I the conctnlctitm nf a uladognm ~ record nctlially 15. T h u ~ C;lnpmsh ( I 'ju0) conia5 sprcdrd up by u u n g oollc uf rcvcral cornpurer pro- mcn ts, 'Time i q a fiindarttc~ltal ~ r n e ~ ~ in~ cvuluJ r on gtanis. T h e PAU?' (Phvlogcnctic Analvsiq W i n g bonar)' ~ t ~ i d j ~ ~t, 3 major god of p . ~ l a e o ~ i t c h o ~ e ~ d Pars~n~nnv) program. fur C S Z I I I I ~ ~ C 'is .1 technique vhould ccmtiilue to be t h r sti~dv rhe diurrtiiii~-at~o~~ , of tvl~ich1nakes thc i c u w t a~sirmptro115 or~iering nf rl~njorg o ~ i p s rclarionshrp to gcolo+cal tinw LII :I in So. hnuing can~tructednn JpprOFnate ~ - 1 ~ d c r ~ 2 1 n qec of nbsen~nt~ons How can we d i ~ n n g i ~ ~ rvrnple~iornorphil: thc ncst ~ t a g c rsplonng rc!~tionrhlpsrr to ram11 111 l blc).;tratlgraplllc r i d r ~ .Fo: (shared prinntrve) from ~vi~aponlorpliit (shard hine this i n h ~ m a t i o l w ~ r h c dcnved) cl7ar;lctrr s t a t c ~ ?T h e most ~ l s c f i ~ l w3y I r thic, Smith ( 1 944) d i s r ~ ~ ~rsne \t h n d o l o p 111 J c t ~ l l 'out,gmup cornparnun'. Here ;In '~ngnup' (ofwhich The result is n phyloge~~etic trrs, wlvclr ' i l ~ o ~ ~ ~ s tllr the r c l ~ t t n n ~ l ~ arer ur~dcri n ~ c c t i ~ d h o n<) <pccifr- ~plittingof I1 neapcs tl~rouglltime and n rii2ctivclip I rally de<ign.nated. and coulp~rcd with ,I closely 'a"be~tertimntc" of thc tree of Ilk'. Very co~lunrn~lrclated 'outg~ol~p'. character present In a vari- thcre i~ at1 csccllcnt correcpniidrnce hr.h\~r.cc~l Any ti11 able state in the 'ingroup" 1rlu5t bc plesinn~orphrcif c l n i l o p m m d the rock record: on the uthcr ha112 it IS alqo fourld in tire s~ztgoup. Likr\virc. apnnmr- the tomhincd cladiqtic and hiostratipapluc appm~cts may thmw 11p uncxperted pattcms. phlc characten arc only prrxent in thc i n g n u p . Her~ruq diqtln,qi~rl~cd rhrtrc kind< nf cladiqric T h c ultimate problenl, not ordy for ~!ddl~111, blii gouprnp. Monnphyletic g o u p s contain the cnm- for all tasnunmlc methodology, result-; fronl con~ mon ancestor and all cif it\ rlcsccndantq (D, C , R, A vergent evolution. Kescm blanccs 111 c h ~ r a c t c ro7 F in Fip. 1.3); paraphyletic groupr arc derccndcd chatactcr S ~ J ~ Cmay have nothmg to do w i t l , from n c o m n o n aricestor (u~uallynow cstinct and rrrcncy of conlrnan o n ~ n but hnm convergeilcc . l~ known as the Ttcrn group) hut d o not include all and it rrlny not always be possiblc ro d i ~ e n t n n g tl~: ( derccndantq (H and C . for cx.~rrrplr,111 Fig. 1.3); rcsultq nf the avo. T1111sW i l l n ~ e r I Wr 1) and lWoor< polyphyletic grnups cln t l ~ catliur hand, arc t l ~ c and WiIln~er{I 997) i l l corlsldcril~g rclat~onrhip\ the result of ion\7ergent cvolu hon. Thcir reprewrr tah c t m ~ c n major ~ ~ ~ v e r t c b r agrcr~rps J T ~ W C tl141 tc tlves are cieccendud hiom ditFcrcnt ancc+torc and 'c1ndil;tic analyqis bawd on paslrrlcrny w.ill tend tlm hencc. although these may look superfici~lly <imilat. minimize and thtis ca~lccalconr,crgence'. and tlic.1 any polyphvlenc group compnsmg them rs attifis~nl. contend that c o n v e r s n c c at all level? i* tllr wort A cladopam I < not an evolutionary tree; it iq an important than has genernllv been believed. T l ~ r r e I( 111ct11nda n a l y v ~ rrlatto~lslupr.AF S L I C ~ I t xc 3 valt~ahlcand ccrtainlv ,I problem here. Even so, clad~stlr of i rigorour wav of \\-orking otlt and rhowing grayhi- o l o p . coupled 1 ~ 1 t h bjostmt~gr~phv srerri.; to be a h t callv hozv orgmuslns Arc rclatcd, arid ~t forccs taxon- hesr way fonvnrd, dlld 3n t'q~elltldprcrcquisitc iiv omiaq tn be expl~cir about pattern\ nild ~ o u p s Thc drawing up nienilin@i~l . pl~vloqenrtirtrees. rnethod~lo~gy cladlsticq 1s cyx-cially good wllcn of dealing with discrete groups wxtt 11drgcrnorpt~olqiUse OF statistical methods cal and srraripaphlc gap\ u l d to rhcsc It h r i n ~ the Inev~tablyp~lacoz~tolnpcal tdxullclnly cnrncq .I errpntenttd fur re,d ol-ycrtivity. A rladogram s h m n tain element of sub~ectiv~tv since t h r ~rjforlr~atlor~ sh nut give a con~plctr Iloiv s~stcrgroup$ arc hirmrchicnlly rclarcd o n thc coded in f n ~ ~ i l j z e d e k ~ O C F arlin~.ilrt h ~ hasls nf ~11ascd-rlcrivcdhotnolcr~ur.but although it record of the stnictwrc and hfc of ~ h c pom-avs t ~ x o n o t n i c rel,1tionship5 in ccms o f recency borc them. There art. partlcul~r o m p l i i a t ~ o ~ that c is of conirnon or1Si11, the order of <l~cccqsion the cnusc troublc. For iiiqtnncc. ynlacot~tologi,?cd in txxnnrock record 1s not taker) into ,~ccuunt (though only can d o little to distingir~hsibling species implic~tly l a d o g ~ r It~~ ~ t.I tlmc JX<O. r nv . \vhich look nlikt n r ~ d live 111 the sdrrle arca but canSo wherc doc? t h ~ lerlve thc potential contrihut not ~ntcrbreed.Polymorphic species. in \vli~ch a o n of qtratipaphy III rrconstnlchne p h y l o ~ r n y ? Inally f o r n i ~ arc present wit11111 u11c b i o l o ~ ~ c . \ l Wh11~t3 few 'tmu~f~xnicd cladiqtr' n c q t c the mEuc specie$, may likcwisc 'he I~ardto sprc-iat~ corrcctl\ of the fossil rccclrtl ~lrogcthcr (3 view v ~ g o r a u d y In pirticular. wlicrc sexual dimorphism i~ stroll;

Divisions of invertebrate palceonblw

13

C r011

ctc
[I.*(

b; 111~lcx,!nil fcrr~nlcr d the onc spccics m ~ br 50 r v ancierl t ~r~rnnisrns their cnvironrnent~l context. in t l ~ r t ~ n ~111l a r ~ appcnr.mcc t h ~ tlicl: havc wmctimer All animals are adaptrd t o their environment in all t 111'1 1 dcrcr~\~cd d i f F ~ & r ~C C I C F ..liarl the true si ru1 :IF \~ l~t of-it% physical. chcmjcal and biolocjcal asp-cts. Each . I ~ I I + I ! I I ~ . I \ IT h,~rrI tn d ~ r ~ n t a n ~(a? with rpecies I S prrclsclv adapted to a particular ecologic~l lc .11ttlllcvlltC"~: Clllptt'r S). n d r e UI w l ~ i c it~feed.; and breeds. I t 1s the t ~ of k l ~ Il'llci~ ~t C O H I C ~ t o t h ~ l l ~ t i ~ ~ c rn ~ iCiI C I T F I ~ palneoecology to fuid out about the rlarure r,f ~ I I P F C i if r,,!irrIl F ~ L ' ~ C F ,I I ~ W ~ V C T .tlicrr arc a number of qtaI adaptanon$ 113 fossil orpnitms and ahnut thr rclnt~r::\.,~lt i t E ~ ~ T I A V11clr to p ~ t 'hiyhrr drgrec of tionqhips of the nrun~nlswith e ~ c h In t .I otlier ,ind tllc~r environments; ~t involver the e~ploraaonrlf both IT, tnr c.u.~tnplr, c:ln he iircd whcn .I wries o f present , ~ n d past ecosystems (ScI~Sfer, lW2) T'IIIIII ~ ~ I C l ~r ~ r t;llli~dtngctlrcr. If n rnllectlon of F Al;houph palneoecolo~is obvrously related to l r ~ L l l ~ q ~ r r i < rntdt from lot.~lity A, the l e n g h / Ir ecosystem ecology, it is not and cannot bc thc s.nnr. \\ dtlr r l t l l l + nr trmlr nrhcr appropriatr parameters rn modern commui~itvecology much ernphacis i~ 1ri.t Iir. plllrtcd on a p p h .I< a fcattcr dlngmta. A plnced on energy flow throusi~die svstcni, but t h ~ g IIIIL, nt'llev iit ( e . ~;I. redr~c-cd tmjor m i x ) nldy thcn k ~ n d determ~natio~l of 15just not possible when dealI\c tlr~~vt~ tl~rougZithe 5c:ittcr. Thi5 givcr a simple Ing 1 ~ 1 t h dead commu~lit~es. Instertd ~ a k ~ ~ n t o l o g i ~ t 5 !. = ;r + q~p11.u'hcrt. it 11, thc padicnt a i d I, thr perfarce murt concentrate oti ertnbhshmg thc cornwtcrtcpt on tlic y ax!?.A sttnilar qCdtfeT fronr n poppnsitioi~ qtrwcture and orgamzation af palaeocom, 11'11lon rollc~tcrlfronr locality R may hc plnrtcd on munities, in attrinptii~g h r n here to work out. r ph, anti t11c rrdncrd major nsns drawn linkage patterns In food w e b and in investigting 1. Thc rclatlvc s l o p c ~ and <c.pnratlans of the autecolow of ~ n d ~ v i d uspccics. al I , n ~ thcn I>c conrparcd ct.lh~tlcnlly.If a ~ Many attempts have her11 made to sunimarizc I h ~ n ccrtaln threshold rhe p o p u l ~ h o n s categories of fossil restdue. to providc the backI ~ c p r t l u d bring of tlir same SPPCICF; ~f ground for iiiterprenng anginal comn~unityscruu.I\ .n the, yxcicq different. ture. The scheme proporcd bv Pickerill and ,. , ~ , , ,nnt. nf ,I ivholr r r n c ~ f pix~ihle v o tern. Hrenchlev (1975) and amended by Luuklcy ( 1 qX3) i? I I I , ~ rnorc clnhoratc t t ' c l i n i q ~ r ~ f niilltiv3n~te used lirre: o :#),I!\$ 1 .jrc. h~sanlrng ~ ~ncrcavngl\Important In LIXlliLt~'llIt. ~ \ > l t ~ tt ~~~~d ~ ~~q ~ ~ v i t . I . An assemblage refers to a ~ ~ t ~snnlple rotrl a glc ~ ~ l v c a ti mlcrocoinputrrs nnd thc pron patticular horizori. pcci~lt\t. wftw,lr~. ppnckngr.i dcqigned 7. A n association refers to a group of assenlblagm, u rncct thc nerck uf pnl,~tontoli)pstq all ~ h o w n g sindar recurrent parterns of spccrm ''1LC; I A I-; Rn~rrtli .mri tl.lrprr. I qr)C): Ry;~nrr composition. lLj()i), me o i n u r n c n c ~ l tlie nnnlvsir 1s becnn~lng 3. A palaeacomrnunity (or f s i community) osl ~ ~ ~ ~ Sr~ti~tic.d d i r ~ l . rnctliod~: I ~ P likewise enrntial refers to an awernblnge, assoc~ationor group of d;tinrny p,~l,~ctlcornrnr~nitil.c. 'undcform~ng' in nssocianons inferred to represent a oncc-distinc;r<lhl~t.~nr>rr+ dcfttmicd fclc.;ilv xr~d of cornpanme them nvc biological entity. Normally tills only repre:,I k9n~ltt'rt.rl mntennl. sents the prcsenrable parts o f an ongitltlal biological cotnnlunlty, ~ i n c e soft-hodicd animal$ are not the preserved, This definitton cerresponds more or less exactly to that of Kauffman and Scoa (1 976).
11 I
(

1'

11.11
111-

es.
111-

:nilur cnreynrlcr, msv hc iricludcd In palacohiulla. palatclr~ulclg(hew diwus~ed with pnlnro\~icr::qnpl~v). fu~lctln~lnI o r p h o l u ~ n ~rlinology m ~ d , :isot wluch scquircs wrne d~scussion. 11

lull
1c:11

Palaeoecalogy

rl y.

\I:,LL rcolog
111,lr

IIF

'"S

r~nlronrnrrit,palac-occolohy i r

the shdy of animal!, i n rc1;lann to the qtudv of

Pataeoecolo~y must alwayi remain a pnrtial and incomplete science, for so much of the infomaoon available for the study of modern ecosystems is runply not preserved in ancient ones. The animals themselves are a11 dead and their soft parts havc gone; thc o r i ~ n a physics and uhenmtry of thc cnvil ronlnent is not directly observable and call only he inferred from such srconchy evidcncc as Ir

Modern 111ar-int. csivirc~n~nt.rit< qradcil =?rcortlarc availahlt-; t h r shvllc iii.it. !lave hcCn tr;~tirj>ortciI .iway from tlic~rnri~qn,llenvirtlnrrlcnts b y r trrrcnr.;, 2nd ingly to cicptli . The littoral cnvlrtonnrenrs af t h e ddl c tt7r f i l ~ ~ ~ l ,IS~C~III-I.I~C~ f01it7C1 m,)v well Fr qhorr pmdc into tliu ' r ~ ~ b t ~ shelf. drld .I[ ~ h cdgt that .ITC miscd nr inconiplcrc; pcr~t-rEcpu~ltiun.~l iii,i~c.n~ticof thc sliclf tllc tontinunt,~lslope gnc5 dowa~to procesrtncni.ry ! i . i \a1tcrc.d thr. cvidrncc. <till fi1rthi.l-. depth; thir is tl-lc bathyal zui~u.L3t.ltnv t l l i c lic tlre ~ ancl t i l e hadal zoncr of the clzcpI )e%pltctlin p . l l ~ c o c c r ~ l c rrcm,rirl\ ,I v.~lld, p,lrtbnl flat abyssal pla~ns q* sririicc. Much 11 Ilrllv knrm.n .~hr>litthe poqt- occatr ttenches.There i s after, a pronouliced zc1n.in~ortctn hiqtory t 3 f /Irmlili rclri.lln.; (t:~photlntny; tlon uf llfc forms in depth zone5 Irlorc or lcsq p~r~lEcl C I ~ ~ ~ C VoT I ~ ) , ,117 ,~dtlrti~~r~;ll f ~j'l~ic-h ' rlitt~en\intr with tlic sl~orr. I n addit1011 thcre is n ~ e n c r , ~ l 111 ~nvnlrcr h u r i ~ l prvcc<sc< (biostratinomy). Tliix dccrca~c abundance (number of individudl5) I7ut 0111rlpq to rli~rrlt,lnglt. thc \Ianoor f ~ ~ t rc\pon~iblc 1;ot r~rcrrsarily diversity ( n u n ~ b r r specie<) 171: o r ~ for d e p o ~ ~ t i n nf .I ~.r.lrttcul.lrfcj';\ll .ir<cmhlagc, s o dcscciit into decpcr {varcr ho111 t h r edge caitlie d~c.ll' li ~a rhnt a~scmblnqc< pr~.ccrvcd r r f rrtrr, which c,ui yic.1~1 T l ~ etautla5 of (Ilc ~ h y ~ a11d l l i ~ d a lre510n~lvrrr of val~inhlrp~l.lc.oc.cnlocp-~~lh r n ~ ; l t ~ n n , hc dl<- ori~tyallycler~vcdfrom tl~oss shallow w.Itcr5 but ill c-.ln t ~ n p i s h c df r o ~ n.~cwtnhl.~ycc i ~ t113i0c' h c c ~ ~ tl tta~lc- are highly a d ~ p t c dfor catchi115 the lilnitcd food avnil.~l>le p e a t depths. TIwsc regini~c. ~ r r Ilowat , porrrd. Riostr;~tinniiivclr prc\crv.ltinll hi\ton. I l n ~but11 ever, irrlpovcrishcd relative to d-tt ~liallnw-water pre-burr21 2nd pakt-h~ln,rl elcrncnts T h c fnrnicr rcgi on\. Anlin~k rit~d plmit~that livc on rhc sea hour aic ~ncllidr tr~n<port.ltloii. pl7y\1c,iI,ct1cnl1c.11n r hiolt~yt i t ic,ll dot~tngc rhc shell, and tlic ,lr~.~chrncnt cpi- benthic; t l ~ o ~l c ~ drict paw~vclyo r slvlm keblv rll rn nf fauna. Po~t-hunnl prr,rc\5c.\ m;ly l t ~ v t > l ~ ctlrc water crduini~ arc planktic (= planktonic) . dirturhn~lcc t ~ ~ ~ r r n s r;~ rn n hy ~ .d srdlrrlcrlt c.ltcr\ (bio- qmuc tlic): are the plankton. Nekton (nektic o r turbation), currciit rcwr>rkilly. colur~or~ or11c.r nektsnic 'CNIII;~) 011 the other hnnii coialpnsrc actlvc .~ntl cwlinnlcrs. Nerltic anitnnl~belong to the ~li,rllou di.~pcnrtic r ~ ~ ~ n ~ n t~h.111gt'~. p i011 watcrs nenr land a ~ ii~cludcdemersal rleiaicntr ~ d and Modern environments and vertical distribufion of wlilch livc above the coi~tineiltnl~ h c l v c ~ fie+ on bcnthic arlir~lnlst l ~ r r r ~ Pelagic or oceanic r~. animals ivater? or i ~ ~ i d ddepths 131 lc Figwrr 1 .4 shnirvr [ h r matt] rnvlrmtllcrit+ w i ~ h i n tilt' Lunaq inhdbit t h r surf;~re the upcri ocuarls: bathypelagic arid thc u w a l l ~ Earth's oceniir n t rhr pr~*<cnt ; ~ n d r1.1r tlic ntlrrlcnrlnturr Sor thc dirtnl.utinn nf rrlarlnc. .iiitmaI\ within bcnrlilc. abyssal .rud 1ind.il nrpnlv-tv 1n11;lblt thr. p a t dcpths. t h r nrt.,~iic.

( )lilt' tl tIl<lllyprrj cr.4 l l t i i ~ n El\, k3cl~cll a11c1 rlowl ilrlally L O l~clotv the

r11~. ocL-;l

O~LICF~ FCC!

.I?C.

.1211c\

# 1 1 l ~, r 5 0 l1 r

rhl-re art. i anw upllti What I

thc*tc.fiorr pan, nf tli rrwnn of Irttnmf. Ing ~ h c l freal toward< rl

Ivi 1 s ~ 1 10
nr \vhrrc

Horizor Thr. 171~1iil ticln of rec tlnrurc idIcncc. T1 I4

rtr.cnn\ if I tl~c ijthcr
c c - d l ~ . Tro

tiunx. an
hvr the
ill

rcl.1tivcly rl1c SPCClC ti~drvid~ Snlit11i-y

thr>urnnd. 1.c. confir few arc e

~ n l i t i i T. ~ ~ l > ~ c 3 l 'l \ v . ~ t r n;Ire r-~p~md y

(

1.2I12rc1 31-

Figure 1 4 Modern marine environments. A, on a drawing in hporte, 1968.)

0 and C in the met refer to supralit+uml,littoml and sublibml environments (Based

gcCt1cr:l car rhc J u ~ t s s trlrtitq whi pninrrr t o tvhich the.

Divisions of invertebrate paloeonbbgy

( j r l l \ the sheli atld <lcqw envirorlnlclitc Arc nor1111 prcrrn~cd111 tlic gc:c.olo~~cnl rccord. r l i r trciic.11 L 1rl.1~ Tile ~byccalplain5 .lrc undcrlnin FO Ih mck. I'nrnlr~lat tllr tliid-11~raii1~ lidp-s 111~ rnnvlr~q J ~ ~ P ~ LfI r c ~ ~ l V then1 to hccoinc
111

15

Woter turbulcncc Irl.ly rrrrcise A s~iLwt.i~~ti~l (-011tml ovcr di\tnbution. and tlit- c h ~ r a c t c r ~ t;iur~,~\ nf tn $1 hi#- and low-etier~y e~l\~i-rorinlentq oftcii vcrv Arc disp:rmtr. Robust. thick-slirllcd and rouiiltcd .;pr.c~c~ arc normnllv .~ihptrd fclr h l d i - e ~ i ~ c~)rid~rior~'r, 'r~~ ' I l ) , l ~ r x L ~ ~ I \ L I I I I . ~ tt l ~ rS U ~ ~ I I C ~71311CS ~I V I I I ~ C~ I O ~ \viicrcas rhin-vhcllc~i nnrl f m ~ l c forrrlr poitit to a l\\*l (I\\-rlw <l 'ccnl~rc trcnclier; ~t lrruvcc ,I$ rigid p l ~ t u ~ .tr~uch clliirtcr water e n - \ ~ ~ r o n ~ r i c u H. i~t n u y LJC J t T 1,- 4 bi-L'rl I! hn\lrnc ~ r r . c r v yo lit^^ ~ c o l o y c n l l v , 1 r tllc po~rihlc infcr n ~ u c h to nbniit r ~ l ~ ~mrhulc-lire In a t~ve n::i(I . . . \ L \ t i 3I...rirntc k1imt.11 tlierrirl bciup uf Tridwc 1I hwil envIronmcnt merely fro171 the type of- sliclls 1 r( ~rr.11nw , ~ p p r o ~ c l i ~ u q a ~i11~iiii~tiv11 C t h ; ~occur. Z O ~ t rill I to bt. cunrumcd \ v ~ t l i o ~macr. Henrc it 1': .w fm- I I I ~ ~ C ~ ~ I ;ihvs~ds ~ d ~ t l l e n t MODERN AND ANCIENT COMMUNITlES of' C ~ H ~ 1 Jtld Pi1 I ~ CT)?ltlllethS. P In rhallow cold-tcmpcrat~ire sew I ~ I J ~ I I ILI I' V C ~ I C ~ prr.rt.n.rd 1r1 the geologc-a1 rccnrd i c brntcc x u nonnally foz~nd 111 recurrrnt eruloy,?cal ilv .i finctloii. .~lbritthe ninst papuloi~r ~01linit1nitic9 awnrI;qhonC. \VI~ICII ,ire irsz~allv OT subIrl, rll I F ~ V I~iotlt.re,rlin of anclent ~iiiies. Thc sedi- str.lfc sclatcd. 111 chew 3 pamcular set of sprclt.\ I I L ~ T Irji tFic ~ . o r ~ t ~ n c r ~ t ~ l ~T <l-~clf il~zludrthose of tlie ;ire usually found rogcther ~~~~~e tlrey h ~ v e v n l c thc 1.rclr.11.~gnnnal, I \hallow ~~tbnclnl. rncdlarl arld r~tttrr l~abiratprcfi.rcnccs. wit11111 tl~esrconinluilitirs ~ l i r '11.1f n-?lnw. C;cncralF~. ~erlirr~erits hccoiiie hncr .inimal~ cithcr d o not conlpctr d~rcctty, hein: kt\! rdscs of the <lielf. thc rr~tlddierrrponq adnpred rn r n i r r o i i ~ d ~ ei~h ~ nrhr Farrle h ~ h i ~otr . w t a '11. r.T17crc niav ire reef5 clcrsc to t l ~ s l ~ ~ > r ~ J V C qtnhlr predator-prev rela tiotlsh~p. r c IT I %rrF .I prono~itlceclbreak ~n lope. I Cornn~llr~iry qmlcturc is norninllv wcll detii~rdin cold-tcmycraturt arrnq, l-rlit 111 wnrnirr scar wllrrr Horironbl distribution of marine animals divcniry ic higher it IF ~ e n e r , ~ lless clcar. lv T11r m;un rontrol5 atkcrinp the hnnsontal dtstnbuPctctrcil (1')lX). working o n the t i l ~ r l ao i thr ~ rlpn nf rcr.rnt . ~ n d fnv11~ i ~ ~ rare~teniperaturc. thc K,-lrsCg:lt.tint ctudled and dcfiilcd mrrie of tlirkc rt.it~ alg 11 m r r tit r l ~ I I~J [ ~ s ~ s ~ 1 3 n l.~and w;ltcr t~rrbu- ur,~lly ~ T , I ~ ~ j: occnirring cnmmunitlrs. Hc 2 1 ~ 0 rrcnipizccf 2:nru T1ic Inrgc-~c,ll~* diqtnbz~tin~l l i ~ ,111lin~l~ tllc 111 two c ~ t c g o r i c ~ bottom-dwcllinp nillm.rls; infaunf trcans is largely n f~riicriniiof rcrnperature, whcrca5 nal (Isuncd and lnvinK warhln rhc sediment) . ~ n d : I C rvhcr ihtrnrq gr%ricr,~llv nprrarc nil J illore local epifaunal (Irving o n the sea floor o r nil I-orks or ilc T ~ C I F ICT .Y ~ ~ ~ ~ C Trcgnus c a r r y the inost divcrsc w ~ w c c d ) I t \\la% ~ o o i f n ~ t n d . i that parallel ct)llimuni;%tncrr, .~nctIn thew thc E ~ C C F P F .ire vrn. uil1slzrau.i tics ucc-\IT,with thc wmr senera bur nor tlie h a n l t . lllrt t l i r nurnlwr of ind~vlduah i JIW c\nc spccics if <yscies. r)n thc. oppodre ndc5 of the A t l , ~ ~ ~ Since o he. r:lari\.clv IOIY. In trmprmtc through tloreal r c p o n s tli~spionrcr wnrk a wholc sc1etice c)f c o r ~ r r r l ~ ~ n i q ' r'lc 5pccicc divers^^ rr icrv, t11ou~E1 thr rlitnlhcr o f ccology ha5 g-imn u p , h a v i 1 1 ~t 5 counterpart i l l i ~,l:v~dual< cpri-zrl; r n r l bc very Inrgr. pcs P B I , I C E I C C ~ ~ O ~ ~Much cffort hac beell r q ~ ~ 111 ~ d . d %Tlrliw 111 thc cc.1 ir of tlic order ut'35 parts per tryrng to uudentand thc cornpmition of fcwil ~otl~and. Mmr niannc a n ~ i n ~ lare stenohaline, cc>i-r~munities, hnbits of rhc animals currlposing c rhe rnrltinctl M w ~ t r n near-normal salin~ry.A of thcrn, communitv w o l u t ~ o nt h r o u ~ htililc nlld. .I$ .,re ~uryhaline,r c. vcry t o l e r ~ ~ ~ reduced of t Tar 3q pw~ible.the controk acting lipon tElc1n (erg. I1rr1n2. hr~ckivlirvarer ravirnnment ic phyrir,- Thnnon, 1'357, 1971). This is pcrh,qw rhc nlwt Thc I~I<IL , 1 1 1 ~ 'difTicult', 3 r d Liuilar Ilrrir~pi l l brackiqh Jrtlvr field oFpal.ieoecology n t present. J ho\t of \\ ,lrt.r< arc nnnnally coii~pnsed of very fc\v sprc~cq, rccent o n g n a l works tcsrifics (c.g. Craig, 1054; *,pel-~.illyh ~ v a l atid ~~wtropodab c l u n g i n ~ 5pc~c to Ziegler ct id., I9hX; Ho~icot.1975, 1C)XI ; Scott and ;I 1117rd .ITICI nficn long-ranprtl Eencra, r h r s r s d ~ n c Wet, 1976: McKcrrow, 147X; S h n n c r rr I(JS1: tl?r,l can hr found 111 sedlnlentary rocks as oId as D(ndd nnd Statiton, lC)CJO: Uost~lcc. a11J Alliwn, rlunnlz. 2nd thcir occurrrncc r particular sc.41- 1995: Rrencl-~Icy n and H n q ~ r1007). . enrs n-hirh lack i ~ c r m ~ d rn~lntle fossils is n vdluablc AT Fiimich ( 1 977) makcs r t clear, holr,cvcr, most , 11nttrto reduced sallnin. la1 the rnuironmctit in fowl a q ~ c r n h l q e s'lack soft-bodrt.d ainm~lc.W11ci-c r1 lilch thcv lived p o ~ ~ i tbr l ~ fossils C J I I he uced to c o n ~ p c n ~ . ~ t c xe fr$r
1

I(

Divisions of invertebrate palaeontology

17

d currenfs

pure 1.5 Disbibution of modern marine

s w r g In Br~ggs, 1974.)

shelf-living onirnals in founol

provinces, using Winkel's Tripe!' projixtion. (Bused on a

r ~ a m p l r .tlir t ~ p f Florida c~rrir.:;a tropical shclf o snl,>redto r l l t r ~ o l - t l i e , ~ ~~ l d11orthwc9t hy a t wlcr ivat?r arm\, biit tI1ouq11 this is part of the &'orAtlannc troptcal shrPtprovincc it ha? hcen iwtrll for <nIiic time alld ther?fvrc itc huna hiis 11:rgrd t n m e ~ ~ ~ f r ~ ttn tlint of tllc Cnnbbean l~a I.rktn-isr ~ m n e n c r a n l c m i<lns~cis { c . ~ ,the .~[apacol) mny bc cnnstdered as pnrt of a general ~ 8 + c c n g ~ prcporl nr provr~~cc, h~r bitt aq their it'lvr~ have been it~iriallv ucdollrzrd bv charire m l g l n t 5 thry rnav have vrw rnanv cndrmir rlc~.!t.rkt~n hlch have evo1ved i l l nolation. H o w znanv L.fthcrr. cnJeinics there arc slras drprncl larzely on I!PWlong t l ~ e iclailds Invc been i~olateri. Tlw drvrlt~prner~t today's iaurl~lprovinces ha!, of brm chartrd throuphout rrlost o t11c Tertiaq. At pr:rcnt Me~ozoic l ~ 1rIarozorc provirlcef arc. Li kc~ P. l l i c hetng docilnlrutrd (Micfdlcll~ls~ 01.. ct 1g71; $Lm, 1473; tIughc5, l(37.5: Gr;iy nnd Houcr>t, 1 "' ; Mciierrow and Scotese. 1 qclO). Whc-n uced in ).i!uncfion ~ ~ i tpahcocun tiilenc;tl rn,tp.: rt mav hc li )~,ibli ta s r c IIOIV the dictributirlll pattcrrl n f ~ c ~ ch~tnas nt relared to thc pnqitior~ f a n c ~ c .cono ~~t ~ t n t a lma<<c+ thclr rhelvrs. Sc>nleti~ilcs aid palacnI O Q ~ ~ ~ ~ L ~ard~ grnlog~c,~l , I ~ I C ~ I ~ evidence may havc b;.annc upon palaentmperntllres 3 r d cvcri allow me snt'crciice to be ~ n d d e rcprrlltlg anclrnt ore;m
llr

111 11.

Modem and ancienf reek Thrcwtghou t gcalogical n m e anirnalq have 1101 only bccornc adaptcd to particular ctwironnients but alqo thcrn~clves created ncw hatmats and cnvirclnrnentr. With~n these there has heen $cope for a h n o ~ unlimt itcd ccolclgicnl differentlatiot~. Pc-rhap the most striht~f: examples of such biogenic cnvironrnenrs are tllc reefs of t11c p a ~ tand prc~cnt.Rce& ( h g . 1.6) arc marcive accurnulatlons o f llrncrtonc built up by bmc-~ecl-rring l g c ar~d a by variou~ kinds of ~nvcrtcbrates. Through the activihes of these harnc-builders grCnt mound$ may bc built up to sea lcvel, wit11 cave< artd channclq witlwl them psowdmg a residellcc for
RA5IN

SLOPE

AIM

LbGOON OR

SHFLF

reel

reef fac~es

~trrtlt syctenlr.

Figure 1.6 Generalized =tion

through a reef {olgol or coral).

~ ii1nn111cr~iL31~ & c d : I I I ~ II,II<. .LII ~ L L ~ I ~ ~ L ~11fi:r- in!: in ~-1~il11pci i 1 1 tllickct~ r l r l t l l ~ rl-cf w r t t ~ r k11I I -JI~Y \Yihc.n thccc rnrlic<t n i rccC ~tlr~crtc~hr.~tcs h~'c..11!1r riit~ntcclfor tlic~rp.:rhrnl.~rrn~hc>\. t ~ c 111 ttw h r r ~ r ~ 1 5 p ~ t ~ -rrrL of tlic t r o ~ ~ 1 WIS 1 1 rurtnct in late Midrllc C.nnhri.1~1 l ~ thcrc \vc.rc tlcl r ~ 1 li ~~ of t<>ri,l\. ivhlch gt.nw LIP t o tlic surt:~cc., r h r \\),inti. iiiorc r c c f nniiiial\ trnt~lcrlln~'h4) M:I 1.1tcr; thr oi~ll OSV~CTI-TICII, ti~rll~tlt~it 1vstrr5 nllo\v r.rplrl cnlriun~ rrc.f\ \vcrc stlninatolrt~c.In Middlc t lrd~)t,ici:rtr1tt11c tlrc~c ~ l g , ~ c ivcr~.joinetl c,\~l-,ll< rrrotn.lrll.ai~tl ~ 1 1 ~ t ~ l ~ ~ 1 ~ ~ 1!1~71cr ~-~3~ltl1lll~7ll~ ,il!tj ~ 1 1 1 s~(>t\'tll,-rll' ~ ~ 4) pritlc.ip.11 I'mliic-builclcr< ,Ire .11,~.:.1c. ,111d c-orX>l\.y u r pnroid\ ( I I T I I L > - ~ ~ L L T I I I\pi>ngv<;Clli.~ptrr . i \ \I rli l . thcrc. .lrc 111:iny nr.1cr kincls t i ~nv~rtc.lir.itcr thc .w by rcd , I ~ ~ , I L - tvhicll trrgctll~.rhlrii~cdJ rCc.ic.nvlin .I I O reef ct,nHrlLltrjb: \pony.r. b17,070,111< ,111il t t ~ ~ l l ~ ~ rotlrncnt . ~ t t l ; ~ c t l r ~I~ C ~ 01- otllrr I ~ I V C ~ ~ C I M - A ~ C ~ cr'~ blacliiol,rlcts ,111cl trilllhitcr. Fnr M I ~l;nc)x~.t nr~long<tt h c r ~ . n hnmc crf rhcctn .i~id millor w,iv\ to i r ~ ~ l u r l i n g 111 the rrrC t;.iii~t.ivcj~.L; t l r c r ~brrn k jt d o ~ v n b o n i i ~ rr,rccliis this type nf r ? c i rnrnplcu Alrl nr,t c r l r \ t i ~ ~ ~ r r n hlI tirnc. T h c rvrf\ tll.lt ,\row I r l and L T , I T I I I ~ -1 [I? k ~ i l ~ nCl the r r r f to ECJ \ C T C ~ br.yc>iid l ~ t c l c \ , ~ > n l ~ n t ~ cuntirru~ll~' beep? p,>cC ~ 1 t h 1 r~iZ~~rdriicc, ~t 15 nlso tilc ~:arhnnifcmuf cvcrc m.i!~ll!' .tl~;11. <ltl-tlllldlnbur i ~ .III b c ~ r ~rontirlu.~lly cr17dcd. In 3 ~ p l t - 4rrornl rerf pnroidf aiirl cor.11r uo I n n ~ playilly \ u r l ~ Irljpnry coulplrx slit. r e c i l t ~ ~ \ l 3 l i ~ r dL-OTL- o i crrncnit*rl tanr pnrt in thcir ccrn~tsucticlr~. I f T h c < . ~ m ckind c r f reef cunt~nll~crl illto rllc ~lynl,ind rot.11 Z ~ C - I ~ T O I I S Lic111g~:cax~nrrl<. c ! t h e ~ n ;I\ tllr sl~rirtkit~r rcrf 5 ~ 1 b ~ l Ji e iFTOkVF ounvnrd\ nvcr I forcrrt-f L ~ C I P C - IJcnrtint~,2nd rti.lny n f I I I ~ F C ' Fnr1gr~c.d t . w.rc ro\c .I[ tllr cdyc ol oc tmiiblrd buul~l:-rx I>rnLeii i r n m thc rcrf $mint. ~ n l : ~ t ~ d of that tlrnc. TIICY Uchlt~d is a lagoa.711\ ~ l t :Il ~ - n r .~rrttd~ c d ~ m c nz~d dccpcr bnqms in ~ { r h i c l l t h c 1~1tc1. ~-crir>rf~r:~Il\ it ~ ~l nr ~ C C .lr, tidal fl.~t< l o n q \IIOIL= r-olr~~~irrdl ~ thr l n rv:uinl~acrcri.l hccnnlc mnrr xaE~iiv IT c\r.~pn~-.it~C1;~ I \I ~ C 1il.r~cv.lpor,lnr>n in p n i i t ~ a l p.lnr Ilchind t h ~rccK 1' t , ('blur-qrrns'). C;rcrrl ,ily,~c c.oinlr,oncclr 111 tl~ra n-~cv .lrc b x k - r c c t ' t i ~ r d ,11q.1t'.Ire thc I?n.itn Ilmc ~ccscrt-r~ wntt-r clr.~wii thrn11!:11 t t w rccf rlric,tl o u t In tll, rr\. Ingoon$ hclllnd. 111 thy Pcrmi.111 rcrh uf TL-Y,I\ .IIICI of tllr rrrt' rrcrlf w'licll arc vet?; I.irgc, t l ~ crrc. Larqr rt-ris r t ~ ~ .,I\i t l ~ t ,* I I ~ Y ~ ? kno\\n nq bio- nurtlicrn Eiigl,~iiri, l nrc (runt s o q ~ .I V ~ ~ C I C ~ 1li.111 <~.fl.~tt~iii.ltcc~ .I\ ,II n l q l \llcct\ hcnns; thry krill dtsc-rrtc ~ n a u t ~ d q< i t q tironi the rr t ~ ~ r n i n g ar rhc rcr-f rrr-rt ~ v l 1 t . ri ~ ~ C . . I C ~ Ic( IL ~ crrcr f ~ sr,l floor. Biostronres. oil tht. utllrr I ~ n t ~ d . t h t ,Ire I . ~ ~ I I I I C'UTIIIIILFIII~ICI rt'~,f-h-pt' I.I~ of ,11111t1.11~ h,~rt-ly lcvcl (Fig. F .i). 2nd T h c uppcr \ust:~c-c \ \ . I \ fiit~iirc.ly colorii7ctl nw nl-iw~. ica tionr. thc -, tl . 1J~roz~ql~nut qcnEoyi~ h r \ r t m t l ~ t a ~ - t .h:lvc hcrll < r r o ~ r ~ n t c ~ l\ivc~ ~ crllrl r r l cwt tow.irds thc I . ~ y o r ~ ~ a l ;1l ii.lck-r~-t.ft:ac~c\.Tlii\ kintl { ui gci~cr,llIrlorrct.c.lfi I ' J ~ ~ C ~ L I F~ I ~ I L nf F rrct' ~ ~ r ~ i ~ l l i i i t t . I I ~ V C ~ whi~-li l\ .1TlCf?ll, H c ~ L ~ I - I s ~ I .I!IC~ ~~ I C C ~ ~ rI Y tI ?~ C t 111 .111 ~ i t l l pl~olngy ~=L kI i T ~ ~ thr-rcic)rc cine\ n o t ~ l o ~ : c,apyroslrrl.ltc. tl11 hll the frarnc-bu~ldr~ri li,ivc l ~ ~ c l ~ i t i c t l nlyac. hut thc stnnd.1ri1p.lttc.n, 13iFip. 1 .h. Whcn tlir Pcrrr~i.~ri c l l ~~~\.rrce..br,~tc. fr;l tnc-l~urlil~-r< 11a\,r beril of (11 tIl.r-c~lit i c a s h.ld dsird out ctin~plt-tcl!. rllc Pcrnmr,ln rrct!and<: tl~r present cijralq .Ire onlv tlir 117r>.;t rr~.r,t?t111 ccrlliplcs typc vnnirhcd, . ~ n dthr-rc wrhry 1 1 0 wort rcr'ti lintll thc +10\vI ~ ~ p t l t ~o fnthr cclml-,~lg.~l i p rrx., 3 srncs o t r r e i - l ~ u ~ l ~ lalil~n.ll< ~ n r : (Nc1vr.11. 1 C171) hm Thr t>lclt.rt k11~7wt1 C C ~ C c V ~ I -30(111 Ma 0 ~ . qystcrn rlrnt nrrbsc in t r lntc T n ~ n l c .Cor.11 r c r m t ~ ~r O 11 i~lade tntlrrly nC<c.rlli~rlcnt-rrnyp~n~ p ~ ~ ~ ~ l h l v up ,11711 11111.r-srzrrt~ng iv.innh.~ctcn-la: h = p~tlk;kll.'tltl~' r CWOriiatol~ St11112of t t i ~ ~ c c ~fL - ~I L - ~ c C ~ c ~ ~ ~ i ~ ~ c l e r , ~ b I t reef-fldt tes r ~ . d l r i i r ~ ~ ~ ~ Cliir.. 15 rcyorrcd frnm t l ~ rG l r ~ t ons rlavc L,tkc repoil 111 C~nndn.IF b e j ~ i r over 4.50 111 tliirk, ,I~A ~cpxrat1115 rli,lllinr.-\hater c.~rbonrttcpl~tL>;71ir1 I REEF-CORE ,I e frtrrri .I dccp-tvnlcr rurhid~te-filled I ~ . I F ~ I I , r l ~ Arc r ~ ~ n o prcqcnwl ~ J I L - [ ~ I ~ ~111J t l~i\e ~ c > I rt't'if. IIL~WCVCT d ~ i d tlrclr ccolog~cnlsrnictilrr nia\. 11.1vc.hccn VLTV rlniplr. Wlth tllr ricc t7r'fr~mc-bu1ldir1y mct.17oat)r u~ thc Luwcr C,llll't~ri.~r~iiciv k ~ t l d13f rrcf ~-ornmuliin. 3 madc i r i . ~ p p c . ~ ~ l ~ Strotrlatnlitic rerh were Figure I . 7 Crest of u Permian olgo reef ( R d r w n from Sm~th icc invarlcd 1 tllc ~pr>iig~-ltlic. . v nsch.~cocv.irIi~d~,n v - 1981.) qr

Divisions of invertebrate polaeontology

19

rlcd : ~ n Hnri~ ~ l ~chr.cl+ l n i c t h c i ~ (,1 1 t)r.r 111,irr T h o ~ n p r t ~(i1l '1 17) it ha< hccii ~ir~rierstood tllc that ~1+trc\r111~- o ~111 r11c c,]rlv C ~ P ~ ~ I L - L -cu;lL>i-ti~,lt~tl~)I I C p.11-[Sof .Jn!- ~ ~ ~ J I ~ IE$ Stllc~ I pcr~ d OIIG of ~ I 1111 I~c'rt. nn r c d ~ ,lrcktlo{r,n (~Iioi~qIi ~or.iI< rci111toTintcra~-tin?fort-t,<.dirtntcd b p I i y ~ c n n ~ , ~ t l ~ ! LIIZ rccli \!I 111:: $olr~c\\.J~cr~ .it tli,tt tinlc). L ~ I I I . E I I C ~ . I ~ I.IW+F. \v111ch hnvc npchra red rhrtlugl~ I) nralq r c ~ n r , c r c d~ l i ~ j \zc.rc 101nct1 i t 1 p o \ ~ t l ~ . ct*r~tral Thc ~hqut,licrc (1 l~oinn.;. i r d I<c~i. I 1111 ~ . IL C 171, ~ I I C p r c ~ ~ ! ir, ~ ~ t Il L ~ i \ r i ~biv~~Ivc\.I L O I 'J'j.3) i r tll~. ~ 1b;il;mcc. pcrccivt.i.i betxvc~-n 'nccl~lcn cot' tc '. t ~ ~ M T I C T ' : , nllr1r.h . ~ nnc p t ~ .16 ~ 1 ~ 1 l o C tL I ~ ~ I ~ I I I S C C Ih i - r n ~ t i~ \ .11icl thc ~ I . L ~ ~ C T I ~ I I nf I pliv~irnl la\vr, a< OI b i $4 I r rlrt Jnm~ti.it,t rrCff r n ~ ~ i ~ - l x i i lYt't~~~~. L - S C caufc? o i c ~ t g ~ rd ciqc ~ n ' . dc ~ i H I I!l~,(l 11111 111 tllc 1 . l ~ ~ C ~ . ~ C CIC;IXIIIC F . cnr~lq C OLI Diifcrrnt Itlnnnc irrvcrrr!~mtc qkclr.tnn~ nl.ly bc I I! unil~rllurcd :rnd ~ l n m ~ n artc cri - b ~ ~ i l d lirlvcrtch- ~i~rtctirsi~,~llv ~ n~ cc>nvcrgcnr 111 the x i n y thcv T o n . . l~-lll-< ;rntl illc rarnc kindr nf qn\vtli pattcrin turn 1111 1 I11,rc 1115 been <tornr dcc-l~ilrin thc F ~ ~ C . I L OC irrcluc~itlv i 11 rcprcwiitntir7cs nt' 111niiy p l ~ v l , ~ . I rIllll l ~ l r i rwfi .lnd In t11r i ~ ~ t ~ l i ( Iw ror.11 qcllcr.1 q ~ t ~ c ci c brc.:ui<c t h c ~ c ~ C C I ~ T I Y C I ~ fen, or.~vc l l( r J I 111 1vli1ih ,181 , ~ n i t r ~can p n l v ailcl yet cnn ptod~~c-e Ii,ird ,~l .I :h, 2 of tlw. T c l - t ~ a n TIir\ 'irr. no\\ c u l l . till(, Ii~cln-l'.~c~~c rc"cgoii ~ n c l0 r 1 a ~ l n ~ l l c rcc)vcri~lg.Invc.lr~~hr,irc diclctnns. hy cnntr:irt 1\'1t17 of ,<,I'L rrlv #c\t ,Atl,~iit~c.I I <L ~ ~ L - I ~ T III,IY still hc tIio4~~ vcrt~,hmtcc,.lrc gcncmlly cxrcri~nl.:~ndtlli\ CCI I\ K 8s.~t~~~utn;. thC rcccq mctc' nc>t r ~ ~ l u l i c a t l ~ lII,ITTL~\VFLI(w~,iit l ~ c tliirt~yl~ y Cpcctnitll of p0wi-11 pncubilitie\ li; . t:.!1 Ft tlic r ~ l r - l ~ t ~ c ~,n c .l , ~ r ~ c ~ t ~ . cl ~c Tllc liltl~rc.ot' ctrll further. Sonic rit'rhcsc y p c ~ ~kelrtons r r J\ of n ,% l~rll! r n i i i ~ n u ~ ~ ~ ttlir. \ . i r mort complck (IT ,111 & l l ! O \ ~ $ , I I I 11i1ir my<teiu<, ct rcn\.>lInr u L qcrii. v Eutcrnnl flicll\ STn\Ylng a t thr edgc5 oidv bv Functional morphology, growth and form ~ c ~ r c t ~of inin z nlntcrinl nlni~g p.lrtlr~ilar o :I nrar111: i~~nc.trr~nk p a r r t t - ~ ~ orga11.i 111 1;3.;<11< c;llinvt rif l.~~51na\ 2 0 1 1 ~o f s ~ ~ ~ wV c r.) r s f i c ~F~I I T ~ g~mvtll th I?.v~r.11~1i~hi~i hy III,IIIV .?c ?T~LTIIOL&J I ~ . JL L~~ ~ ~ rcc~1ltqill n lcly:~richiii~c A V rp~rnl41tlll fit 111 br,lchio-lil~l~~!y~t+. I < 5tlU ~ l O ~ ~ l l . 7t'3 20 <OIllC \VdY l?llt 1L lr pods, ccph2lopr1d+,~nri cornlln nf cefiGlnl l l r ~ p l r 111 ~ ~-nr,~l~. t 8 t \ ~ , ~ ~ t l x l ~ l ~ ~ t ~ i n g p ~ r r ~ < t lO ~ < \ I I I K r< IKI~Y !~r ivorkr~! : v 1 ~ 1 1rl~r.\tiirn.>3th,rt btwr thcln \vnt .~ln.v. Such Eutcrn.ll <krlctrrn<o i p l a t r i - d i y r ~ r ~ ccontlqlutw t, t,r ~ ~ v c r l . ~ p -i ni>nmnllv wcrrted alcltlq J i l u c F ~ p ii~ I I I ~ I ,~ I \ ~ I I , I Ii l i t ~ - r p r ~ t , ~ tIIC>IVCVCI, Il~ilitrtl 15. ~ ~ n ~ ,111d III ztnilc wllirh 111,iy13~. hut I < not .lln.,lv\ i i ~ n r ~ r ~ ~ a l . A I I I U I \ nn\,< i i h.lrrl tr> ~t 1>1-\'01icf.I ~ ~ r r , p0111t. ~in ( r rLn tvllci~ tlic i ~ ~ n ~ - n u n,i p n ~ r i c ~ r l . ~ rp n 1s oi or good cu,~mplc i s rlic ccli~rlotlr~ni~ skclcrati 111 \~.lliclitlir plntcq oiicc fnnlicd ,Ire ~lc.rrrl;ancritl! 1 I ) I ~ Y I [trrrion 1 I 1) I I ' d ~~onto!ngirt~ S ~ T ~ prth~cntrd t t h t ' ~\tfitIl .IrC lockcrl into plncr, though thcv in,l\r th~*rcaftcr I I c &T~I\L in~livirlually ? ~ h .~cr-rcrion n ~ a t c n a l corlof I ~ ~ : I E I \ w,hn<c t i ~ r i r t r ~ ~ n 1, nor clcnr : ~ t ~ i l~ h r i h t hnvc I rt 11 rc>ntitcrp,lrt~ Irr rnq . ~ r ) i r t ~ a l s . w ~ t h r ~ i l c In 01h cciihjc zoncr. t717,lrrc ~ppc.1r~lw-c xx~lirrh.11-cy ulud1fic.d horn o Estcn~nlskclctnnq ,111 f~~n11r.d t the m e tinrc.. a Tlir .irthrt>pcld cxnqkcl,:tr,n I F mart n p ~ c ~ l . I 11nrn1.11 wpr fnr tlic rarclil t11;tt t l ~ c vt~ctifv[ O tmx,rr:tllc ,~d.tlitatir)iis. <n111c nttt.ntll.rh ran hc ~riadc C;m\vth hcrc i r dif5c~ilt thc ~ k c l c t o r ~ J ~to L,t= for II I I I V l r d \ 1rlrt.t-ptetin~ thcrr- n1orpholngt.5 111 tcrlrlq of pcriudiunllp ~ n o ~ ~ l t n d , c proccn knrr~vn.ir ecdysis. W11ct1 thc old t \ ~ o ~ k c l c t o n caqr tlie ,lrtIiroIF 1'1 ~ v 111t)n .jild I T I O C ~ = I PC? T V ~ I I C \ 1x1 tli rn III.IV ! r ~ d t I 1 1 , e ~l(,lrt*r i~i~drnt.~nclinq . v n l t r t ~ u proccfrcs. of r ~l~~~ pod tnkr-\ u p Ivntcr or ,nr and ~wcllrrn tllr nest I' [ I i ~ * # tprol~lcm': . ,ire r n tic. t,tc'Ll~-LI. then ,I cohcrcnlt lorgyr w e , nnd rhc ncr~cuticle xvhr~hui~dcsl~cc I I ~rl~nrfoln~icnl c l ~ r . rI i ~r~ ~ c d c d . 1 c p,art~rul.~r , < \ ~~ 01 thc old o n c thcn I~nrilcnq.C;rt-nvth 1s thus raplrl .:%r8 in ~f npprnnrli. the p.~r.idiyn TIIC~JIC>II of .~rhd cpicoctic .und I < ol~l!r pocsihlr rlunll? moultifi~p. Tllc dr~,irlv;untngc oFtlilr ryctertl 1% tlit v t ~ l t ~ c r ~ b ~ l I;*l[!natk 4 lLJOl), 1wc.n ~ n ~ i c h diccur\crl, but i t i l r ; Itryrly O I I ~ E I ~ tlw SI.C)PC C ot' 1111s \t.r~rk ,inJ ir\. ro c t ~ m , ~ .g c r dprcdatinn I . ~ L I ~ I I ~ r ~ o u l t i n g . ~r < ,lnl\ cnmr cx.1111p1c': ~i ~ t \ . I F P ~ I ~ ' . I ~ I L I.trc II , I ~iin~!iitd. ( E\g~b~ltt;tll~, kind o f ckeleton. ~ntrrnal o r ,III\I T~rv)rrl~tod,Irpcrt\ nf p . ~ l ; l r o r t > ~ l\ r > ~ ~ c:111 c*rcinnl i k "lighlv cnn~rrriiii~-rl g c ~ n ~ c t nrules. I hich by t' Il* L!h*iii~c.cri trcvl~fu\r~lrzccl rcrnalllc ,~lotir- p-t~wrh ~ o \ v t l i~ ~ O C C S F C F2nd t h r proprrtlcq n f !l~:itcrinls .IYC , ~ 1 1 ~hn11 Fn!In\viii~ tlic C I ~ I ~ \ I ~ 1 worL nf ~ 1 ' A r c v Thlr wggc~tq t I ~ t . q v c n ct~ougli titiic and an
rc3c.i

11

(,I

cxtren~clv Enrpe numhcr of cvolrit~onary cspennlcnts. the rlisco\,en I I ~ nrgnnicrn5 n f "good" dcsigns - t h u ~ cth.it are viahlr 2nd can be con5mnictcd \\qtl~ .1rall3ble matcrinls - w.1~ incvitablc and in pnnriple predrctnblc. . . tlic recurring d c r i ~ l s e w n h w n ~ c n t t r J c t n ~ ,ordcrfy and ctablc contipuranrt. rlotls uF nl.ltter that nllirt I I C C C F S ~ ~CIncrTc ~n the ~V course of cvolutlnn' fTho~na\ and Rcit; lqq3). It1 p o i t ~ tof h c t ~ I I C p ~ t r t ~ t i a l ,~vnilable11~sbeer) vcrv wril exploitrd by Ilvin y: crc,itltrcc. t h o u y l ~ speciiic constraint< fcldoin ,~llnwthc product~on of idcal orKazlism.;. Natz~rc~vorkqnr .I r ~kcrcr, rathcr i than a< JII c n ~ t 2 r e (lacob. lEJ77: ~ l \ Chapter 3) r scc n - a point to be bonlr In m i i d in ccm~~dcrirlg liv=llI in? 311d tbssll org.lt-tIFrnr.

stratigraphy nnd chronostratipaphy. aU o f wl~icli are w n y of ordering rock stsdta Into m c n n i n ~ r l utlitr (Hedbcrs. 197h; Hnllatld rr n l . , I W X ) .

I< t ~

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rhq-

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rill- tlir~ct
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Lithostmtigraphy ~l~llllil~l rIIIIIIr~llIr[' Lithostratipayhv 15 i-ot~ccrnrdwith rhc c-rec-tlnn 171 IlT<,lrl linlts haccd upon tllc characten of t h c rocks and drffurentiatcd on typci o f sack. e.g. silntcri~r. I~n~ccmn~. 7 I l l \ 1 % I10 o r clay. I t is ascful in local areas and eyscnt~nl Ir8 \ \ 1)c.n ~ h t . grolocjcnl rnapplng, but thcrr is J ~ W ~ ?thC d a t i y ~ ~ ~ . ~ . t ~ r ~ T that cvcn in J small arcn rock utdts ctrt acrns.; rrrnl ~ Y I I I V , I ir.1 p l . i i ~ e ~ . instance, if a rhorclinc has becn advnniFor \ L e t \ prf:L-I( lng in one dircctinn a particular sulte of ~ c d i ~ i ~ e n t ~ I l l .ltlJF 1 probably o f thc carnc g c ~ ~ e rhnd. will be lcft in 11 al rl 11.11 l l ~ i nltc c t i n ~ \vakc. Though t h ~ sbed will appedr ill thc rock record ns a alnglc urlifnnn laycr, ~t will not all 1i:rv: f t t > t 4 \ . tfll? becn deposited . ~ t the o ~ i e tittle; slilce it cut\ acrnr tlorl \ \ ~ 1 t I i z~~ne plnnes i t i q snid to be diachronous. Sucf, i l f * ~ ~fl~ ) l l l l ~~i.irl<cr\ ' dl~~chrnnicrn comrnon In the g c n l o ~ t s , ~ l is rcccird Scdiinrlttary rockc havc hccn built up by laver I I ~ O I I F u r t h e m a r c , ninny s~iitcrof sedin~cntr lnrcrallr are \~i~u71(>~7i~ layer of ~ c d i m c n r \ ~ l ~ i cqnrzlrtir11t.s Plas been much ~~rlpenistcnt: , li diffcrcnt ~ t d ~ r r ~ t ' r r thcier nlav hn\;~? ~ ~ \ . t l ~ q l I.,< ny the sanlc for I n t ~ p pcnodq of ~ F I I FGut a t other tltncr es~creda t the qamc time XVithiti a qnl;lll qyacc - ,I ~ t i l a , ti>r M has ch.ltigcd itz ch.br,irtrr r,lpidlv. The 11ldivid~131 ~snrI%ronc l.l\tl.(l \(Wll Inycr, for lnlta~lue,passlnp ~ntna <l~al! layer; uritliln ~cdirncntan,rocks arc separated L ? c [-.\i,>n some diaance nw,rv. Litliustratigmphv IS t h u ~ only r)l bedding p1a11c.s.T l ~ c hcciJ~ng ~c pl,~ncs time honare i t 1% $111,<11' real valuc within a rclativclv srrlalE rcglon. zons, and the ~ I F I O T \ ' ( i 3 rock rcclliciice rs rcflecred 3 pt*l3~3d\ WI The rlivi5inn~ ercctcd in litho~trat~graphn a in its Inyenng. E ~ r l lavcr 111 tilc rock sequence i1uirt armn!ged 111 3 hicrnrchial cyturn: graup, formai i n ~ ~ l l ~! n havc been Itlid dotvn a n a psc-csl~tit~g l.~vcr, t h ~ t tion, member nnrJ bed. A bed ls a distinct I~ycs so rt T l ~ r * pr thc a l d r ~ trockc arc ;kt thc b o t t c ~ ~ t ~ rxpcr~ed a rock qequcncc. A rt+crlibcr is a p o u r elf hcd of at1 114,ll C V C t , sequence. tlic yonnyc\t ,)I r 1 . 1 ~ top, u n l m the succeq- uillred by ccrtali-t c o r ~ ~ m u n cliaractcr~.A fonnntirri I t y i r ,II ~ U C sion I ~ n s I-recii ~ ~ c t n n ~ c a l l ~ Thlq IS the prin- is a group of nicmbcrs. a p i r l uuited hy ch,~rnctrr invcrr-rcd. (11 I I ~ P ~ . T1 I ciple of superposition,rccogtlirrd a5 long aso sac w ~ t h katurcr in romnlon. It 1s thc p n i m r y u t ~ i 41't thh') bv t l ~ r l - l . ~ ~ ~ iccicntifi N I C O ~ J .Ctcemrn Ilrho<tmngr,aphy dnd i% mo5t u4efill in grolo~c.ll <h ~IF (Ttrno). nlappiilg. Hencc it i x fw-tnatious that arc nnrnl,~llr Str3t1grvphy 1%coriccmed with thc study of ~tratl- rrprescnterl bv rlitTcrcnt cnloun oil geolopcal in;rr tied rocks. their cF~w~ficntioilr l t o ~ ordered ~ I I I ~ ~ F and cross-sect~nn~, .md and a Fnmiatio~~ 11nr111,111~ iq their histnncnl inrr.rprctatinil. It bcars not otdv upnn detincd for ~ t r<r n p p i n ~ t applicatiom. Finnllv, a ~ o u y paqt gc:>logcal C V C I I ~ Sb u t J ~ C Oiip~11 hi~toryo f rallks abovc forr~lntion;it 1s corl~poqcdnf hvc-~ tllr c-r' 11fe 2nd is p c r h a p thc rrlrlrt bn.;~upart of all geolop more for~natioiisand 1% oft~'11 S C ~ s ~ r i ~ p l ~ + i i i ~ U for (Harlni~d and Annqrroi~g,I I; Uc~tton,1W 5 ) . srr;lti*qcraphvo 1 a ~rn~ll-talc rrrap. r Mucll nf ~ t r n t ~ ~ a pq hmr~ccrrlcd~ v l r hclxonoli y 09: ge:t'olopc,llrccord h.1~to I'c CIIVIC~CCF into thr. ttp Biostrotigraphy tulle penodf w h ~ c h qt.~nd~rdizrd, h r nq ~ O F ~ I -in b i ~ s t m t l p p h ~ fo~sil arc AS thc content5 nf the beds nn ble, all ox7crthe bvnrld. 011c pi the pnnlnry d l i l l ? nf used 111 it~trrpretingtlic Ili~toricalfcquencc. It , rtrnt~~rnpllv has bccn to protlt~cc a n accurate based upon the pnnclplc oi the irrc-vcrslb~lir). 1 1 chroiiolng in tvlrich nor crnlv the ordcr nf event< cvol~ition.Thiq rllr;lnq that at nnlr nrlc nionlellt 11 but alro thcir dntcs src known. S r r a t i ~ a p l ~ l c a l claisi- the Earrh'5 histoqt thcrc \ v , I ~ livrng a unique a n i fication is hLiricto ,111 of tlii~. specla1 assernhlagc of ailirn:~l+, cl1aracccri5tic of t l r l l Thesr are thrcc p n c i p a l c.~tugnrjcq o f rtratl- period and of no otl-tcr. As tlmc went un t h c ~ c \vet qaplucal cla~rific.lcinn, lithostratipaghy, bio- replaced by othcrs; cdm \ ~ I C C ~ F F I V f ~ ~ s~ iClS C F I I ~ ~ S : C
F ~ ?

Divisions of invertebrate palaeontology

I ? .I

21

tllc bnlc that thc 4. Since fowl speclrs could n l i ~ a t c t'ollow~iigtheir dcpo~ited.Thus d u n n g own cnvironmeilt through tmlc, thcrc Ir alw.1~5 a tht r',~rly l'~I,~ct)znic rsilnhitc~ and brachiopodq were possibhty of diachronouc f i ~ i ~ ~ The zolir as ac. rhr niurr rrvnnion ~OFSIIF. Ry thc M ~ S O Z Ct11c nlost IIC defined ill o n e xrcn may not therefore be es,~ctlt ~bundnnt prrucnrnhlc ~ n v c r t c h r ~ t c cwere the ti~nc-cq~~jvnlcrltthat In al~crtllerregioll. to ~mrnonitc\:tlicv too bccnn~c cstincr, and ~ n ~ i2nd ls I ~ I \ ~ J I V ~are rhc rommoncFt rrlrcg niCcnozoic time. < III the example n f a paptolltc. thcl-ciorc, for [he TIII~t5 I ~ n wi t .~ppc.lnon a bronc! ~cnlc.kIr>wcvcr. rcaqoin nutllned in ( 3 ) 3 r d (4). t11c total mnqc or \! licn tl~c rimc r,rniy< ilf i n d i \ ~ ~ d af,k ~ i qpccic~ ~l l art. biozone of a species 1s not likcly to be prc5cwc.d In uamliicd ~t ili rvidrnt that wrnc nf t h r ~ c Instcd ti13 anv one nrea, 3 r d ~t 15 therefore hard to draw ~ d c a l nlt a h,lctic>n of ~ c o l n i j c n ltirnc. ch;~r.~ctenring rsochroi~ous boundaries or time line<. #.q. prcriqrly a p:~rticul:~r hricf hicrnrical penod. Ideally, zone fossils s11~>~tld ; p ~ r t i c ~ ~COIIIIIRVPI l;tr In In\ lr~calxcn. nncc tlic xcqueiice of-fbsr~l fau- h a t i o n of cl~aractcnto make t l ~ r l nfully \iiit,~blr , + h:t\ ~ C n Cr r r c i ~ c l v cscahlishcd thrclu~I1 awduous for bio\tntigrnphy. T h e w would be: ollrction ,ind d o c ~ i r n c ntlon horn cspcrccd qrct~ tlti\, th~r Lr~nwn~ ~ r c c r ~ rail o n uccd for cnrtcla~ i hc a wide h o r i z o n t ~ ldi.itrilv~rinn.prefer,~blyin~ri-oli with clrhcr 3 ~ ~ Ccrtain fowi1 qpcrics havc ~ 5 . cnnrrnmtal; b?ctl rn~ind tn bc p.lrtir~~l,~rlv good q r r a t ~ k ~ ~ p h ~ c a l ~ 1 1 ~ verfic~ld r q e so that thry could be uscd to n t-t r 11rl;crr. T l ~ c v ch,~r,~ctcnrc .rhort kccnow of thc define A very prrclse part of the g c o l o ~ c a lcalrldLipcn1 F U T C C E S ~ ( Ikl-to~vn zones. To rake -111 ~~ nq Ifnlll; t,m~plc, ~mrnnnitcq pjrmrulnrlvs{,rlorlzonc foq,Ire c n o u g l ~iiinrph~tuqical h a r ~ ~ tto rcnablc rhcni ~ e s 1% Mcqrvnic crroticctphv. Thr J I ~ W W pcnod IC to hc rdent~fiedniid d i s t i t ~ g ~ l r l ~ c d easllv: 17ttd 50nit. 3.5 Ma, .In0 in thc mnr(.ird R~lricll 5ucStrong, Iurd ~hellqto enn hit. tliem to he cornrr\rnn tlicrc JTC over h 0 ~ n ~ r ~ i o i izonrs hy which rte r r ~ o i ~ preserved: ly I \ ruhsln idcd, 40 thc 7niie$ .1rC drfiilcd hrstoncal indcpcndt.iice nf fwies. as \vould br cxpccmd , cnodr i ~ h i r hhavc ,In :Ivcr.qC durnrlon nf l c s ~ 'than holn a fiee-swimirling anirnnl. 1 n)i111oi~ C S ~ F Y c,ic 1 . 1 The pr.~ctlc,~l proh1i.m~ In b i o ~ r m r i ~ a p hare, y ,411 of t h r w cnlldltior~s scldou~ arc f~~Ihllcd torill nri rvcr. vrn: cninplcx, .uld FornL* part$ o f tlic gcoFils used for 7nnat~nn: perhapr thc t~crlrlc nnlrrloi~ircq )8fm31 ruct+cvCion,Irc t ~ ~ RlOrC c'li7\~1y C I I I C tllnn come cltlse~tto ~ t and ~t 1s not ruq3rlslnF r h ~ t111~ c h ~ L~ , t11~'~. i ~ ~ ~p ir n ih ~ c ~ arc~J F fnllo\vf. Tlir i l t~ p n n c i p l e ~ reallv precise stmrigraphlcnl c n r r c l ~ t i o i ~ nf werc tint worked o u t hilly with ~ S F C F C f o ~ ~ i l ~ , I Minv k~nds f i ~ r r ~c~pccially nf l~. rFtc>w of bottom- notably bv thc Gcrlilan palaeontolopist A. Orjpcl 111 d dliny invcrtcl~rawr, u arc hcicc roiirrnlled. Thev the 1R.ills. lqvcd In particular cnvlrnnmcnrq nnly. c.g. I ~ m e It was Oppel too w h o fimt recoplizrd t l ~ there r ~ n u d\chi Ruor. sccf. <and nr q i l t y (en Ronr. They n r r vnnous way of wing fnqrllr iil s~rati~grapl~y wrrc nhcn hiq11lv &~daptcd p,\rncuEar condifor whlch partially circurnvcnt the difticultit< rnrnt~o~~q of'tcmprr.lturc, wI1111ty o r w h ~ m r c ~ n d , are tioned. nud hcncc diffcrcr~r types of bmmnes. There not tburtd prcwnwi c>titsidc rhls cltvironrnrnt. arc four l r n a i ~ ~ I I I ~of ~ I O Z O ~ Igenerallv uxed ~ F ~ C Thn rncnnc that thcv can o i ~ l y uccJ for come- (Hcdhcrp, 107h). Assemblage zones arcL beds or. ht* Innng p ~ r r i c u l ~ r cnMronmrnts and ~ I I U nrc not proups of beds with a nnmr,~l~~5urnblagcd fovcilr. ~ 1 un~vrrmllv applicahlt T h e y Inay bc based on dl thc lo9qils prc~civcd 5nmc k~ndr nt'forrilq nrc vcry long-miipcd. 'Thcir thcrein or on ordv ccrtain lilndq. T!wy art. u5u311y r.lrl.5 nf rvoliiti~n.iry c h ' ~ n ~wrrc vcn. FIOIV. c vcql nnich cnv~ronmcntallvcnntrollcd and thcrcT1it.c r.in only hc 11tctl 111 ;I L ~ T c I . ~nild g c n c r ~ l h r c o f IIFC only 111 l o r d cnrrela tioil. Range zones ~ (cnw filr Iony--p~,norIccm-t-lnrioii . u ~ d of v c y arc. perl~npq nf nlorc prlcral applicatiutl. A rnnpc nrc littlc IIW for ~ v ~ ~ h l i s l iclmr r~lbdlvi<ic~n<. lng ~oiic usually reprewnts rhc total mugc of a pal-ticuI;cloJ znnc fo~sill; <iicli 27 t l ~ c p.~prol~rcf dcli- Jarly usut-izl relrctcd CIC~TICII~ C I I i~a u ~ ~ O.n e rrta! nrc 111 n rntc and n n l ~ prcrr-n7cd In q ~ i i c tcnvrrnilmentc. tliercforc r e h r to the l'.~iklicm.iplr~rrr~rll;~ zonc, hasrd bc~ng ilc~rro]ccd trmrc turlwlcnt r-cn~d~tto~ls. UP011 t h ~ 111 CPOIIVl11C)ltF ,ItIllllcllllte that dt'fillcs the
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:.tr>~~lphv stage, utl thc t ~ r l l c rhniid, I* :I p-oup A <ucccq~ir:c zones Ir,~vingp r ; l t p r d c t ~ c a lu w , LTPC11v \ i ~ ~ ci t i<11orn1.1lly thc h ; ~ r ~ r e lvorking tirlic Baoks, treatises and s y m p o s i a it nt' T ~ ~ ~ I > I I I I F ~ ~ , I ~ ~ < ~ J thc ~ .n;lrrowest t h . ~ tc;ln ~II ~uall!' I>rzbsccl cjn ,I rcLjotl.il sr,tlr. A . . 1 I i I M C< ; ~ . I M . IIy nr t11r ~ t . ~ g r r h ~ TO&< 1 3 c u ' I C ~ C I ~ It'v~*l t Ilrll. Ncw Yr~rk. (U\rLll bntlc tcs~;t) r i c s c;>n bc c-orrc-lntcrl. A i :In t.sL~rriplrs Ll~rringc>n, F .J.W. t 1 CEt,71 hr~t.rrrhr,~rrStnrrrrrrr ntrd ~. z p x n c dijtiiulticu in t ~ ~ n l i i nprcii5r zo~r;~l Ivriri!lcvr. NC1<tui,L o t ~ i i n r (I.iv.~lu,~blcr j c > l ~ )trcut) y K !II-I.111L,lI.I~ C ~ Y C C L ~ I . C ~ O V I tril0hit~--br~i~lii0- Drntntr. M.J. (cd.) (14'45) Tlrc Few! Rt,rarrt'.?. ( Ii.~~~rn;ltl I I CMI~ Hall, London. (Invdluablc rurnrn.lrv o t str,rt~gr.~ph~c.iI ILI t:l~rt~ar i r dt ~ t l ~ e - ~ y ~ ~ i vC i~lu i-~ a~ct 1 t sraprcl~ , r ~1 ~ h d i ~ r i t l u t i n l ~ kntnvn hnil genrr,~) ul',4ll r.\, r;nprol~tcrare r.-irrl!. prc~crvrd 1 1 t h c siltstuilcc 1 4 limt-.;rn~~c.; o u r c d by rhv hhcB1y fo\siIs, aild tlic Do.lrdtri~n,I< '5.. C:hectl~,mi,A 1-1. and I'owrll. A J. (cdr) tv ( 3 'W7) I Irrt,rrrchvntc~. BlacL~~cll. (.)~L,rd I.lt tcr, I ~ C I I I , ~ f l t h l i ,C O ~ I ~ C 11z3t lillrnll~t I~ ! the st:~p,n,~nt 111 1 1 t 1 < m v-h~cli the p P ~ ~ t c r l itvrrc l w t yrcscrvrd. rc~ Ill runlc RTC;IF, ~ l CC)LII\C, ~ I I C t:1\111~\ dtn ;iltrr113t~' f 111 p , r l ~ ~ r . n r n ~ ~ t r t ) ~ ~ tI iI ~~~I t ~~ ~ I~ A l ~LISFI~S ~ I _{rt3111 ~ ~ I;~OICI~LCOI ,itis~lxt.cl\irn'l-r sincr. t l i r 5ircs of rlepourioii of Socrrry of 1 nrrclotr S~L'CI.I! T't~hl~cntlnrl No. H (11mc3 i.\c nlrl clcl~'5 Huctu,\tcJ ~ v i t ho ~ c i l l a r r i ~ ~ r c ~hu !i*\,1l11r rhl>~lyll prccicc 21111~~-r0-201 C C ) S T C ~ , I ~ ~ O I I S lr EI~cvicr,Am<ttrJanl. (Adv:i~~rcdc s t .I ~ I : I ~ I1I~I. ~ l i r y ~ ~ 1 pni~llllr.at SUIIEC Ic\:c~c~t i 5 f ~ l l i ~ill ipl r ~ c t i c ~ ~ tli:lt diutrillur~nn) ,r,I+?~-tci:l~ia p ~ ocl i znijCs ~ - i o m t l ~ I v s t ~ v i ~ l wilt11 I>rdChjopt>d ~ t tc ~ IErjuc-ut, A.J. (1 9% I ) I'ritrriplrs q{ Rr,rrfhir- , I h~rirrr :!:rm< dciinrtf all ihcllv t&il';. P n l ~ r . o r ~ r ~ b ~ ~ ~ . Prccs. N r w Ynrk. (l'n111.1 if Acntlcll~ic blc Io~91lr v r a rcl,~rivt- l i r t ~ n l h o\ v l ~ i c lr.tn bc p ~ p ~ idlr-lu?wcr.it~i-) ,::J , I ? tlic prl111'1q- i~,wi\ ( % < ~-l1ri3tjt1strat1g~~pl1y, L I ~ - ~ I \ C 1'. I ~ I~I ~ ~~I l.irprr+ f ) , A . T . ( 1 '1'5') k ~ ~ - r ~ . i p t < ~ , , ~ ~ , I . I ~~vt~nliclcrc,is r r t i t * ~ ~ it h:~td to inrrclntc p r c c ~ r ~ l y f!t-ol(y]> ,.it/(/ /~~'i>I~~tiotr. p n l > nk 1 l a ~ t / . 1 2 ~LIOII, ~ Cll~ ~ 3~ :#,I, rrf cq11iv.1lcnt 4 1 ~ r 'ill \ v i ~ l ~ ~ l y srl-r>ratc~l arc:)$, (Nc\.v. V ~ ~ ~ ~ .IIILI I rc;~d;thlc.tcst) U I F i II! fn'ni~il xrcIur.nrrn\, tl~cu ?I, \vull d o u ~ n r l c n t c i\r'itli l i i$ricgs+D.E.C;. ,ant1 Crc~\vtIlrr~ . (I t3tJO) f'L71L~r~~/jr+ll(!tf)~: i 3 ir ,111 c l ~ ~ c .Irc.i, rn,iy collt.iin vcr! fcrv r . l c ~ r ~ c ~ ~ t c .Sptrhcsi~. Ill.~c,liwrll. CJxlrrrLl, 583 py. (-Tlir. ~ L * < I ,111~1 in . ~ l ~ ~ ~ r~f~ i ~r,d ~ ,.1r1d ~, ~ t E1nt.r. d i c y hcltrr~gtn tin~ 1 ;ill, Inmt 11seh11 4 r l ~ ~ r u r ~ t o lC ~ ~ IC~~ I~ R - T 11f:dI) J ~ ~ I p o~c I I ~ ~ I . Z - o J * / ~ ) ~ . IIII. ,i~f:rc:\t hillla1 p r o v i ~ r ~ - cM . ~ I I C ~,>rr l i ~ r dt ~ 1C - ~ r r i - - I I r ~ g p J,C, ( 1 {)74) ilf,in'1r~ ~ ~ ~ x ~ i - e i ~ tbl<C;r,~\v-l < I N c w Yirtk. (I>cllnlitntion of G~ln.11 proviilcc\) !.!? nc5, IIC)LVC\~~,T. tilt- I l n ~ ~ i ~ c lof r siir'l~ ~ i~% H n ~ t o ~ i [>.I-. 2nd Harprbr, 13.A.T. (1 9')4!) Micm. -$:,I\ y I I J I : ~ n~cillatctlto . ~ n d '['F~crcnr:~! fro. L ~ O I ~ I I J I I ~ C T \ 111 p ~ l : ~ c o ~ ~ t o l o.p~ ~ r ~ ~ r i i ~ rqtt/' i ~I//(-> ~ f '. ~n .!~lri.~r)rc ,Iilllcal- d c r l ~ r - r ~ ~ c~ d l n c ~ hullal prtlvof nt P ~ i ! ~ ~ ~ ~ + I I tI X t tJ i~.Ill~~o~ ,t i ~\ -1 ~ ( # pqwrs) 3 ~ ~ ~ I I / 370. ~ fE. IU.A vrrtic,~lSLICL w i ( w , \ I L I $ C I C I ~ ~ T : Isrrilti- ~ 111 ~ T ~I~I Cuwcrr, I<, 1 L)LjJ. Hi$loly '!I' L+, 7r1d call. l H : t ~ k ~ ~ v I l , ,.*!,,1 I I I- ,II L O T T ~ 'ti011. Ar nloir m , ~ r ~ ~ ~ . t p hori~ans ~ . ~;~ l~icaI ( )xfi>nl. ( E l r l i r ~ ~ ~rc.nd.thlc) ~tlj: 1'1t.r;. ucu.~llv ,Ire .;clrnc uhl q i ~ t t c m ~ \rorld\vidc trwils. I)odrl. J.R. and Smntnrl. 1 t J . (13c)ll) l'tiEnrar.rt~l?q),, 5 . 1 r i 1 . 1 ~l n t e r c r ~ ~ r r i n c ~ ~ t : ~ l rr)rrt.l.~tii,n tr 11ot ilnpcrc.iiblc. Gvrrt'pts ,rrrd .-.ljy~licrtrii)~,-, cdrr. W ~ l c y .N c ~ v 2ntI York. Ill ~ . l l r n ~ - r o ~ m . l t itg i ~ rc\.~tivcr ' ~ 1 i l c n ~ c c n l r ~,~ ~ ~ \ i ~ r gv .5( 1 ~ 1 1 .(111trrc..;ti~~,c 2 approacl~) !,: :hc t;:t~r~I, is cuppI~.n1~11"'1 a n d i - n l ~ ~ ~ l c hy i E~TII.III. (1053) Z ~ ( ! ~ V I . , T tht- , ) i ~ 1Sidpvlck .111ri cc S. ,!/' ~ 5 J , ~ ~ll\rllutc datci \\,hicli c.111 hc .rfi;ucd ; ~ rertair~points t J.rcliwl~. Cr~ndon.(All o l r l c ~but still uzcftll text 0 1 1 tl-iannr lifc 7 0 r i e ~.111dfa~l11.11 ~ L I V ~ I I C C F ) ~ ,! '):*rc.vcr approprialc r o c k rxcnr. 'Tlic~c.nrc usunlly F ~ I c I ~ c L ~ ~N. L a t ~ Cir ~ c r n t t1 ( I W I j) PIiyltyt,r~,,ti~ f t t r ~ t q c, ~ .. i'~ '.~V.I\ b(=~i-krtcd hrt\wc.n ft7crilikmur seditncnts, :tiid NNJ Evr~l~rrioritiry rllr Pro[-css.C u l u n ~ b i Unuvcrc~tyI'resc. ;~ .'I:'I~ ibTCLCrCCtlcc I\ 1 1 1 ~; k ~ 1 COIIIIII~III. i\ m o s t tt Nc\v Ynrk. (Exp1,11nsrl;~iliat~i. n~ethnils) #:.~lil;rl\.. thcrcti>rt., r l r . ~ s r,ldinnwtrii cl,itin: lvill Fnlrhndpr.. R. ,1nd J~hlnrlski, I). (ctls) (11)7')1 '1111. W. .!~!+l*ncilc l . u r o n r n l o ~ ~ ~ - , ~ l p~ ccvrcl.ltion; t31t. two .arc I ~ ~ ~ c ~ * r / q j~ p t * ~.Sr;ct~r-tx, ~ Efirt/~ l i ~ i Veil, V l l + TJtt*L't~yc!ryi,dr(r 1il'r:I~ ( - t ~ l t ~ ~ 3 l e l 1 .~ ~ ~ ~ tI1c~ l ~ ~ ~ sLlc.c-r-5$ l>f lFTLi t , p-vGlr (I{ Pftlt~i~trrolaqy. ) n t v d r l ~ . H u t c h ~ r ~ r r >a~~i r d I G l c s , I : l ~ ~ ~ ~ ~ ~ ~ ~ ~,p~tt-t d 11%tl -~ ; i ~ ~ ~ l l mvcs ~ ~ ~ . it] m ~ r ~ g ~ i t ~ ~t .i v Strnudslm-g, i'cnn. (lnv.11u.tblr rctrrcl~cc) II,;!YII to 17vtl1. Tlic 11s~ c~f'.rutnln.~ticc ~ t a - p r i ~ c c w i t i g G n i c I r i ~ ~I<., (1Wl) F~uit.<i ~ t I/I~-FICIL: !1~fi1r1~,1r11~ d ~ rk*trirv;11 p t ~ 1 2ih, L T ! I W ; I I ~ c ~ ~l - ~ l ~l ~ i F~L,~ t t ~ r ~ Ir~~I l~ ~ l ~ III;W ( ~~ g ~ [ ,;~r~dysix, L O I I ~ I I I ,l-l.irlw~. ( I - ~ ~ I ~ ~ ~ I ~~~ ,'lvll ~ r ~ r ~ ~ p rti)r sC~I t~ cI ~of Ct l i ~c u ~ ~ ~ t r a iitn~ ) ~ \ y a l n c o n r r r l ~ ~ ~ itr<unnsticrn i~ Fatl~crt\l in tlic ficlrt) ~l ~ cal i;n~ild,S,I. (IQC)O) Ti;l~lrlr~fiirl( i , r rSrr Ifrrrc~r.jsSli,llr ,irrr! rill, ij ,:.lcr:t \imtipap!,ir-,~l ~ ~ ; ~ ( - I T C L > . }
:1

K.C. Tr~rherr, Rohlqnn 1i.A .III? KJCIIC~. U., 7 'rratrrt. 011 It~t~~ft~I~rat~ t i Ccolog~c~l Socicty r ~ i Atnrlrican ~ r l dthe LJti~vrn?;ltv Kans.v I'rc<<. Lswrcncc K,I~I. (771~t ~ n d ~ rrferrncr work u11 irivcrtrhratc tbwc rd F I I ~ ench phylunl truatcd a l l r c p m t c ~ . o l ~ ~ n l e s ) . MLKKIX , J.W. ( d )(IXi5) Arlhi L I \irr~t~rtc/*rm,~fr i .(1 ~ d\l,~mfi*q\~/( Lonsv~,rn.f l a r l r ~ ~ ~ ~ . IJalnrontolr>pcnl As';rjr1.\rir7n for rlie (Exccllcnt phocopapllic coverage of ~ n ~ puuprj in bun.i\) l'.n~l, C.!<.C. ( I [MO) T/w >\Tnrt~rd Hi<totj# Fvs~r/s 1-1.1lk11~i. ( I cJt){~\ .-It1 4, t1111i- Plt,~tic-r,~zt~ir l?i~~i-tlrrc~p/qj. Wridenf~ld and Nicolwn, Londnn. (S~rnplrbut IntcrClaibrd C T n ~ v c n ~ r v Prcs~, > \ - h r d . ~ rrting) Il.~rlnnd, W H (cd.) (I"7h) '17rr. L l r $ 1 1 Rrt-t~rd - o I ' i v c t c ~ ~ i1. (cd.) (1 % - I W I ~ ) 'rr~~rti j)ti!C[~t!f~llo?rr~ , dto M,rr.~tln, 13.1nr. (The Frerich 'Trcanw', <I~ghtly cllrlc* 13d~Errltrfitliy~' L~s\ilo r d c n nnd ch.1rt.i c h c * w l r l ~ drucr\~wfucru.~rlv~ls t11,tn tllc A n ~ c n c a nTrtaticr or1 Itrr~rrf~brtr?r. but of vcry h ~ g h qualiry) tliro~rglitimc) F1.1rl.111tl. W .U . ~ t ~ d~ I I I F T ~K.1,.I <1,')4[ .-IC;t7tdt)qiin! liaup. I1.M. and 5wnlrv. S.M. (1 97K) Prirrrrj~k.s L>: A ~~ / 1) Pr~lcotrrt~ll?yy, 2nd rdn. Frecinn~i, S.ln Frnnclsrn I'l~trc Ffltt>. a r r i l m d ~ rPJruvcnin; P r w . C~nlhriiidjic. C (Exrcllcnt tcatl~ookcmpha>inng apprudchcc 3 r d rnn353 pp. (l:wenn;ll nnd up-m-dm) icpn. 1ir1 n o t i n o r p h n l n ~ c a l <trati~mpl-r~cal t or dct,illi) F - I ~ ~ ~ l 11,D. ~( ~ l ~ r r lL)7(1)I ~ I ~ ~ > W I ,Ytr,;fi?rc~pfrii,Grti~lr. ~TI~~?I~~I/ ,~rr,f (:li~.<.<ifirt~rfr?r;, -/TI< WiFcy. Ncw York. (~?fFici;~l ~ i i l cto \tr,~tihnplGcal lG4lcy, M . (lW5) i<i~td~~tiort p Rtfiin~rnt~t~lr C l n r l i * ~ ~ ~ .n p l a n . Harlow, ?I 1 I 1 1 ~ of Lu pmccdurc) (C:lcar trcnmtcnt o i dtfkrcnt .\ppronchcs tn cla\\ific~I I c ~ ~ q ~ j tW. +i r d L , ~ d dJ.S. ( I c 5 7 ) 'li-,vrisr ~ I (1h r i t ~ r e l~, + I ffLtl/iicy,irrrl C ) ~ ! ~ T C ~ I P C L ~ ~ F ~ I~ ,J, J I ~2, h4c111nirror thr VL~L I ; ~ = o l o ~ ~Souiim i$< Ain~.rica No. rb7. Gcologcd ,lc~l Sorich- 17f Anicnca. I .~wrrnd-c.Knti. (Sundard test.
Moore.

R.C. tci~ccc.~$ivr cditorz frorrl 1453)

Londnn and P . ~ l a e n n r o l u g x ~ ~ l Assoc~ation. Schsfer, W . (14)77) ' r o h ~ y rritii I~alorclr.colcyy4 .?.lrln)l[ E Irt~r~rrt~rtrncrtts (trant;latcd c d i ) (cd. G.Y. C:m!g). Cll~vrr 3 r d Uoyd, Edinbursli (St,indard work on Kercr~t North Sc;l cnvirormlct~t';with applicatio~nfor p.t!aeoecnloyv)

S~=hoyf'.T.1.M (cd.) ( 1972) Rhlrit.ls it1 Pr~l~ol7iolrrq Frccrrlat~. Coopcr, San Fcqilci~cn.(10 ont$nal pdpen. l w r ~ n n i L I U ~ ~ ; ~ IC :. I I I \ ~~11.1nrd0 1 1 gloh.11 n~ap\) O I ~ t IIIS\IY\. N F ( l t l S t ~ )l,<vci/< ! r r / ~ > n t t ~ r,\rw r ~ ,I< ~ ~ Rtlct~rd1n2 nlanv q g ~ ~ f i c a n z ) .4 (197h) Stmrt~irrarrd nwrl Stmml Corrr.b~il~~n L-rlrr~rqtrt.r. Carr~hnclpeUniversity Scott, K 1 and West, 1t.K ( e d ~ ) C i n j ~ i f i r ~ ~ tofr r ~ ~ ~ ~ o c o t n t ? r ~ tDowdcn. hutch ins or^ u P nitrn. TErrw,Ccrmbnrlqc, J-tpp. and Row, Stroudshcrg, 13cnn. (1 1 nnginal papers) lowcv, K.A. 2nd F r ~ d ~ v , A.E. (IqH?) I'~clhl[,rrtr of I)(t)d~4pcmr~rirt Rt~tt~~r~~rrrrtwrr. Cyqtrni~rrc\ Atrorratinn Speci.11 Skinner. 1E.J. (cd.) ( I O N 1 ) P~fcntrtolngy irnd p~lt.wrrr~~r~~r~rirrn~rt.*: Rmdi~l~gs.fionr .4rw~siron S C I P Irr ~ Kalr fnlann. 1-05 I J Vt~lurt~c Nn. I I , A ~ n d ~ n ~ i i Nt-w York. ( 1 1 Prra\. Altos. Calif. (21 cclllcctcd papers) Sttiith. A.H. (lL)94) Syrtrrrratirs arrd r l r ~ -fossil rt~orti D a r ~ ~ ~ t t c . r ~rt,t>i~~tralrr~r).f t c m r r ,Wl~ckw~ll, ritr~ ~m C3?rl'nrLi (Lucid rxplanatiun o f rladiitlc mcthodulop) Iluckwnrth. Lnr~clnr~ (E:nit~rnr~n~ty n i ? ~ r n i t ~ t ~ n n r ) 5ylve~tcr-Hradley,1j.C. (cd.) ( 1 Y 5 h ) 77ic Sperie'cr Colrit.ppr rrl! rrr P I r b t ~ t ~ t ~ r l ~ /y ~ tq y ,~ r t i c < S ~~ cn ,4~~ocintio11, London. (13avc M ~ - K c t m \ vS . ~ r l d , SCC?~CCC,. R . (cJcI (IC)C)O) Palaeo701c. C work nf r c v c n l papcrq on pdaculltological tas@narriv~ Tcicl~crt,C. (1075) 'Trticltrsr o ~ rIrrr~cn~bro!u Paleattr(~lq~y, I1nrt W (Suppl. I ) , 7i.iart. F ~ r r r l r ar~ti Pro/~ir.t~ini~m. p + ~ r s ~ I I C B U ~ I I I ~I'~~.I'C'ZCIIC \1orI11 n1.11lc) Crolugc-af S o r t r w oKhrlerica, Lawrmcc, Kan. M~ddlclii~rs, A,, I<,l\v+nn, I'.F .111d NCU,111. G. (eds) F. (FC)71) i y i t r ~ ~ r rPr,l~vtr,rrrri .+II.I. ~l t~rrri'lirtll,. Xcc.1 f-.lol!';r 'Thomprot~, d'Arc7: W. (1917) Un Grorrlth nrlri Ftrnll P r m , L I r~ > ~ ~Ii3I . ~ r ~ g i ~Iprrql C ~ LI I I I ~ I d I~ < t n h i ~ - C:nnikrid_ee U n t v e n ~ t y Prec~, Carnbndgr (Ind~vldq ( n i.~ ~ u a l i ~ t ~ clawir work i>n phwlcrl lnwr determtniny c

K , ~ n f h a n i l F . G . .)nil Scrltt. K W (147(1) t l . 1 ~ 1 ~ . ctnllicprr qnr\-.'tl?: t l nbrtdqcd r d r t ~ o nn f this honk nr.isc.rlitcrl I.? n y 1.7, l k > i i ~ ~ r rI ' W B I ,111dI I U I ~~111~1islict-I 111 by (.'3111l~i~!i~- u t rcirrlrrulllir> r c o l o q .~ardp : ~ l , ~ c n e r o l (ill, Sr~rrir~rn, Ii1;d C~~t.~~~!ic~ttorrT ~ ~ ~ : ~ ~ O I ~ I I (~ c Ih~ II t .IW. P Sc~ntt L! / ) ~ I I ~J T Univcruitv 3 'rc.;.;) 'rh0r.t311. (;. ( 1 L):l 1 ILtfi?i t t t l ~ SI>J,UJor!d U ~ i v ~ ~ k i t y ,111tl 1 t . K LVrstl. Ilou.rlrir, I ! U I C ~ > I I ) F ~ I I J I I ~ J<n\<. i.il>ram,I.OIIL~OII. (Si 11 I ~ I L * . IVC-I1 i l l ~ ~ ~ r r ,text: ~~iri~ls Strcwdrbrrg. Perin., pp. 1 - 3 . it~ ~i Locklcy, M. (I"X.3) A rc\-ictt. of bmclilopnd doiilzn.~rcd y ~ l . ~ c o r r r r ~ i n ~ u n t i r n ~ i > t l ~ t - tvpr C)rrlnulcinn. tir P,~fi~ci>t~ttdoqy 1 1 1-47. ( T C ~ I ~ U I I O \ L T # p>het>2h. tix coluililt~nir\) struuturc.) M n t ~ r c . nnd Willtncr. 1'. : 1'17) C:~mvcrgcnr c\.i>lutint~ J. 1' j11 i11\~crtel3r~t~<, ~ * I K (!- I ~~ ~ * I ,~1~ \ h I~I ~ ~ I I ~ #~ 72. -O( Ncwcll. N.13. ( 1 1j71)Tlic cvolurlon uf rccfr. Stit*rrttfit .-ltrir~rr~~~rr 5447.5. { l n f i ~ n n a t i vrllort p3pCr) 226, r P ~ l i ~ i r r .. R . (llJOS) 1310tnrrc. - 3 IWYV k113cio *trLltlA f yt-*~pI~i~- ~ i i t t l f P~:/~~c[>t~i~>ft~yy ~ ~ -{011nid 39, I 4'1-53 Individual p a p e r s and other references P ~ l ~ l i c)\.It.( I 984) TIlc hjclnlerc pmlllctl~.c\~c)lution r, of I'r.llg. G.Y. {1"?-1) T l ~ p a l . i ~ = c ~ ~ c n l r ) ~\? ~ * r t)1' t r Top Hucic .In idcn. J>r~rrl~il ~ ' d r l r t > ~ ~ t39, 1y-IC]-3. of oh~ 511:d~b {Lo\vcr Carbonlicrnr~<)t J. ln~-allna rlcLlr Kilsytlr. I'ctcncn, C.C,I. ( l V 1 H ) T h e rea bottom and it, prcjduc( ) r i l ~ ~ r ~ r ! y j ~ ~ thr t C ;r~liyi[td ,<orla.ry t~fl~vtiio/r110, i?f ~ r ~ , ~ / tuon of fishfnod. Rqlr~nq of t l ~ c T>at~i\h R1oloyrrll 111.7-1 0 . (Clnrr~c pnl;r.-nc.cnltl.~cal .irit(ly) Statton 25, 1-02 ( T h e tint drtinlrivr 5urnrnni-y niljrnFortcy, R.A. a n d Jcifctrcc. \<. (1"'X 1 ) Iihvlog~-:iy . I I ~ J tl~rc o n i r n u r ~ ~ h r ~ ) c w < ~ c ! n ~ r i-s .I roinpr!nt?n<c .qiprr,"~rh. n /'rtdj/(3~if.i r t a!/' Pitkrrill, R.K, aiul U r c ~ ~ c t ~ l 1'. v , c (1'175) Ttrc appl~c.~t~c-rn P!r;./opr.~rrri~8t-i-tltrsrnrr;rijrr (rdc K A . Jovsc-y . ~ n dA.F.. c t f thu ccr~nrniin~h' cnnccpt I n pnlncontolov. ,17+1ilnti1~r Frld~y). A C J L ~ C I ~ U C NCIY Yurk, pp. 112-47 T'FC'IF, .C;rdr~rrrrrirI I , .p. ( T C F I I ~ I I I2nd R P~~~ E C ~ ~ ~ C I I I ) :H P~O ~ (Whcrc r.\.\di<tlcr1% ~ x c 1s not t ~ h ~ t i t l l l ktudwick. M4.S. (1061) TIIVli'cding I I I L * C ~ I ~ I ~ ~f Ithe ~~ II Fii~ii.11.ET. (lt177) C o r . t l ] i ~ ~ U p p r r Jlir.~~qic) ~ n r i l u r I'crrnl~n hrac h~opnd Prmrlr t l ~ u / ~ r ~ i rJol,lcj ) r ~ ~ ~ l r m3. [i ~ n. t y bcntinr . I F S L ~ ~ - I , ~ ~ I ~hI m l E11~1;\nd I I ~ N w r ~ i l ~ ~ ~ d y 43L-7 I . ( l ~ ~ r , ~<~pproacl~) I+ . . dip~~ P ~ ~ ~20. , 337-s5. ( ~~ ) ~~ ~ ~ ~. I C~ ( ~ ~ I-I JCI~I, I ~ ~~4>c ~ S ~ ~ l~ t lD.U. ~(1OH I) ~ ~ ~ L I I , ~ ~ ~ + I l l ,~ ~ Ttie Magncsi.iil L~lnr'itonc (Upycbr brc,llvc c r 7 ~ i l r ~ l ~ l n ~ t i ~ * ~ i Pcnm.~n) rccf cun~plcxcr m i nurth-raqt tnglnnd, rn 1 , 5tr~1ti>t1l~t~j1cr1c>. ~ l ~ i ( ~ i ~ / ~ i i +lirrrr~1>cor1 111 P ~ LT ! ~ ~ ~ ~ ~Rr$ dl : Cr~iiycrt~~!i~ 1 I> 13. lrrdtb (cd. I>.F. Toonlev), h o c ~ r vn i Spti/fcs:> ( L ~ D.E.(-i. 1 3 t i ~ p:and P. I<. C r ~ n v t ! > c r ) , S Ernnotrnc l ' ~ l ~ , c m t u l o ~ ~ *u.~ dM u l ~ r ~ ~ l Spcrial ~ ,t : u~\t Hl.~rk\vcll.Ocftlsd, 574. 3-' Publicadun NII. -30. Tl~lcs. O k l , ~ l ~ c ~ tSEPM. pp. ~l.~. kI.~qi~r.I ) . A . T (I1+*.')) i~r,ii/r~ty,~i/s* I P I t11r L J p p t -/~I l(>l-R. .-lr,~frrt~/i~itr srl~~r..<.r,rr! rC1~-i/t>~lri.i~11r) Cirf1;r1~ l s t r i c l . Thoma<. K.IF.K. 2nd l i r i L \ -E (IC)'43) T l i c q k c l c t n ~ i i f 1/14. ! ~ V .ii,~tFir~~i/. IR,1n2 , 7'1- 1 3 . I f . i l n c o n ~ c l ~ ~ p I i t Sn ! ~ c r v c oc rpncr: a finite S C ~of ( x - p t ~ i ~~ ~ i g n cEl~oErir~or~ cl . 47, \Innnyaphs (tnnnrrmnlr. prtlr-ccl~~rr rr+trcd to it1 text) 34 1 -60. Harpcr, !,).A T. n r l t l I<v.in, 1'. ( I ( W \ )Tow:trtl~:I ~ t a t i s c i c ~ l T l i n m o t ~ .C; (1957) I-Jnttnm conlnlurIittr\. 111 Tri,,rr~w nr >vst<-~n r ~ ~ ; ~ l ; ~ ~ t ~ ~ ~J t nI ~ ItI ~ I !q!!/' t s , (Aott!qii (11 h T I l ~ i s tltr ,\Jtlrll1~, Er17lty). fll7ii P t ~ ~ ~ r , l r i o l < ~ p ~ ~ , (d. , Val. 1 J. \~)irr.t)' I~*rrrla*tr ~ 3 5 - f~ i (Fwcn t ~ rt- ding) I!! 147. . ~ l Hcdgscth), M c r n o ~ n of chc ( lct>lnp-.~lSr)cucrv of tl:jiv. J.-L. (14X-I) A t ~ . ~ l y \ c c-I,~d~.;ti~lur c Trilob~tcs:~ : t l t A n t c r ~ c n No. 67. (;rn!ogcal F n c ~ c b of Plrtlcnrn, pclint LIL. vuC. h / i , l ; , l 17. ( ) ! - ( I . (L;<c:IIICI I~rnlt.ltinn< f o L ~ w r c - n c rK.ln ., pp. -!(,I-53.1. (Standard \vnrk. in> ~ l u . rl,idlmi) ablc, well i l l u ~ t r ~ t ~ ~ T ) 1 I 1 ~ I I , ~ ~ 1 tC'. I-I.,Aucllr\--CI~.~rlc~. I. M.C;., I5nl;wtt. M.G. 1.t Licylcr, A.M., C;ock\. I..I<M. a17d R ~ t ~ h ~ c hK K , ,A!. (I1l7H) A gtlidC t t i \tr:~t~g.~llllii. m r c d u r c . p (I')t,Hj T h r c n i i i p n ~ i t ~ r ~ r ~ d ~ t n l r r u r c r>l' Lowrr dn t;er~lr,~qc.tl Sncicn. nl' ILlmclnll Syrr.i.rl I<r.lxb~t N n . 10. Silurian rnannc coninlunltrcr. LA rlrttt,r 1, 1-77. (I'lir pp. 1 - 12. (lnvnIr~.llllrrntlrlcrt~C S C ' . I ~ ~ I I L ' I I I ) tint iiinjor work a n Lnwcr IJ;rlarn7uic r o n ~ m u n r t l c \ ; \l,:ab, F. (1'177) Ev(b1utit)n . r t d t ~ r t k c n r y .S4lrulrtrb 196. very wrll illuw-.itccl)

[(lt--

Evolution and the fossil record

t>\'cr ~ l l i
I~II~F

11C)t PC' l i i ~ \ t y ~ ~111 c ~ Inttr ! :\t~ci I nr

111 11I?I11\ , L71

111~1

cytolnl

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t3I,li I c - 1 1 ~ ~ it1 \\

i r ~ - t r F>rniip tllcny ~ ~ o l ~ z c t = ~ cyccpr ~ ~ l innlcculnr h i o l o p Arc tranxfornii~~g .1i4diiig ltu and Thc rrcc thi~ ~ynthc+i+, nclv v i r w ~ n tlic Iiaturc. o f tlir nncl o 1 1 wlf-rtylc. ~ gciic Fcun to bc r h,inyi tig our wlit~lc conccptinii ot tlliclltlv d1y Amon<-t .ill the sricllcCF ~ 0 1 7 c ~ m~sirllnrg.inlc ho\\' organisms rv01ve. ~d .111rl St.llllc.!, cx-oluhr>i~it is n n l ~palncontolnp thnt har thc T thc following tcst. tlicrcfurc, I present fintly .I n TI<-cdud licrt ~it~rquc per\pccrivc of ~colo@cnl urilc. It i q thc rock qunplificd account of classic neo-Dar~vinl;ln evoluAltllnuyh rccord nlonc th3r providcc nn Ilisrot-icnl pcrspcctivc tionnry thcory. .iirncd p.~rticul.irly ~t xtudents nf thr. f,irht.r trl for tllc ~tutlvnf cvnlonon.ir\. cvcnt+, ,111d thi< ciinc Enrrh rcicricc who rnny only I T J V ~n Jimitcd backd i n i ~ n ~ i ncould ncvcr l ~ a v ccnmc frrmi 3nv other groutid in b i o l t ~ ~ T . i l ~ ii y l inuludc~ Foinc ~rlfonr~at~crn \vt-rc. nlaiiy source. Ac~-orrlin~Iy. input of pal,~coiitolog-yttn or) thc impact oi ncw i n h r n ~ ~ r i o n thc from mc~lcc~~lnr tll.~t,1111~~1+ of tiiric cvnlurinti t h c o ~ hccn In unrfc-r~cni~rlrng hi$- grnctic?; o n c v o l u t ~ n ntlicory. This section i r not Einq thc i i t > i ~ prL,-c ~ to^ o f iik. in inrc.rpristlrrG p,lttcrrls of ci-oltttrori irlrcrlcicd tn be o o m p r r h r n ~ ~ v[lor does it pretend c, I j,lnvlbl I l l 1 (c.g. adnptix-c r.~di,leinncl ;\nd lirrcc rlf cic.<ccnt, .tncl. tn bc a guidc to all rccvllc dcvcloplt~cntsnnd Jir\ ~ ~ I I ~L ~ 0 I~ 1 1 I ~tnpnrtantly.in a$wwng rate\ rlf cvolurinn. N n n , it cnvcricr;. Fullcr t r r d t t ~ ) c narc readily avnilablt. elsr=t~ ,>i ~iioclrni h d q tn he ~dinirtcclt11.1r rhc For~il record i< incomwhcrc v i r , . , Simp~nu (151431, Maynard Smith (1'375, FrlttlltBVC J plete, ~11dto iiitcrprrt i t can in \ornc. in<tnnccq hc 1C)8?), M ~ y r (lOli3, t Q7h). Dawkii~c (1oSh). thi >llyllWh 'Ilke trying to rc,lrl .I di.ln7 wilth h ~ l thc pagc-5 13oh7haiisky rt al. ( 1 9771, Garnlirl and Vilics (IOX7), t lir>nt rhc 11 m~qcing' (Shrlilon. IOSX). Yct it <till rcr~lnins nri Ronner ( 1 9881, Ertdlcr ~ t l d McLclEarl (18)XX), Inrcrtw~~ irnrnenwly rich qc3urcc of pnrn'ln d.itn, and can Hofinantl ( 1 9H9), Campbcll and Schopf (10?-1), fivt--tingc.rc g v t ' , ifrcqcnlvcd fincly c t i o u ~ l i a f.11rprcrlrrc o f cvo. M a y i i ~ r d Srrlitli and Szathtnary ( l c ~ 0 S ) ,F u m y n a h r a t c ~ ,lor r lunnnnry cvcntq t h ~ ~ i t u . ~ ! ltot~kplarc, I~owcvcr ( 19961, Stricklwrger (1 906). ILidlcy ( 1 c)Oh) and t v or \ v ~ y <TI . long ago. othcm 1iirtt.d In the biblioyr;lpliv. whilc Valentit~c r1T rrlariric T h c ~hzdetltof p , i l . i t ' o n t o l n ~ ~ nrcds to harrc some (1073), Hdllaln (1977), Ctanluy ( 1 979), Copc ,ind rhe hoof5 ( bnckqnunrl in hinlogc.~cvol\jtinn.ln. thco-rr).,otliurI Skclton {lC1X5), Lcv~nton{I'IXX) and S k c l t n ~ ~ (1'3Q31 J ~ L . #dl v ~ r i wsr he x~ill in die po~itlnnr)f "rhc rnan ~ t a n d ~ n garc inorc directly ror~ucn~etl bc with evr>lution ailcl rlrvcrqzty in by the rondridc +~nrl ~zatchingtthc car7 go p x t but the fowiI rcuord. 111 t 1 1 ~ rrc.oild part of. tlliq chuptcr thc G.11.1p.l ~ t ~ t l ~niiy ~de,i ant oil>c>\v their cngincq ~ r ~ o r k ' . UFL' t h c r r is a inore cutendcd auccrunt of ~ v h a t fosril to tlie r~ic(~ii~ Umziwtr's ( 1 973) p ~ p h ~w t ~ i l e . For tlic fos~il rccord can tcll uq about thc n;\tnrc of rvoIutio!~nr)' c EX.73. Icd rrcord cannot wvc n~lirli ~nform,inr>rl the xncch- changc. nu d~vcrrrty in .11liqrn of rvnlutintlnr). rhangc; this has to hc progroup "f bl iriiled by gci~ctlcq,c v r o l o ~ , niolcculnr b l n l n q and , origii,~l pn population dyn=irn~c<, hullding upon thr nriyrlal F P C C ~ C F had contcptloilr of C~IJTICF r)an171n. Mnr~v yt.ar%AFcl the ct1clc'. Crrc)] fint rcal n~ulttdi~crplinnry nrnnlgnm of dat,l u7ar pubh n r m tvl Ilrhcd 3 q Ft,~11tt~totr: tllr ,rLldt.rrt .Cyrt~/tr.siv{Huslcli., I T1 1 the ( I15 1912). From tl~ic higl-rlv r u c c c ~ r t i ~ l attempt a t lvrld- T h c thcnry o f cvnlurmn liiikq togcthcr J nlliltipltc~t (Y-F;. l l l t l ~ it~g tosether iiifnmi.~tiou from v:\nnus sources, thc of hioln@i_calphenomei1.l and i\ undcrp~nneilhy a ~ndic,-~tcd .I 1x0-Tl_ranwniansvn t h e i t t;rkt.r it< ilarne - ~ n td ~ i qig t l ~ c l evidcnuc of thc gcoIo~c.ilrecord. I t rcmainq ,I <tib5equrrit a ~ i ~ e t istarting pnnit. Ifttt rcwnt devcl(>prncntc in thcnr)., n o t n Frovrn f i t . Llr thc irnnirncc t i ~ l t \ r 3 ! r tl
to .I

Darwin, the species and natural selection

27

' v r r n'l~rsl~ rvolutio~ ~ r v li,irlgcr Ii.lvc t.~kcriplncc CCIV qliecles to EIII;LII ,Irc,li only \i.llcrC c~r-I111nd l ;ill:.< liot penllrt thcit ~lircctc~hsrrv.~t~oti. Tllcrc 14, I7ccome addptcd to i t 5 utvn cr~\:irnnrr~cnt. I~ll\vect.r. o ut11t.r ~ F ~ r o~n l i i c lc~icoinp.isccc$0 n r ~ L>nnv~t~ p,~rti~-\rlarlv \vas it~tcrccrcd i l l ~clcctrvc rrllltli ai-rtl ,tee-ord~ ~ v i t h l ~ r v ~ ~ l c oicr>inp~rLitivcbrredinp of ai11tllnl.i;md plant5 ur1dc.r rfc>~nc.cricnnot~. t c ~~cc ~~lrrnmr., hiuclrrmlrrrv , I I ~ L ! p h v ~ i o l o p ,of ~ C I I C ~ ~ CH r rt-al~zcd I [h;rt tlnr prcscllt g c ~ varicry nf r l c r ~ w* t i ~ c f t ,~.td c vrolr~,g-v,ant-l t r f dit- rcl.~tir)ticl~ipc hrtu7ccn c.imlr had bcct~p r o d l ~ r e d111 only ,I fc\\ thnu.;nnd crqiil#nl, prrcc~vctl b\. ~ . ~ Y L ~ ~ I O I IWVh ~ r r , ~ < ve;ln f r a n ~ I the oldy one o r ;it must a fcw .inct'str.il hpr.;. tr~drnccIn .;chrrc ot thrw ficldc I.; clrcun~st.inti:iI. Hc cancltided t h ~ t.I g c a t pstcrlrial firr 'tic\c-rnt ~ I l c brolyht toqcthcr ,lnt{ ~ntcrprctcd hriilrlr up j ~ ~t wltli nlodificat~on' irlttst in all . ~ ~ n i i i i ,3nti ~l~ rlr .r rhporv of tioriii~J,~blr o r ~ < c y u ~ ~ ~ l c c , c and .I\ plant5 I Z , I ~ J C I ~c ~ ~ i I1.e cpreiicd up ld wch ~ r t ~ t l l ~ i ~ l r)rl!vl~,~nckv s,ilJ, 'Notliij~git1 i>ir,lnp ~rnnkr, L > ~ ; ' ~ C I TIl Ii~ . Id C ) ~ I to tllc 1,rli~'ftl~nt ~ 1 1 7 ) i l . l ~ 11aq I l r < S ' I ~ Te~ x i r p t in 111c Iiy11t o t ' c ~ r ~ l i ~ t i r ~ r ~ . ' proccsscf, t l i o t i ~ lprob,iHv nil ,I clinvrr timc ~c:rlc, ~ Ille rc,ccrlt Llcllr ,~:tcrnptc r o Jiccrcdit c v u l ~ ~ t ~ n11nd npcr.~ted11) t h r \r~ildstarc; i l l r9rhc.r \\,orcls 'il.ltn I!\. ,clf-wled *crc.,~t~oei >rrc.r~ri.itr' IIAVC' I>CC-II \ O ~10- u r ~ l e l e c t ~ o n-l'li~rsbruudmg cspcrimcnt\ .irc rhr ~ ' lll~~ntlv CI1sparc11cd h j 1.lu~t(1 OS?), Ki1~I1t.r1I W 1 ) foundnnotl nf cl,u;s~ral yrncnc5, whcrt. rhc mcclin111tl Yt.lnlev ( I c ) X ? ) t h ~ trlo f~rrt11c.rrolnrnc-nt niciri o f hcrcdiq rs rindrrstuod 111 term\ o t - ~ r cct'rcct~. n p r d l d Iicrc. I'lnnmii n r a F also r o ~ ~ u c r n ew ~ t hp,llncontologv d 4ltliou!:F1 Cliarlrc Llanvij~I < y c t i c r . ~ l rcgarrlcd ,r< hut hc f o ~ ~ t idint tllc hcqil rcsor~l r ~ + l~ il n <OIIICIV~F:I~ f o :Ilc K~ltl~crf e~~csllition c r * ~ o th (13,\r~v1~1, 185'3). t l ~ c r r 1 d i ~ ~ p p n ~ i ~ t l111 r i p p n r t i i ~ s I I C cnw i l ~ r v o l l ~ . il w~t ~ c ~vtri'mnnv prr-Dnnvj 111.111 \ L lcnrl\t\ \\.ho po~tul.ircd tic>n:ly changu. He I1.d Iic1pct4 IO find c v l d r r ~ c r :lilt J I I I ~ T I , .~iid l n ~ i t ~ L l l ; ~ n p -olrCr A~F p h.lci ~ l Ion? pcn- nt' gradntint~nl chnllqc h r h ~ . r c n, ~ n i t n ~ l ~pccicc. of I\ of tirr~c n tlint n c n t\pc\ crt '\prcic\' h.ld ;Inwil 'ir~tinitely i ~ ~ i i n c r o utr,~r~'riticli~,~li k ~ '~ 1 r 1 1 1 ~ c t i i i c ~ d s l~i ~ !tnnl prk.-culftlnt: O ~ L ~ ST IIC'\C T ~ I T ~ ~ . IVLIT~CTI ,11111 ancc5ror5 tn d e s i e ~ ~ d a r land nf \ t r ~ t ~ ~ r r a p l i i c . ~ l l ~ t~ Ihnnltl h~nlcclf ~ J c i i t i t j c ~I l I A I I Y dl tT~,rct~t I point; LirrangcdS C I I ~ Ss l i a \ v ~ ~ ~ g t with ~nocIitic.~~tor~'. 'clrqcrn 1v41cl1 cil~~lil rrqnrrlr-cl '1.; cv~dciiccfor thc o n ~ n I n Lct. ht did not I;rnr.l w11,1t lic I1.1rf cspcttcd. 1~ cjt' ~nodrrn EPCCICF jj-or~l prr-cYi\tIIiy ,111~1 morc m.lnz*in as~lirn~cl the inrpcrfcctioiir i ~ i rock that tl~c~ ' n n i ~ n \,iiwrrrnrr. 4 1 tllcqc .Ire acccpterl tr~d,~y, sccr3rd 2nd tllc lirrlitntianr of ktrnwlcdpc . ~ that rlrnc ~c 1 t tllwr~chr l - f i n d nnd ~ t 1 J ~~ tCl I T . I U ~ I I O I T~I v o . F , ~ v c w h c t o n rc.;pon.;ibfc. f i u IV.J\ irldcurl part~~lllv I< . I \ ~ ~ the II~'< ~t the lirnrr of cvt>llttivn.in t h o u ~ h t .,tnd clrlwl!, cnrrcct, hut to this pniiit wc \l1,111 rctrirt> I.ltrr. ~~i:~.mv~nctl I T I < cornlp.lrJrlvc aii,lLrlrllv. Thu \xitIi C3f tllr prc-1Janviui;lii rvnlut~otii~t:, tl~c. ttiort i~l.-finp-rrr\, r prilt.ui,~c l I i n ~ l ~ higlicr vrrtr-- p r c > ~ n i rt~ r ~ i the Frcllc h i i ~ t ~ ~ r , l JC~~~1-l%ipti<tr n ~ . li\t Ilr~tr'r, ti7r cxnrnplc, 11.1q hrrri i i ~ i ~ c l ~ i in d v,~rict\r Lnrl~arck (1 77&1 XllcJ), \<-ho propn~crl I O I I ~ h c h r c i c ,J r l " \ ~ d \ ' ~ . ~ I I C g r i ~ ~ p i th~iirl l q ~f'prhtn.it~*k,i ~ t l fl~ppmc I>an\?ll th.it ,111 luv~nqclrgatllsiil<!1,111 oripriatcrl from (I' nlinltc ninnlm,~lc. tlic lvie1yr of h ~ r d c,111~4 h t ~ : pnrlllhvt niicestors ~17d that i t 1 t h ~ ,ilow plnccce oi :Ilr hoof, of h{>r<ti- ttlcv ,111 1uc)L di\.;imilar. b11t ~ c change< h ~ c bcconlc ad.~ptcd I i v ~ iin ~ h l iilr n p.lrticI:rJ ,111 vnrlnntr nrl ,I r.r)itllrlc-ln tl1c.111~. I_ikt.ivi~c the ~il3r i i v i r o ~ ~ i n r n t ~ . concept oKwrll adaptation c Thc ,Il{r.nr~ qtructurc ancl ti~ilc't~on tllr hunk\ n i nnpnatcrl w ~ t lLnmarck. He .~pprciintrd ~rPi:tt IT In of ~ ill? G , i l , ~ y n w ~ I L - ~ I C ( T_;rtu1*r: n ] . \\ h t c l ~n.~rwin nrdcr to livc, animal\ 11,lvc to hc rtt;riciitlv . ~ d , l y t ~ d ~I ? to L*~~(rluiltcrcd diirir~g thr I ~ C ) Y J ~ ot' thc Brnqk, in all the phy51cal and blnlugcal pr;lm~-tc.m tlrc c ~ i \ ~ i . oc 1111%. lcd Itirrl t o 5,ly 'Seeing tlric q , l ~ i , ~ t i n r I~I ~ rmlrrrcur5 they inhabtt. I r t a qeiihe, of<-clunc,an .ilir.I Irr.cnrn~ srnirtlirr i r l c r cln,iIl. itlrirri,~tclv 111 m related r . 1 I F 'dl adnputinli': i t ha% t o 1lc .~rintninicnlly. n1 ! i < ~ u p h ~ r d \ ,orlc rr~ightrt*.~ll\ ~ n c v nf I tl1.1t (ion1 .11t pl~yrnloycnly aitd trt~pl~icallv ,tttL~ptc-d II\ ctlwrontn kryn.11 p o u c ~ kof Iv~*tlc111 tli~q :~rcli~pcl,~gcr.r melit. and ~t I T ctltlc,ll t o cx,elution t h c t r y to undcrnn c.rcrlrq Ii.ld Llecr~ r.rLc11 ,urJ modificd t;,r difirr.iir <t.i~~rl, q~ichad~pt,itlo11\ how c.11rrc to bc.. , IIIL' Gcng;~pl~ic.~l d~~ri-ihlitiun + c r n .I? irnpnrXV.IF While npprcclatlllg the iinpoi-t.lncc of arlapt.~tinn, r l ~ tnr cvolutlnnnsv rhnuyht 111 clthcr x v , ~ t \ tnn ~ however. I,anlarck 1inkt.J hi5 1n\isllt \virh w m r Tliu\ tl~crulrtencr o f ' r r l ~ r r ' . I I I ~~\ol;ltcd \pccit.~ cnilceprs 110 1o11pr bclirvcd to Ilc sciiablc. I l t c . ~ lunyii~l~) i l ~ i l i v n t I11 pnrn o r tht, ~ ' n r l d wrplv behrvcd tli.it , t d ~ p t a t ~ ahaci cnlnr* .ibout thrn~igli n l;hil~f;ltcd oriqin,il wltlcm~prc.nd an ,~n~.c\trnl typc, :I snmc kuid OF iiltrrnal d n v ~ n q forcc. n 'vltnl ~ p , ~ r k ' ~ri'wquc~~t popul,ltiou i.t>\I.i17sc.,~1111 rrstncnnn nf n ~ v l i ~ c h made nni1nL~l< bccnrnc iilorrnC ~ I T I P I C E . frlt HC
1 1

28

Evolution and the fossil record

clrpnrlq ttruct iri\tv horn lrciv I I C ~ C L \ and t h ~ a thu nnturc of' v ~ r i a t ~ r ~ r l and hCrrdity wal I~scc there '~url~ritc~d c.h,~r,)c.tcr<' tvrrc inl~t.trtcd, 111 hlc unknown, 50 that l i i c views oli tlli5 wcrc ~prcul,lt~l n\ cl;i\sic prvrtlilntc th.ir thc 1 1 ~ . ~ 1 ;trf tltc s~mffc had ~ t l d t i ~ i l f f ~ ch~r~tlir ttllenry to hc fccn to v.lorl: i r~ Ilt.comc lo11gc.r in rc\ponuch t t ~.I ' n c d ' tc reach Tlic pioneering work of thc Au<triai~tncd Icavcs lip un thc trct.. 'The t11c.nn) <IFiiihc.tit.rncc o f Grcgnr Mcndcl i l l 1 Xh5. ,111d of rllc 1.1ter schc~nl nm ac~luircd 1-1l.irarr~r~ I I ) ~ Ir I 111~41ly rrg*~r~ll-d w a d . ~ y ~ T.H. M o r p n which b c p n in 1310. 1,ud the founno .11zJ if gc~~cr;tlly ~irlrc.;t,illlr. ( ~ i l t l i o u ~ hl ~ c r c i u d ~ t i o n gcrlctic cxpcrinlc~lt t for ,rnd t h c u n ~ It \ V a l rhl\ . coiiir evi~lrncc.or .\ kind of' ~ c u ~ t kcdhack fro111 t l ~ *i~ipplicd l ~ c I C ' C C ' s u~ - ~ ~ r s t ~ ~ ~ of~l~crc~llc t t I \ ild ~ iirig the rnvlrmunli>nt npcrntlny to prtducc. dpparcnr ity ccccntial tcr thc a m p l l f i c ~ t ~ o n I S J ~ V I I Irt~eon of 'F 'L~nrarc-kranc h a n ~ ~ - \ ' 1J~nv111, thc t ~ t h c rhand. ). ctn p m v ~ d c d.I lopc.11 ~ t i d rc\r.il~lctlico?' f i ~ cvnlunoiir .IT L ' I I ~ It-iiic fhdt h,+s \tr>nci thc tcct a( time ,1nt1 ~~': Inheritance and the source of variation p r n v d d c ,1 Fr,lrlins: pc)int fhr I.ltcr d c ~ c l o p n l ~ i i t ~ . excc'pt I The f~dl ~ d c f r > c ~ n ~m i ~~~ r q t o ~ q xvt~rk 1350 WJS In the cells of;ill euknrvntes (a11 O ~ ~ : I ~ ~ ~ F I Iin:S ' vlnicee, bactrna dnd C ~ ~ I I O ~ J CtJ1crc~ I , ~ ) ~ C IC ()I! t / 1 t a [ l t Y T f ? l O f ~ ~ l l ' i / l ' < /I)) 1 1 I t ' ~ l F P LI/ / 1 ~ 7 f f l l ' i d .il'/<'ltitlfl, OY ~ nucleus (Fig. 7.1) c ~ ~ i i t ~ ~ elotlgzted t11rc;id-IrkL r~iilg thr ~ P L - Y ~ V I J ~ o/~ ~L i~ ~I o z t r t d ~ 1 t ~; I3) fir1 ~!ri!qqlt~)or !$>. J / ~ l I r nrc T h c 1narn prjlrltq of clic dicr>n. arc rtniq,rl~rtc~nvnrd. hodleq. the chromosomes, wl~icli t1-iaJc of prntc~n ;111rl deosyribou~irlri~acid (DNA). I . A~~imal cpcrin rc1,rcxlucr ariorrx r'lpidlv than ii ~ r r r d r dn m.iint.1111tllcir nurnbcn. Anlrnnl popilt smooth endoplasmic reticulum lntioni. lio\i~c\~cr, thnligli flr~ctu,~tins, tcnd to rcn~.linst,lblr (Hc-rc hc ii.,e< ~ntluc~rccd tl-tc I?): En~lisllmnn M.iEth~i\. tvhn I ~ a d w r i t ~ c noil t11i;i vacuole , / mitachondrfon subgcrt <clrrlcyc.arr cnrlicr.) 7. Tlrcrc. must thurcfi~rc cnirlprtiriun \ ~ i r h i nand bc hchvcct~ ~pccics rhc. 'qtru~glc c x ~ . ; t c ~ ~ c c~ r 111 for k ', food. f;>rIl~irrgILL^ :~trd(~vitl~lil c l t l b ~ r < nl oftlie qamc qpccicq) tbr IrlJrcs, it'thc ch:lrdctfr\ tli.jt indiv ~ d u : ~ h ,\rc c t ) 1 t r . ~ ~ ~ ~ ~ tn~i tlic' timt gcnbcdr c . ) t~cI t cratlon. 3. Wir111n s p c c ~ c ~ .inirrl:~I\v~n', 2nd th~u nll varl:ltlon Ir ~rllicri rcrl. 4. It1 rlte ~tnlgglc r lifc t l ~ c > w h ~ndividt~;~ls tittcd host to currivc 111 , particular ciiviroii~nciit ntc thc i (>TIC$ tu IIVL.nnri tlr tc-pl.cbdticc. T h c othcrk Arc wc.cdecl out in tlic intcliw rr>talpctltir>ii. T11c hvour,~bler h , \ r , ~ r t c n < tth,lt ~makr >llch sliwivfil ~c porsible ,Ire it~hrritcdby tiirurr yvilcrG~tionr, .111d thu accuintrlatic,rl n ditkrcrlt th-otir.~bIcc t ~ ~ r a u C trm Imds to tlic ~ c Ir,ltiorl o f ~ p c c ~ c sv c l l p ~ , ddaptcd rtl p ~ r t i c r r l ~ r chnvircl?lrrlcnt*.Tl~ic tvh.lt r).lncitr 19 c~llt-d'n.lt~lr.11 \clccrlon0.

t h ~ iirw t

~ I ~ L13,1nvin'\ I L ~ I carli inqk~ncc, u - y c d thdt n ~ n v i n , had 11ut rually c l ~ c r \ v k ~ u ~ vL ~ ~ ~ C ~ ~~ r ~ ~ C wcr:' ~ ~~ < ~ I MU Z ITJ >

All thi\ ccchrn< luq,ric.alcllo~rgh. ~

C T I ~ I C F ,M l v ~ r for t

actually accumulated. only huw d ~ o s ca i i i r t ~ ~ Icw l< fittvd to their ~ ~ ~ v l r o r ~ ~ i i t o~,unrivc. I11 t l ~ i ~ Cnlcd c ~ t they \~c.tq: 17rlt ~ii1~1>und. t l l ~ ,n105r S C * ~ ~ I I I I~C ' c ~ k fi>r ~ivw tlic r h c t ~ r y , Ir pr~,<c.ntcri T ~ n n ~ i tlvnq thnr Figure 2.I of b\ l,

rough endoplasrnic reticulum

Eukoryote cell structure, with orgonellel.

Dutwin, the species and natural selection

29

Tltc DNA ~ric\lct-~tlc ~ r r .I~ Ir>ily,n~,r<tcd. p i r ~ l fi ~\ ~ t11.1t g c i m d~emcslvrsarc inatlc ut' hvo lut~ds of 1:1lcrnf \r.hacl~tllc. ' i ~ p r i ~ l l t comtrt of ,iltet-r~atc cnimponciitq arrsnged in series. T h e w ~ r exons. \' c Nltk4 of pl~orpllor~c .~c-id.111il thc pcnrovc sug,ir \vl~ichcndr For ~ ~ O I C I I I Fnild introns. \vhic.h rlo , ~tw,nbow, whcrcar tflc 'nri~gs'arc rn~tc-hrlip ~ 1 t s nor. Whrn thc unclc.,lr D N A I F rrarl~cn 1 bcd t o r c l , i t ~ l . ~ <III.III ~ i i i ~ t \1, 1 ~ - ~ l c ' l ~P.ICCS.t ' .l~ ~ ~ ~ ~ r i d TII~'>C nuclear R N A it r-ctnln? thc orpniz:ttion r d t l l c on?L , ~ c ~111, PNA, .lrc . ~ d ~ i ~ ~ i i c . \ gu.lninc. ~liviii~rrc~ c d 1i1,il 13NA. but \vllrii nuclcnr ILNA is rcprcrc~..;<cd nr tn P, tn\ilic. TIIUV,Irk- .~ttacI ~ L % to t l ~ cpcil tow sug.lr I nlewrngtr R N A tlic untroiir ,ire Invr 2nd rhc c ~ o t l c u ~ l l t t. ~ t ~ d prolcL.t iil\tarc14, l i ~ ~ k i r ~ g rocctlicr In up .Ire spIiccd r n ~ e r l i ~ ~ r . docs nor ~ l \ r ~ a \ . < -Tli~s tnkc placc 1 7 ~ r i AJcnu~c c. 11 onlv Iurr w ~ t h 1 tl~vininc, .lnd 111 thc nngn,~E ni-dcl-, hon~evcr.dr)C( ~ 1 1 ~ 1CXOII ~ l l ~ l f 1 .ro$i~ic w1t11g t ~ ~ i i ~ ~ r ~ ~ . f ing n i ~ y tlit' haw of mmd c ~ n l ~ ~ noo p i o t c ~ r l r be iir Iri tht. chrorno~nrric- thc I3NA rrr,intI< are thcinsrl\-cr IT] iiclrJ c n r r ~ ~ u n n t i n ~I ~rcn ~ ~ ~ d te ld~ t ' . l , i:lngcd in discrcrc LIrilrq, rhc. genes. zvhich arr rlicsc arc tiiilctronal. t m v g e n e \\'FE?illr 111.1~ ,uimc 11111:' togrtI~cr .1l(r11;: r11r I C I I ~ T ~ O C the chrnmn~liroii~li cnnip.ir:itivel\' fexv ~ ~ i s rllutflallg -:~'\~cilt~. li )In(, art1 i,f n~hichtli:.sc. nia! Ilt- cc\,urnl h~inrlrcd 2nd In ~110rtp c r l o d ~nf tinie. TIIISF :I neudy ttncPerI 1.r cllromoro~n~.h c g e n o m e t $ . I rcnll uccd to stor2d phcnnn~riion.hi] t n l q li.ivc rlrlportLlntcuiiwT ~.tcr~t~c R ~ I I O I ~ tlic Fcnctic c o i i ~ p l t . ~ . Thew gcucc q~rctrc-csfor cvcrlut~ont1icor-y.) r the pn1nar-y Ilnlti of I~c~rr~din., : r-.irrviny rhr Whcei mcwcnprr RNA nrn\,cr n t the ~ u r f i s e01-n yncric rode, ~ ' h ~ .1 4t inyc~Iv~'C1 ~ ~ C ~ ~ I I C I I I 11l)oqorne it does not ~ ; ~ I - I Iprotchin dirertl~.. but l 111 pT17~ I I I I + dnri 111 illrcr.tily the cic~c-lnprn~.~~t fitnca d ~lirtlughvet. ntlotller ultemirdiate ~-l~ulcculu, transfcr I l l n l n g ~ ) rht. \\ htllC ~lrqnnlsrn. All tlic nccc<s.iry t KNA. ,lnd when tl~rs romplcx proccw 1s coir~plrtc, ~t;tr~u~~trnn li,r tl~cst.cn(l+ 1.; rnrntd 117 rht- I J N A , the teclilt I \ ,I platcln 5eqrierlcr. rcldcd h r by thc 11~1 what I\ Inipnrt,lnt I< rhr w q ~ z r n r cnf tF~cin11r lluclrnr 1TNA. t h r traii~ferIINA n ~ c ~ n w , h lrcttirnlc lurl~,ntidch.~\c\ ,ilo~ay its I r n ~ ~ h . Though r t ~ c ~ c tn thc cvroplncnl irlg I~'.~II 1 Lrnd of ~lyl~.ihct ron<lchty only n i fmtr I'he ui~derctanrl~ng protrill ~ ~ ~ r l t l l c c i shccn 3 uf ha< ~\tt(~n, ~ . r>t~rnl-r.rr t t ^ pnscthlr- cc~mhin.ltinn~ th 111 rn,!jor n i u t ~ ~ for ~ ~ ~ n l r r ub a l l ybut thc u ~ n l c pl ~ li r c ~ . o l I ~ : ~ h cnn cu~ct, thrv coding !I;x <ye(-itjcprnt~iiis. ic cular p n t I ~ w ~ h ~ lead frcm ge11t.c c .~ct~ial tv t o corp.iiw N.mrl%Lmvur. Tlilq rc=rlilcnrc ai ilttcleonrle h,ise< ITIA cl-mrartcr:: arc v c cnlr~ples,dtld a t prrwnt ~ ~ %':pvrmrnt.\ rhc vhcltlc.ncc 111 ~ v h r c hany nf r l ~ e 21) lnrgclv- ~ l i ~ k r l n wH a.w rhc protein.; snd o111crcon)~l :rlIlntb nr.~Afound ~n l i v ~ n gnrpni?rn< .Ire ~ t n i n f i p ( > ~ ~ i i d \ nrhlch arc produccd arc o r ~ a u l z r d and llullt ,I, , m,lkr protrinr. I t h . ~ r hrcn F I I ~ ~ V I - It h t up into f i i ~ c t ~ 1 0~ 9 1d and whoIc 13odlcs scn1alnc o ~~ 1% ($1 I < nf t l ~ r r c x c c acting t n ~ c t t l c r b r-nde fim n r ~ e t h r Irl,un tarkq for ~ r ~ r h c c t ihlolo,~ t ~ r nf l~r f thp x ~ r ~nnrl ,icrd'i, : ~ n d rhcrc : ~ r c tnnrrl. thdi~ tiinlre. Some p r o q e u has alrcndv hccn 111adc; 1 1 ic I ~ L Q J ~ C ~ ~ F I I cc~rnl~lr~ntinn\thi3 'bacr-mplct' I ~ o ~Ic ill [low krrowu t h a t fame penes Are r ~ t ~ ~ l i ' t l l ~d, t ~'d ll , \ . . t m ~ r tn m ~ k c rhc htnci,l-uc ,ln-unu ,ICIA\, a114 to i all concrnlcd orllv with the yntlwsir of rnntrnalr, '1.4rhrtri try iir tlic nzht orrlr-r m iiinkr- n specific o t h c ~ r Are 'regllarnr' Srilrs. w-h~rh tontrul dncl yrs'tc ti. 0rgs1112c the c t ~ u ~ p n ~ ~ i ~c iddirrct thrir iiuildiny. .i n c Irc r\ntt1c~i7cd, illlt in rhr iinrlrtls, but 5kirh gellef releaee cheiiiic;r~prnducts \\.hlch st,lrt a t ~ t n c ~rhc ~~'11111.11.r-vr~pl~lsin. this wlrolc ho5r of c o n ~ p l c sre.lcrioilf. In ~ O I I Iklildt in ,md ~ b , O I > 1 1 1 1 , ,111 tli~, i infim11.1non h , ~ rtn bt. tran~fcl-rcd of dcveloprnent the priies ,Ire switcIlrrE on nnd otf oi tllc ~ ~ u c l ct~ rthew slrcc c ~protclu .;ynthc<i~. r part~cu o r f u l,lr wqurslce, relr~sing proclucrs wluuh F L ~ *Ill\ tr+ r,l\:t pl,~r-c.thc ~ n ~ c l c . DNA fint pro- r r x t together 111 rvntl~cslzl~lg , ir conlplcr nlc)lcculc~. .I , ,I wqlc-\m,rnd~=d ropy o f itrclt. n ~ ~ ~ l e .N r , Scn~ctuml3cliz5 c.111bv nctivatcd a i d cle~ctivated K iA artll 11r3c.il rrpl,lc~~ay rhynlinc ntirl i-iho~e wgar xvl~cnnccdcd; cvsdciltlv thc ~ n i r ctlti~tilu~ the ~~l For ' i l t ~ I+ I I ~ drvlxvnhr>w. F n l l i ~ w i n~ h l cp-occ5s of sw~tcliingolr of stnicniral gcricy rs FIVCII I V ~ I C M .I ~ rr,murription', ,I fu~rthrr ~ r ~ ~ l ~ c u l c (mcwcngcr wnsnr gciie receive? a n JpFropnatc 7 tllnnl~r c. 14 t~jnnerl.ivhir 1 mcwcr oiit of' rhc t ~ ~ ~ c l e i i c , Slriiplr orcnrusrlrs such A S bactcr~a and f ~ m p have I tlir~utgh prrc.5 ln thc nuclear u~crnlir~ir~e. of;l$acent grn.ut-sk r ~ o ~as operons, cndrrrg h r cets v~l Ill ; ~t%clt' thc nlwconir. (TI rn 116 I C 1101 a a p a r t ~ c u l ~ tl~ctaholic path\v,ly. a'l~chc can b ~ . r a1 nl 1'1 ''CiF- b l 1 3 ~ . c ~ ~ ~.IT tli~s I for r . st,i?r the Tencs r~vitclicdr r f i a n d or1 tngetlrcr. In Iugllci- ~ ~ ~ J ~ F T I I S tls*~~~\r.l~cr arc innil~frc~l. IlaX 13ern chorr~ii It rrccn tly Ilnwover, g e l w ~vhicllror~tsold~tlircritp.lrts of .I
111:
Ijllr

'

3 0

Evolution and the fossil record

%conrdin,itr.rl Ilrogr:imnic, m.iz hc <cnttcrcd n r o ~ ~ n d t ~ l ~ ~ nbcing c n ~ i v c t ~ n o ~ ~ n l l y n c 211. 111 hcr dch~rcrf 5 A"m n l l LTmupr. but o n drtrrrcnt c h r n ~ n n v n n ~ r ~ . W l ~ c i i thr bndv (somatic) ccllx of a11 atrrn, . .-.^,..., Gc~lrrir-nllv ~dcx~ti~-al org.ttn<rw rcnrcd lri diFcrcnt dividv ,I< the o r z , i i ~ i s ~growr, t l ~ cchrotiin~nn n . . \ c r ~ v l r n n i r ~ c n t ~ 110: dcvc*ltsp r o c s ~ r t l y \vtlf tllc r;.iralc divide hy Icn~$trid~rlnl fr<~inll, i d tach 'daughtt a 6 4 , . ; . j , >. <1 , form, n.i I \ I Z ~ I ~ I T C < hv ~ hi^ iiififc-t~~iit FC .q~pp,lr.liict~ cell tiil~crit\n a ~ of cvact copy of tllc ~ l i r n m n c o t u c ~! - '--J-7--, :b. nr . vcgctdhlcr polvil 111 r ~ i hnnd pimr ~ n i l r ,rcqycc: -4.. geirllc5 111 the parcnt rell. l'111.i pmccqc, knorvn 1 :L.?' rivclv. T h c ~ n h c n r c d ~ciicttttrrntcnnl 111 a i ~ v r p n - ~ n i t o s i s o (Ftg. 2.2). is ct?;ctivcly thc c:uiic for .)I1 t ; ;- ",;d-~ . ,- 7: I~TTI, kntnvn nq tl~c* genotype, t s smc-~cct Ilpnn b, thc vnnintic cclls in niiy nnc hodv. ! *,. 3.. 2 1 ; 5: rtnJlronnlrrlt ( p r r l l v h l ~ tl~rfrugh rciirrrr gcricq) t r ~ W h c i ~ ccI1 i< nnt d~vtding, c h r n r n c ~ r n n ~ c ~ n thc a -P--. ---....crcntc n dcvc.lnp~-d ~t~ri~virJt~.tI. il~divldu.~l, Thlc thc I,lrgcly inv~viblc:t h i ~ knnivn 25 intri-yh.lsc. A< 1-0 ir . prodt~ctoi botli Ilcrcclit~,111il c n u i r o n ~ i w ~ rr t tlir dlvlrinn ~ C ~ I I ' hinlo$ical F h i n l n ~ ~ . F , wvralc thc rhr phenotype nlnsnii1es. c ~ c h alrr~,~dy dirridr-d Into tr+m c+hrnn~,trl Interphase S o n i c t i ~ n c I~ I ~ I C qC17CS t-017rr01 <inyIe ~ . h . i ~ 1 i t ~ n . tngcthrr hy a small rphtncal c-rntmnicrc. \vF~is F I hcld Morc nTtrii rh.ic\ctc.rs .lrc polygenic: r h . ~\ , ni.in); Lint.< 110r kt3iil The fint rtngc of rcll d~vtctnii I F;CTICS. C.IL.II ~i \rnnll ct'fcct. ~.r,i~tril>~itt' h r bro- prrrph,~w.t.rlhcrc two ccnrnolcc moyc to nppn<l tc) t chctnic~lp.itl~w,~v.; rccl~ltti1 thc fi>n~-r~trt>t~ pnlcq nnd a ' ~ p i ~ l ~ f l c ' r r n t ~ i n t J i ~ c . ~.ippc> thnt of a (>T d~ thcm. A t niet.~ph.lwthc ntirlc.i~tiicn~hr /--*\ 19 pnrtic~ilal- c l ~ . i r ~ c t ~ A. r . so~i~c gericas .ITC h ~ h % , ~ { : n pleiotropir; thar ~ u , tl~r'y i k h t .;vl-cr,ll cli.lnctcrc vnnlsJlcs . ~ n dthc chror;lmolnc< 11nc up .1Jn11g t l y \ ; . . uncc tht. v n l c gciir- prod~rrtqmav uwJ 111 d~ffcr- m c d ~ n ~ ~ ~ 3 ~ 1 1 planc, cpllthng 111 h' FO that c.v 10 c n t biocl~cmic~il p.1rl1w.a~~. p l ~ c r ~ c ~ r n c\n~ li ~ i ; ~ t chrcrt~r~ttd A o ch bccortlcc 3 Jnughtc.r cl~rcrlrlo<on~c with .. ( * finr ~ y l l tf~c1115 1-+117?3111y th,it IIILIL-11 of thc ow11 cctitrrrmcrc. T h c n . d u n n s nn.~yhn\r. tllcy 171 is ; ?A-+I grnorlic c.nnsirrc nf rc-pi-atd 13N h FrTncnt5 (multi- apnrt sc that cach b t ~ n d l c rlnughtcr chrnrno\nrr, nf t<*: L-L-;~ -.* g c i ~ ct:.rrn~lic.s) tvith , r n ~ t h i i ~frrlrll twll to rcvcr.11 muvc\ t o t l ~ c y cq~poc.~tc pcllc. Fin,lllv, a t tclnp11,~rr.. t h o u c ~ n dc n p r r r~f c.lrnc. :ycrlc. Thnuyh it inav I ~ C Wn l i c 3 ~ ~ r thc ir~crnbsanv 7pp~;1n~ O ~ I T I C F cnuh qct , J p F C N that i i ~ u r l ~ I-1.rrrhnnromal m,~tcrial1s rcdi1r1- chro1no~orncs. thc rp~ndlc vani?;hc< ,mcI tllc rl , . dnnr, thcrc co~rlrl n.~-llIle t ~ ~ ~ p ~ t'~011itini1~n. dlvtdc< llitn h t r n den tical rjnughtcr ccll~, r~.~iit ~rnpl~cattr~iir .ir i c di<c~i<ccci hcrc, Pdtc'r. O n tllc other hand, ahc ftmnatinn ot' cgpq . %perm(gametes) In thc tc~i-CF and ovaricr rc'ipr , . . trvrly itivnlvc~.I VCT difi2,rcnt proccw: rneios , 7 +-. .\ (Fip 2.3). .+":< ' . ' ' i. Where d o e s variation come from? I' . . '* ' ' I licrc .ire hvo ccll divlcinnq T h c firqt n f t h c ~ c rr ' ' wit11111 11~1clcu\ ~ " ~(ell y thc. hndy of'rnc~rc rc.ductroti diviqior~ ill a r h ~ c h cach C1311ghtc~ 1 '. tllc ~ t i 171 ~ livlnfi orq~iii~tnc 1%a cpc~itjc ~ ~ r a t lr~f'chromo- inJir~irs thcrc ~ bcr onlk one chrninair>r~ir &in1 311 1111111010y04 Cornc? x ~ , i t ltlicir gcrlc. Tlic c ~ i i r ~ ~ i ~ i o IcI o ~I n~cC T ~ UI I I$ p r r . Herr n p t n tliert' ; l r t f011r ph;lr~f but qoriic k- . '. / & alw~yc COllGtdrlt ti rr tlic I ; ~ C L - I C <I n l i ~ s ~ i i . i t l ~c ~ cI ~ L ' thccv arc ' i ~ ~ n i f i c n n tdlft2rrnt frr>iii tlicir rnltnl . ih , lv 33 pair< clf slirr~rirt~r~ntic<, ill the' f;uit flu' countcrpnrt%.1 )unng prcydlnsc nf tlle fint inciol 1% hilt telophase Drtlst>pElillrrhcrc arc. t i l t ~ s , ~ ~Tlic tmc~ubcrsof c . , ~ c - l ~ dlvikion. w l ~ t n t h C t-hrnnlo~c>~iics p n. pair tip, thr pair arc. all homolognus. 1.r.. <irrr~l,+r ,lppcar,iilc:. drvidc. longitt~d~iinlly, 11) fnrnling grc'rupq of hzir clitt dt~ri 101gtl1. c*;~i+pta l l m ) ~ ~ c ' tllc ~1 c l i r c ~ r ~ l u r t v ~ ~ ~ ~ , (tetrads) h i t w t l r o l i l y nnc Crnhnincrc 6 f p.~lr. mntids upr111 n.hrcIi tlic yr.nc< tc,~q,lrlat~ng +cstr,tl cl1nractc.n cnch p ~ i of ~ h r ~ l i n ~ tThesc four ~11rnii13~iEj'r r id~. ,1 .~rc locntrd. ITklc ccu ~ ) i . l titldividu,ll dc.pcnds 1ipo11 ntict~ ~ tangled a t ccrtdrn polntq kuo1-n nc chiasmnt thic pair nf ccx c h r ~ ~ r ~ l u < o r r t~ tt-- \ i.i ~ .crbs thc chro- and ~5 chcv ptill np:lrt thcv brcak aird rc-t'orrn. c.br I ] r Inc>somc< IC~C*IItic31 (XX) 111 I ~ C ' o t l ~ c ccs they .lrc c ~ ~ l ~ ~ i i paiwl g Jrc r g i i nitfir q.inw lcngth w ~ t ht v honln i d~h~it~li!.ir /?CY).~ I C C S ~ I T O I T ~ U ~ ~ +L?-~dI m <i r t h ~ r n p i i q chronioeomc. Tlir result of cuch 'ctossin T ~ C C gc11cs tllnn t l l o ~ ~~ . L - I ~ ~ c J rcql011\117l~' \i.sual \ L I~~ fn1over' ir that t l ~ c uiipn!rcd chlnincrcrr~nr\ pnr\?,2 The stages of clinr~ctcr< ,111i.l t h ~ i i-r-rt.t~t~ pliy\ical <h.~r,~crcrr arc dou,ii to thc dLiug1:hrcrccll nrc rmt tdciiticnl \ \ t l ccs-l1nkc.d. 111 maiii~rialr,thc tciii~lci q XK and thc thncc n f rhc parcut ccll. n f they nrc cotnpovcd of lvr FNTlrrrs hnm ~rl,ilc XY. hut in llirds i t i < tll~.cithrr \tr.ly rnundj. nf e.1~11~ l f 1101n01ngn11q p d l r ~ dc l r r n i ~ i o ~ o11;"~ ~ r ~ ~ l fflI1111 tht i Il~ 't]~ \Vl~cnr - h r r ~ r n n r c l ~arc s i ~ ~ p;~~rcrE chic thc c r p like T h e scrond nlelntir div~sinn1s likc a soinatlc rlrO*'oFo'nCc-lrU niz.atiorl 1% ~ . ~ l r11) hc. diploid, tllC r-lirnmac{~inc 1llit0F1~. t11;lt~ I I C l so c~I(+ prnduct nf tlir hvo c~~vr<ia:~~lY I~~~~~ "f th"

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h'. ;;: E +

'

Darwin, the species and natural selection

.-..'I

"'..

,
.
interphase

,.:
"'

centromere

prophase

metaphase

telophase

!c~8:?.2 The rtnges of mitotic cell drvision in a wkorptic cell. For explonaAon see text (Modified from Kershow, 1983.) p m c t c r froni a rillglc p.lrc.nt cell. Each iriarcri,rl war exchxlged durir~g crossing over. c ha)?half the nu~nbcr oKp.rrcnt chromo~omes: T h c ~ c panlctes containing unpaired chromo'1. ~'lronrotornc\ arc uupnircci, ,1nd none srr iclentl- \ornrs; arc caid to he haploid, and the chromosnrnc , ' : l l . l n ~ ol the p ~ r c ~C t~ I ~ O ~ ~ ~ SIIICC T ~ C S l ( ~ I genctic nuntbcr 1s H.
'Ir

I..

32

Evolution and the fossil record

~nterphase
<--

-L. ..

prophase
,-,

--

--\,

7 -. ..
% .

,--

./I

!; s

q,
., ., /'

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C Z s
'\

.,.. -.I-<

'

.
rnetaphase

, "
chiasmata

anaphase

second snaphase

telophase

gametes

cell. Figure 1.3 The stogas of rnerotic cell division in a eukoryot~c For explanation see text. (Modifiedfrom Kershow, 1983 1

hn~vever.arc not ~lccccwrily d c ~ l t ~ c fur ,tlwv 1n.l i ~b V[rl~cri ~ lire* icrtil~zcd cpcn n tllc I~c)riiolug~~:i,us c p 17v CX~FC a IILIITI~CI ill of ' C X ~ T ~ S S I O I ~ Sknucvu ;rs allelc~ '. rhron~usomc.\Kri)~iicllifi-rc~~ t p;lrcrl tf coin? tnger11t.r ar~ctpar up. In t h i s proccs\ of recombination ,I nv\\ eacli nf tvIilc11 Ir rl~ghrlv ditfcrcnt cherzuc~llvJIwill lead to thc dcvcloprllcnt nt' diff>rei~ces111 tl gc~lorypc rcwltz horn JL~T.IJV t-stqt111y C ' T I C \cieriving ~ . d l ~ ~ I.dI1L' fkorn d~tlL,rcrrt<ourcchc.Thc ctf~hct of 110th c r o c ~ i r ~ gc h n r ~ c t r r sit CC)IIXTOIS. SUCI~ l f f U ~ t tlldv ~ ovcr a n d rcuoml?m:rtioil I \ t l ~ \.ari.lticrrl \\ itliir~thc fro111thc mlnor to thc very \ u b t t ; ~ ~ l t l d . ~ t S o ~ ~ l c o m c s . ill thr Fenef controlliiig Iiut11: .is producrs o 'rcprr~rt~~ct~nr~ It 1~ T T F . I I ~ ~ I Y f I F higli. i1pt711 r y r rulour, t l ~ r r cma!' by only tn.a ,kllclcs. d c ~ l c thls v a r i ~ t i o t ~ t 113 tur.11 C ~ I C C ~ ~ C K I I~~ C F . h O~C~ . n.4tt.d R 2nd r in cnnventlol~~ll notation, R coritrtll h l y brown t.yc rolot~r;111d r cnt~rrollingbiuc r1 culour. A l l r l e \ (or eyr uolour nldy cxlst 011 homolr Significance of a l l e l e s g o ~ nchrornn~o~nc\ Four pocsiblc t o l n b i n a t ~ u n 111 RR, Ar, rR or rr. 111 rhc hornozygous conditrrl ,tllclc<o n I~orrinlogouc L-hrornocnnlcsnrc of t l ~ c \,In.

Darwin, the species and natural selection

n.
>I

33

111

thc heteto~ygouscondition

rrrnt kltidc. Ilr .und rl?. 1911 ind~uidi~;~l cliatigcc in gcne I*' ir><~trgR will Iln\.t. brown c y ~ c ,w h ~ r c tlic ~ ~ ~ T ~ < - ~ i o t l . R .~
,I.YSO~

Evolution can thus he cnnudcrcd as haccd upon frcquenctes tlunugh i~atural

'9,

at' 17 will hc blllc' ~ v c d ULIZ c r j n n h ~ n ~ . thr Rr or r R o*iII.il\c7 lc.1d tt'r Isri>\v~~ C S . t l l ~ C~ fur I . I,. rJ :,. A. ,I: 1 Ir uwtllinant nvcr r. 2nd ~ n . ~ < irc cffrci. 11r 1 kc i,><ca r 1 ~2ld hc t t ~ c , tcr recessive gcur. Natuml sclrction vpcratcs upon tIlc &?cat rtock of ,+lrrtrtrmcr ,~llrlr\.Ire ~n~iltrpl~.; snnlc ~ c l l c fart. ~ I tn c x ~ it n L I (0 11 ~ 1 1 1 ~ 11 ~ I~I ~ I ~ J E~ l o o d v;irintinll illllcrerlt iit the Kcnc pool o f ,I poyul,~tion. I ~ I ~. l I ,rrc uontii~udllyrc~huflcd the mccE-t.~hy I.\, irrt r~\amylc., ,lrc c c ~ ~ ~ t r o l hvd ordv a ~ i n y l c Thc gr11e~ lc I nh~cli or.c.un a< three ,rllrltq d c < i q ~ n t e d q lmin dkcursed above in nc\v cnrnbillanonr or geno11, 1, 4llclcr 11 2nd 11h ~ v r cclu.31 rlnm~nancc ~vlirrras types. But i f evolving orpnisrris arc cvtLrto +c risc radicallv ncw there muLt be a murc-c. of thc lixir htlrn.111 blond po11p5 C2 to nrlvth~ng iic~n. vananon. Where doc? this x~,mation comc ' 11(7ninzy~t~rc &711p A mav carry P 13; +nup U qll nr qr, u,l~ilr g o u p AH irnrn? the l l l L LLlll l r ~ h ~~t t = r o ~ y < c : ~ l t ~ . ~ . ~ p17. Ir rnrnerrlrlcs happens that a getrc will huddrnlv t ~ t . L~tluvr. cx.irnplc-r dc.11 \vitl~ gci~csaffcrtrng and spontnncou~lv change tto a new allele. This kid o f rando111 chan!gt' ic known as A gene mutation. I t ~ l c h.lr.~~-ttrs, !hew :lrc incrrc rnmp!cu pntt but ir t h r~ = w lof a chemical change In che DNA helix: ~ t 1s a i \l~ri;tdun\\,lien t\vo nr niurr CcncF control or :Ic rc. T l i i ~ ~ 1-rliun.lgc cnlourr 111 p.lrakcen A wtr<tin~no~-rreatrangenlcnt of rlurleotidc bnsc.;. un :OH trol of' two gc~lc$ xvhich 111 dlffrr- or .a dliplication or loss of a rrnall part of thc I-i Inr ni,n yrcsdi~cr. l~fuc.p r c n , yellow ~ c y u c i l r t .T h u rimplcst kinds of mutatloll nnsc bv o j ~ ct~uclcotidc haw, or a pair of base5, being ~ r h ~tt r ~ t h c s .a ivtC1rr C L ~ T ~ I \ J I ~ I Ji~ ~COOfII OI U ~ c ~ bv Ir rruuld I yossi'tllr with ot~ty.I s~ndcp n ~ of rcpl~ccd anothcr b x e or base pair. This i s rt~l*r;tl, c t l'~\ tiition, hilt mutation5 cnn also anse h additJon r3r y tklction of h a w pairs and t l ~ i s may di~ruptthe film!'hrrc rnul.ce.: o i vari.ltron - cm$+inp ovcr. rc, I \ ~ I I I . ~ ~ I O tloinu- w d l i ~ t c r r v ~ g t > \ ~ ~ , II, inultiplr tion of thc g c n e Mutations on a larger ~calc~ 3 1 1 afict from tcrls to thousands of base pain. and It-,, ctc. - arc all cclnqcqucncc\ of dipluidv. III chrernosomal mutations. wl~rrl~r c the tnoqt a inon, ulally plants increase thctr porcnti.11 vari1l11lbr1 by polyploidy, 1.r. dupl~cationof chromorstrcrne kind, may ~ n v o l v c inversion, rearralipe~ l i ~ c d ~ l o ~ l l ng r, l greater gii~lnbcrof l o c ~ L)r allclr<. ~ ~ i c r l t , addition or dulction o f long rrqucnceq nf DNA. Thcve t n l d to llavc larger-scale phenotype I'ld<yli~~rlv. hnwevcs, I F r;~rcly tiilund 111 anlnldl~. I~ICCFFIVC ~ C I I C nipl lit b r espcctcd to giht lmt rffcct? a d arc njorc often disadvanrageous. ~ A l t l ~ o i i ~tl ~ i was fnrmcrly held that mntationq are ,I 1~ii~tvol1it1on, In t i c t the proportianc of allrlwlt It,, a ~ t h i ipnpul~trot~q l ~ ~ r not ttnr~ndlly c!l;qngc. comparativclv rnrc cvcntq, it. i q no\v apparent (llrakc ' ~ n l ~ -t\h ~ r IrF 3 <\~cc~.II l i ~ i ~ , l l + ~ advat~tagc ppotse+\- r f nl.. 1983) that Fpontancouq 'Icsiolls' arc contrnu1t1 one pfimculnr type nf nllrlr. If tllcrc is t ~ o tuull ally occurri~lgin s h r o m o ~ o m e sduring repl~caholl; lrllaqc a pnctlc cquilrhnun~ is ~rlwltair~cd in there m y be mailv pcr cell cach day. Htit such niisnr tlit. Hnrdy-Wcli~hcrgeqnation: pairnip arc coutinually h o ~ n g rcpaiterl by a complex e q n r systcm (DNA pnlyrncrx~c acting In conl~iilctionwith other cnzymcs). This acts as n 'proofrcadcr' which picks up wrongly psired ba~cq, ~ n d 'rc 1'' ~ n d rcprcqcnt the rrlativc proportlorls of putr thcrn hack in tllc right order. Even this proofq' ttn.tnt hulnoqgotc< (KIE) and rrcesrivc heterurending systeril sonirtitncf dipq up, and mutations )rn[rr), rc~pcrcivuly. Tllc cqt~ilibriiumso dcfined then are f-tucd in tlie rhrornnwnrc. tn have thcir nnlv hc ~ t i ~ f i c d whcn thcrc ir ,I drc~ded c l r c t ~ v erfftcts up017 the descendant. ~ ~tlr,igciri pn~srscing,I part~ciil~is ~llele. Thiir if a While 111utatin11shave bec!l tmdi tionally undertcul,lr phctlotypr 17 precenwI bv natural qclecstood as being due to DNA damage or mispmcrn. thc b ~ l ; ~ r ~~c1c1 1nltcr . ~ n d propornon of thr rng. it has recentlv becn &own that rnnny 1 the t ivithitl tllc popul.icirni u ~ l Incrcasc. l spontatlenus rnutatiom. C'FPCCI~II~V ~ I I O F C o n a Inr~er
I,'

itrrir

34

Evolution and the fossil record

~ c n l r may hc rl~rcLrt ~iiohilc c ~ ~ c clurnunts (vari, g t~c is not I~kcly rh,lr inutntmlrs xvill rprcad v r n (1st 111 a owlv nnilicd >urnping genes', 'transposons'. n r Insgr pnp~il.~tinr~.o t ~ l i thcv arc dom~n.lnt. z t ~ d cv cvrn 'molecular prasitcs'). \\ liich drc DNA r n u t a t ~ o ~ ~ ;lrc high. Hut 111ost t ~ ~ t i p o r l ~ l , ~ mtCr ~ p~ < ~ ' i ] ~ l C l l C " S .1h1~< l I?FO\'C :1111?1gC \ I T C I R I < I F C I ~ ~ ~0 r 5 . 1 1 nonq ~ r c .not cvcillv d~ktnbutcdthroughour thr'lr f C < 101 one c f l r c ~ n ~ n \ n ntcl c .rlj~>tlrcr. ~ TIir\. <.In altvl. thr qpcc~cc; raijgc. T h c v tcnd tn 1r rubdlr.lderl intn . wlielc gr.rtc.tlc str~ic.tlri-c.uf ,111 org~ilic~rl throt~gh rc1,itivcly m , ~ l l units ot ~ C I P L I ~ : I ~k110\1711 ,I? ~ O ~ , rc.n-ranyr.nlci1t o r c x p : ~ n \ i uc~r~i t < TrrlctIc nlcjteri.ll. garnodemes. Esaralplcs c,f si~clinatural yop~il.lr~orr l Scmc tcilcl to att,~chthcmrc.4vcr tcr parricul.lr wcr; y r o u p i t ~ g r:~bhitw,lrrel15. cchoofs ut'tirhcr, par~ri arc othcn <c*cttl, I ~ I c SO irt in , ~ I m o . ~tn n v h t r e .Thc cnnirnul~itic';, ~ the 6 u n . 1 ~ e n c l n ~ e d nf lagonrls a ~ d 111 , capncnn fi>r n ~ c ~ ~ c r nor t l l e ~ ~ rn 111ob11eor yztnf;l- thc h u n ~ ~ n <phcrc,tllc inhnhita~its m\vn+ , ~ n d of \?Istnhlc ~ c t l c t i rt'Ic~i~t'nt\ ~UCIJIJ\,I r.~Clicd311d r;lpid lagcq. Thcy rcnd to hc pi~rtlallviwlartd koni nrI1c.r 11trr.ittot~ o f t l i c pdttcrll n i p r n r esprecslon bv a quch ganic>dcmcq,hut nor co~ilplerely fcjr thnugk sn, ~ r n ~ y c r t i c c v c i ~ t- ,I p h r ~ ~ o n ~ r n o n ln 1%-Enrhhas c.~chp ~ n o d c r n c ~ r r i c s t s ~ v n c ~ o gcnc pool tl~crcir clc.~r irripl~ratiorl\Lor r.lp~drvnltrtlon n l ~ dfor t l ~ c come pmcihility of r s c l ~ a ~ i g e IICIIC'C 31id q c i i ~flnv; nngin of I I ~ X V tvpc'c ut'orqnn~z.lt~oi~. hcrwccn the pools. hcntab'c A II~IY .~llclc,f'ornltd llv ~ r ~ i ~ t . l t i n r ~ , I t i s within such nn.111 natural popul.ltiol~r t11,1t and will prnducc chatlyc5 In our or mnre chnr.lctcr~ f;~vourahlc tnutatiorls. eeprc~allvthnw nllorvinq llc'r1 1 thc ilry3mstn. Thcsc. rh,~tl<e:c'; lrfunlly of cmnll tcr ~C(;lptatic)~~~. 1 .lrc arc able to spre.id. A iiiut,lnt condlccdc .]tad uf lin~itcd t t k t : 1 they .ire I . ~ r ~ tl~cy c f c r arc tion c-qt,~hli~hed cvcn I o n e btnmly Ins!, n l i 11 vcn7 oticn lut11.1l. hlclrt tnutat~ons .ire iicutml. Gomc pnrsccqu\ sdcctive ~ d v ~ r l r ~ l p e . sprc,~d~limuyli roon 3rc ~ ~ F , I ~ V . I ~ ~ ~ I ~ C ' t ~U C ~ ~ I C IITI I J ~I C A ~ tn ncw or thc. p ~ p ~ ~ l ,it1 tthco gnmnden>c wlth~rirc,l,~t~vch h I ) C> . t ~ i ~~ m r n c a ~ h . i t ciiff rcrit clinr~crsm ~\pZl~cl~ msv c a F fr\v grncration~. It 111;rv then ~ p r r i r lt ( ~ nttlc't qomc .~r.lcctive; i i f v . i ~ l r a ~ ~ . ~ c l v a tagcuuc muta- p x i ~ o d r n l ein thc v i c ~ n ~ t )iucrcaqiny tlic c f k c t i \ c An n ~ ., tion. wl-tich IlIav ctnlv II.IVC J n i ~ t p n n l l y briirtjcld rnngr of the u s e f ~ ml l t n n t coiid~non. ~ Surli n rnuzint cfft.ct,will nrllv be ~1-3cto sprmd ~f (1) it ic rccurrcnt condition, especially ~f occurring in 3 peripheral w i t h ~ n rhc poyu1,itroil 40 rhnt net%, recruit5 arc isolate (i.c. a s i n ~ l e g~rnwdeitle.nr small cluctcr r)! continu.illv addrd to thc rnurnnt crack. (2) ir g a ~ n n d c ~ l ~ e s tlae fringes n i n lnrgcr ~vidccyrr.~d oil ranics rurilc selertrvr advanr~gt. w that fhc populatio~~), he the finr step toward7 thc frmn.4will Hardy-IWcl~tberg eql~ilibrrutn ~ h l t i ~ d I ~ Ffavnur tlun o f a new and disr~nct 19 ~n specrcx. The nccurnulaticlr ,111d (3) thc p o p ~ i l a t ~ o11r qtnnll c t i o ~ ~ g h ~t t o bc o l a fe\v more inutattnns rnny thcn i<ol.~tc corn. n fist ~t r~mid and spread ~-cl.~t~vt-ly ~lnickly. plrtclv, as ill rhr c l a w r I3ai-\t,ini:ln madc. Tllc orig!i All aUrlrs havt rc~iiltcii&(>in rnut,iticlrl: tnultigik of spec~esfro111 such periphcrnl 1xolntr.s i q kuowti ;llldt.v h ~ v c x-I\eil .it d~tf'crrnttiincs. 5 o m r inutatlt allopatric speciation ( r ~ ~ n c 2.3). Wlicn the mica ln allclcc r~iav p o ~ ~ t a ~ ~mzitntc l ~ ~ > g i n to tl~c. h - 1 7 ~ sterile wittl the pfirnt srnrk, Iuvinp rliverpcd s e o u ~ back is nriyinal qcnc csprrlclorl from uhic-h they arose. c l ~ o u q h grr~etrcnllv, thc 'parei~r' 2nd ' J ~ i i g h tcr' T h c ratcq of 1n~tt.iti~111Iwztll ti]? sr-1lC. SOIIIC \;n ' arc spccics n r t then c r ~ t ~ r e rli4titlct. ly niucla h ~ s h c rt1i.111 ntllcts. but rhcy ,lrc h~ricr;tll~ lLct-cssive niutntlons rprcnd c-nn>p~r.~t~vrJ\ %lr>~vlv low, onc n ~ ~ l ~ ; ~ lt t~~c 0 t0 1 ti)r .I p.irti~l~lar e n t ~ r l d E l l1 0 ~ oidy in urlall pnrr~lnnnnc ~vithclosc i 11hrt.c.dirie bcing c a n ~ i d c r r d I~igh. ,25 The t7tc of rrultat~onc ~ u I F thcrr any I~kclihaod of ulorc r ~ p i d+yrmd~n: Zt 1~1crca.icd\ rt~dl.itrnt~.t i c l ~ ~ d i n g ,. b ~ rxpo\trrc to Holvc~rer,recewve n ~ t ~ t ~ t t ~ o l i \OCCC>~~ICc.111 Junlrultravio1c.t light, u r 1 cl~etrucnlnlr.itnr. v . I I X H ~ i~ridrr certnln circun~qt,ulccq Fspcrirner~tt have c h o ~ m that chan_rcs in tcrnpcr.irurc .ilon. mag bc ctio~rgli ro rnndlfv tlir dornrnnncc of .. geuc. I r l n n h i r ~ ipoplilat~on< thcrc arc. Fomr prne' Sprecrd of mutufions through ~ 1 1 1 ccxist only ar modificm. c h a n g n rn \\~/lir-h h nu\ papulations alter doiiiinanct. to r r ~ r * r < l v c l l C < < ;I ~ C I I C , If' J 111 Mutat~rm1% t h c ynrtr:lry hourcc uf v.Irinnot1. but i t IF rcccsnvc ~ l l e l eif fiv011r11~1~ will 1~ w l ~ c t r d ~t 61: onlv t21rrrtish rhc. ctkr-cs r l f c l ~ p l ~ ~ J ~ o r g ~ i - AC)II-IIII~I~CC. 111 Iitghcr isms t h a t thc rnut~tiunq d n \prcdd . t~lellt~oncd. i k q ~t

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Darwin, the species and notv~ol sel~ction

tlation and species formation

35

genetic drift. hrcu~nesilrlportarlr in tl l c ~ c51n,ll t pnpuiatinn~. I'his n <miplv ~n rvnlut!c7tzap ch.lt~::v tr~rtuYdt i c w bc clC,lrthat w n l r k ~ n d f r c n l ~ t i n i ~ m prne f r r q u r n c i c ~ n ir ttirough r;~rldn~cl ~ , ~C.n .c r ~ ) r II n t,\~tn111\ cwcnt~nl for syc.c-jr.<f i ~ ~ n a t ~ c > t mnv hc .I l ~ l c n twrthout selection bcirip i nVolvt-tl. Hcnt c c r*rIt r . , .l~g,~pl~ir .I] rct~l,~tron. cnmc pnrr o f a pnp~llntlnn tain locnlrzed pop~iIarior~< r.c. iic~ct'~~cIt'cl I I I .ill 1~11r1,ll j; c > cut off frtlnt tlic rest h\ ,I hdmcr, 2 % norcd p ~ nr few paln IiIay hecemc qurt~ ~ I I ~ I I ~ I C ~ ~ L L . ~ r r. But thcrc are ntlicr k111tIx n f iqrhnrlnn \~,hich may cvcn dlfferentiatt. ns new rpt.cic\ H I C ~ C ~ \ br r.yu.lllv c.fTcctivc I ~ ccrt.~lltc.l\cs. Thrrc. 1% bccnuvc the folindcn o i t h e pclptilnt~cll~ I l~np!>cnrl-l to lgc.11ict31~tio11. \ ~ h t i l \pcci~'$C C ' U I ~ ;1~1~1pt~d it1 l ~ IC r:lT a pamcular wt of nllrles. 1urruhobjtat5in tllc sartlc gct~cralarea. I'hyioIn m y gcnc pool thc etTecn nf all t h r f;icrorr c r r w a nr ti~olo~,lval c t r ~ n AISO IFOIJIC pnp~ila- ~ ~ d c r c d f,lr are conrinu~llv lIIFCrat-tlll~,tllnr~ch l t~ niay m ,to thdt tlicv ~pccintt', c.g. C ~ . I ! I ~ L = F t h r tii~ilng m.iiiv biologists brl~c-ve tIrat Inon r l u ~ t a l i o ~.Irr ill ~s IIlrccdirlg cvclc ttr mcrclv qizi- clifkrcncc5 prcncutml ;lnd ocillr a t a more ar lcr5 cpn5t;lril rarc. In: cclpul~tion.Ay.1i11. ~ I ~ C T IIIJV L ~ c .I ~ l ~ ~ i p Gcnc f r c q ~ ~ c n c wc t h n ~ gene pool n r c rclatril n r C lc ~ ir the 'tic incun~pat~biliry; icrtilir,~cinnrn,~v tnkt plncc. thc chrornnsomc lcvcl to m u t n t ~ c ~ n , cm\rlnr: ovrr ~ ~ L T , I ~ Y C ciitTcrc,nt cllroiiiosnrric ltngths or {>I' ~ r l drcrotnbination. Thcy m;ly 1 3 ~'st;ll~1117~:d' ,I 111 tlon\ c?i allr.lcr, rcc-unlh~natit>n trnpo'i\il7lc .inti pnrticular kind of sclcction t h ~ tnutu;~ll\.prcvcnt< ir I1ybnd\ nlwrt In r.~rjhn.r~ Jrr ~tcrllc. nr cvolutinn i r i ,I I,lrgc population, fnr ~f rluc ~ p c s l c ~ ir 111'rcArc r ~ t l l I~ r Y ~111 wlilrh i.~ol.ltlc>n VJ might bc $ ~ i c c c ~ d ~ ~ l , wiJc.;prc-ad and well adapted to i n cnrrlr r c d . A pnp~il,ation~ - x p l < t ~ u t r II~IFII, dcnvcd ' o ~r" ronmcnt, t h t o i i d r ~ ~ m h l are \vccdcd unt -trid tllc ts II J f ~ i l l ~ d c r popillation. rn;lv initl.~ll~ cxpnnd OIIIV i-qwr.~tinrl,~l srlecrion I C stabilizing s~lcction. I lnlanv rwn arras in M hlcll rorr~pctirionor prcThir tl; an~logouqto what Mivarr pmpo3cd in Irlc III I\ rniii~nral. A h tlic tint HII+II. IICI\\JC~,CT, criciquc of D ~ n v i i ~ . thc ~ r h r r On hand, qcztc fieI k v ,111~equally~ sudden crash. which 1c:lvcs qucticics bcpn to ;ilter when thc rnwrniiincnt ~ ~ : J irw rtnall tsvZated p o p ~ i I , ~ t ijvidcly s . ~ ~ ~ ~ ccat- changcv to becnrnc castcr ar harhhcr nr whcn .I popI I I ? ~ ~r3ch \i-itll its own sarnplc ot'nrhc total gcnc ulntior nligrgratcq into a 11cw area and. ki11~11117littlr I I b e w In turn t11,iv act a5 foundurr of new cnnipctitinn. hcpnq to dift'crcntiatc c . c o l n ~ ~ c ~ l l v . :$, at Ica\t. sonlr of wllich may runrivc. WI~cre Thc ~ntcnsity f hclcctinn unclcr t h c ~ e o latrcr ell-cum1 1 1 , : r a i Y ~I J I ~ C ~ O ~ L I I . I I I C I I ~ I C hccn bur ivipcd 5 t ~ n c c ~ much LTCJ~C~.. F I I C ~ nIrlc5 rhr'rr 1 ~ 1 , i j r is At 1 qrinic ~ ~ t . ~ ~ t ~ o p I i c , tllr qvcirs nidv rcc0vc.r occur directional selcctio~i tnw;~tciv ollr cxtrvrrlc ir, 1 ,YFIIIL~ ,~q.~in. !i(>u w r , I L ]XI> p.iqscd t l v n ~ i q a ~ rc<ptmsc to A l ~ ~ ~ g - ~ ~ i i t i n i ~ {~~<~FCLli-~. e l s~lt~11~111 ~d 2nd llk bnttlrncrk' 111 ~ v h ih nluc t~ uf t l ~ r c puterltinl tY115 I.; sccn an thc fo<\ilrccorcl ,IS ~ 1 evolutionary 1 11t' <I I. hrcll lost. '1 Ilr lash of tlicst. allclcr I c n d ~ tscnd (though oitcu it 1s hard to rcc ~ E I L . rc.,IFnri fnr I:cllllnzvyrwt)., and sllc-11 I,irk of vnndhorl tndv tile trend) . A l t c n ~t~vely disruptive ~ c l c ion n I . I ~ ,~ c~ 111'1t8'ly prow c!etnnlcii~n!. occur; tlic rolnrtloncst phc.nc>n.pc. in n popul.~tion can l>c wlcctcd nkqi~i%t, i thc. dmrn ~.xtrclnC\nrr ~ ~ positively %elected, S'iwnuu!: twt, dift:rt-nr spccicc frcrtn at) 0rigi11~1t l O ~ 1 7 l d l l d ~ ~ t n h t ~ p r d ~ ~ l ~ i t l o l ~ . ~ t np Genetic drift: gene p o o l s One \~oulcithcrefi.,rc cxpect to h i d I r i tlic firwil ! o ~ ~ ! l u l n r ~ ir ~ i i c v c ~ ~ l ~ n it ~villcuntain otrly n r ll rccord relics uC l o i ~ g period5 ~ ' ~ ~ c c ' \tnl)ility, irhpIc's , .%~. n d i . ! cuidui~ls.ir~lplc thc tot.11 qcrictic v.iri- drntvd by hruaks rcprcccnrlns tirirc., \vli~-ti of ~llccin~iofi ' - v i r l r l ~ r i itllc rvholc ~pccic'i. i.c. only .I fractlun W ~ very rdpid. and t l ~ r spin qtablc pc~iodq. h n ir F l T . ,i\; wcr.lll >rlic poc~l' n i thnt ~pccics. 7'hc yen7 uticn ~ v l ~ ir t fn'ound, a$ will bc cuplatl~ctl.IT a :Ilk: :11< pnpul.~rion.thc f c u ~ c \vill bc thc ~ I l ~ l t ' xw n i e Icnptli In thc rwxt section. r , I ~ J I c ~ dh\ivt~t o fit~trtI gcncr;lnr>tl<. I11 a n y evolvi~ig 5yctert1 thc lrlnrc nt~cmcnvirnn! ~ I ~ rtlc1nsi<(>lily n ~ i c h r c ~ r ~ of ~ p i o , ~ n ~ I II c ~ J ~ r rnt.l~ts tllcre arc, tlit grratcr ~vill tllc* prr\\thil~r!r ot br :. ! I : I . r,~lirln~n 53tnplc c ~ i ; ~ l l ~ ' P.ISCL-CI , AC~WII to A~tErentiatiot~. 1% I ~ ~ Species thus unn be 5a1d t o arlw 111 ":, ,:,~rnctc\:1 1 ~ ;~llclcx ,1 ui ,I [illy popula tinn rhnt scrpollse to the cnv1rclnmcnt5 ~ c t ~ l a l avail.~l,lc. lv Iiwr t~~~-ol~~hincrl in 5-rtilization arc. rhcrcfnrc It 1%well kilown t l i ~ rlatural ~clcction 4 ~ p ~ r ~ ~ i l i g t i ,.I 1::.1 .p~r.i;~l cvolut:c~n,il?;proccsq, knt>wii AF ~t thc prcscrlt tirrlr (Ei~dlcr, Ic)Sfi). Evidcncc fru- thi\
-i l.
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36

Evolution and the fossil record

t + t > n i ~ t T I I , II I - I ~ W I ~ ~ . ccuirccq.

Tclp1c.11 ~\.1111plc\ O L I ~ ~ tuikcrcr'. The w h o l e orytusr71 lr n sinqle rv\rc%nl C and thc ar1c1 Lir~znnuo f itc cntnpui-Hrn im-llidr t h r rrsc o i rrb\muoim,ltcr\l\-tc<t<ttllit c;tSdin! <>i ~lm-pI~olol;.y pnrtc h.we t bc compronuwc amonpt rc>rtyv,tll~ 1 IIL.b.lt ,ll ; o mbhitr o r , ~ i i r i h ~ c - r t ~ z - r ~ . ~ i c t ~ ~ ~The c,l\c o f h.lctcri.1. ~ t T l l ~ * b l ! \ <<> t h dein;~ndsand bascd upon Fcnctlc and phyic.11 ni,l 'inrFil\tn,~lrnc%l;lnr\~n' ~aic)tli\ hccn r.utr~~5~\.elv in II,IT , lI>lL..I b j - 0 I hc .;turlri.d .r~-td i t . Hcfcrrc. thc Industnnl terial which tnay ~tsclf lunliiny. I , > t I l ~ * Il For csarnplc, the lenrer of .I tnlohirc'c cyt- .I. l-lcvtd~~tror~ the. north of E~ylarld,the nloths In o : g v l l t , 'blltl c ,1 inhabitir~t: thtv I~c~l~~-n-rnvc=rr.~l ~ tree$ l ~ a d t m i i b tbf a made of mlclrc (Clisptcr 1 1 ) . JL ic ~ h rc5t of t l ~ c11' , 1 1 1 t I 07 r:.rlil t p.11~ \pCck.lrd c ~ ~ l uh,irrilc~iiizii~q 1 t thc crlvirw- mal's cuticle. Prrniln.li~ly r i l n b ~ t u\Ircrc. uiinblc r ~ir 3~ h secrete ni.~t~ri+il than cnlcirc for any part r ~ f t l t s I * t r I I X ~ I C A I orElcr 1 t w r t 1114 T ~ I I c Z Ctllr~li# I I C O ~ I S ~ I C ~ I When. ~~~ I O~~~ ~ ic ~ , rccclrclk lvcrr tint kept 11) the carlv 15(51k t h c ~ c rxoqkclernii d u c to inh~-rcnt c r ~ c t I i n i ~ t . ~ t i r,lrl i ~ ~7-1 l , l [ l ' [ < I <' I,~~II'~LLII although calcite hac the urrtur o f tr.~nsparciicy,I* riiotl~\ 1 l . d LIP' E ' J i ' r ~ O~'TEIC' p ~ l p l t I , ~ t ~die~other I1'L 1 ~ ol : <I uJcre d.~rl:-colirllrt.ili~ npr? which \vt.rre more eas~ly dirndvnr~tngefor 211 nybc.ll yrcrn ic ~ h d itq n Itiyj:18 I l l TI><+ I 1 1 1 1 \ ,!~\'C n 1-iokcd our bv ~ r ~ - d ~ t Hut . the end o f thc ccrl- h i e c h ~ l g r t niiiicr.il. psnducl ng douhlc i i i l ~ y \ on L I I ~ Iti11111 7 difr't.rcnr depths, f i r 1fgl:httrnvc-llitlg t l ~ r n ~iti rn l211, ~ ~ tun. thr. wc~odlr ~ t i *nr.ir tire citrcc hnd brcorne poli ( I I I ~ of Kt= ;r Iutccl .lnJ thr. I~l.Ocll~,J I ~ C I I 2rr w r y scnvtlvr to other dlrcction\ than thc t n u ~ r r ~ l oig ~ l <-,?\I W t r g x t l i kji1d1 u r ~ c l r ~ Jtr. I>.III I l c c . ~ i krllcd, cx7miny thc b1nckent.d Trilnhitc~ha1.c ovcrcnrnc thiq di\advnri t;yc fint 11 in n t l trc'e t n i i l k ~ tx=loi~r 'I'he vnst m y o n n o f thc rnlothf In c>ricntatiiig thc. ~ - . ~ s in\n ~ ~ no tlie vi.;u.ll susf~u r ) ) l ~Irclll3r I,\L,Il \I of t11c.w JlC'Jr WLTC t J 1 ~ 1 1 t l ~ c dirk-cnlni~rcd tbr~iis,thc and cccoiid h y r n a t l y rngctlirj~l\rnuc!ifir~tic~rl~t h p l r v.~rir-tie< ~ ~ ~ c t ~ ~ i i t110 nmore than l"0. crysr:~lI,~tnc-c;thcv II.IVC JIW ~v01vc'd ICII\ dr>uljlr.. l l ~ ~ ~ l t ! r r , for ~ g i \ t r.!~<illltl fouu\. I11 <pltc, tllct, S~ncc. it had brco~llc. .idvails.~ccour ro br d ~ r k - m-hich brill9 11q111 trr n c11~1t-p I ~ I I L . !:~,llet prcrrurl cnlourc-d 50 tf1.1t prctl.itor< 1vo~1lJ lcw cari1y scc thc~ fnrc. o f ,In iilrnn61c lilnitation. \rlcctirb~~ , I \ Il.lllllC c F moth5 ni1 tlie d.wk rsut~kr. r1nt11r;tlcclrction had bet-n ha$ 'fnrccd' t l i ~ , Ilc\t fi~nctiur~:i! Y F ~ C I I I t l ~ l tvrl trlr.llt Ic' rntciw in thc sl~.~nt;.rr~g cnvlronmcnt. Soiiic 311 p0~viZ3l~. 1 Ii~rrl:.~pt Such ~ r ~ n t u i ~ ~ i c aptlvqiological ~ d ~ ~ p t , ~ ni o i ~ s and l t spectec i l f i n n t l ~ krirr\vn t c i h.lvr hrr-11 3ffl.c-tcd b y arc. l't*rl~,ip FHCII i n d u ~ t r ~ nrlr i c l ~ t r ~ <xvitliir~ ihc vcrv qhort onc rcs~iltof nnturnl ~clcrtiun.but a1urn:ll pop1111 n~ I.. tll.lt 111. alqn ad.lptcd to tlrc~runviruzlrrlcrlt rcTrc1 tinle<p.lll '10 vcnn I3t1t only .I few grncc .lrc tions ,~rc rht=\+ aft-rtrti htxrc; t r rr-(l~iir~-fw r r chnnwc r inihdtc ducrivcly and tropli~callv. 111 this, rrrpcct, atld I,, 11i.11i n o gcr~cml rcnr~r. two kindc of ,tdapt~: t I r i l l ~ c , ~ n ~pc'c~att"ni~. n l v i r ~ g ~iw :r pcnrt-r variety o t ' g c ~ i c ~ r-ton- !>mad ~ n d ~tratc+c-il ~ a v c bucn Jirtm~q~isl~erl. arc d ~ t l k ~ l ~ ~ I . l t i . < !KC1 'I'11~y trollrri~:ditTcrcnt I-11nr.lctcrr. d11d r e p i - o d ~ ~ f iOr1 3 l l l l l l f ~ ~ l l t ~ , l n l ~~g A rnnrc rcccnt cu.uilplc c-nnccnir chc r;dnIc. cnt WRY< 0 f F U N 1 V I I I ~ I < , I t 1.t h,lt Gn1apnyo:nc f i r t c l ~ ~ C ,; + ' ~ V ~ Z = ~ I ) 1 1 ~ ( ~ rtu<li~ri I3~nvan. cnurgy budget. Whdt src knnwil .I< 'r-.;tmtcgr~ II 4 ~ n N o r, O n c cpcc7r.u 1% ,I .;pcri 111'it Ii.c~lcr or1 \ttrall 5ccd5. Ilavc very Illell fecundity s that p a t iiu~iibcrc 11!,111~ier : rapidlv. Ir~dtvldt~nl'i b mdv Fc>llolv~rlg 3c.ttcrc. drnuglit 11-1 1077. rhc av,~ilnhilicv uffqpring arc prudu~cd .I I ) N A t!I,l1 o f ~nlnll5ccil.i hccatnr. ~ ~ l i i c k rctlirrvcl. Biyycr rccdq 5hort-livcd. rclyiilg upnn tllcit supenor reproiiul ~ O' I~ou~c\.cr, wrrv irir,rrb : ~ h u n J ~ t ~ t . t l l o ~1;aryt.r tlve nhilitics for spccles F I I W I V ; ~ ~ '. K - s t ~ ; ~ t r ~ p i ~ t ~ 'I .I I ~ I ~ C h:~ 2nd bird? 1~1ths r r c r n ~ ~ - r b~-.lk~h t r,71iId CI)PC u ~ l t htlltlfe thc other harld, produl-e fewer .off<pnilg, h t ~ rJ., hdjrlrc r l < I I t !:CTII'. I1111 nild tinit wcrc nhlc to \urc'~uc.I t li)llo\w tEslt t h c prcvsurec of ufuallv slow g~ow1115 live a much l o l ~ g r r n a t u r ~ lsr.1c.c tir~iiudn; Crorn p l , ~ c c pldcc nrld fro111 they arc better at competing fnr food nnd Iluir~ !T-ILL> L l t i l r i to ~lllllllpl~ c ?car t o y ~ . i r t t d ~ thElt~IICTL- 1\ 11t1 T ~ . J S O Twh!~ ~ ~ n a l l - rpacc- T1-~iri s an a t t n c r ~ v econcept 2nd in con! I rcsprcts still ~ ~ t e f ~It l1195. hn~vc-\,ct.hccn hra~ll. 4 >plr\ o f t i . ~cnlrcvc~lutinn\ l ~ o u l i lt l t b t bc rcvcrrll~lc. 111 tflc cu.implc. ~ ~ u t t t c d sbovc- thr \clectivr ~ d v a r l - criticized. s111ccso111c Fpecles have both high fccr~n I !l%I'. r<>171 tngc tni rhc grhnci\ clcar. but i n mnnv c,lrer. ~t 15 lefs dity nlld s~rpcriorcompct~nvrrlcw,;tud .ill por~rbl l i ~ < ~ 1 1 ? 1 , - ,(3 t*c\ 1P ) c.l\v tr>uiidr.r\~;lr~d rclcctivc ,idv;l~~t.lyt'r pnrtic- conlb~nntionr; of tlle~c C I I C I - ~ O H I ~ F 1113)' nc( \I thc nf i i.rluiirl.~tit (Stcan~s,1977). Furrl-rer~nore.tlic coliccpl of rli iller c11.ir~ctcn. 1 1 11\ I It.,lt Scltction, thrn, i s srcrl .Is 3 pnman7 ~ g c ~irl trvu- 'Iluuted enerp' Is~ldpct'1s not lnvnnahlv vnlid. It i ~ L E ~ I OIt I doc\ i11)t. 111nvrver, p r o d ~ ~ c c I idcal Kcno- thcrrtbre not very r.lsy ro catcgorizc adnprivc <tr.lt; ' I ! , ( ) t i < * Ill $ y, hpcs. hut otll~. tlic hest ~va~lahlc ?;.lrrcn the pes very ~ ~ r n p land ~t vccrnx rathcr t h a t ~~nptilnhur~ , l l ! l t ' ~ l T 1 iiLr1IEcro1Ix C O ~ ~ E ~ ~ . I 1vliic1-i opt.r.lre at anv t m c . will tcnd tn nl;lsi nl1z.t. whntcvcr rcpmdiirrir~ca)., t fli <.t\ ,)I-. III~\ ~l T h u ~ .IF Jncob (1'1771 hag well s.ikd. 'Naturnl ~ E c c - ~ u r v i v potrnt1.11 tllcy have. 1t1 thc c i r c n i n ~ t ~ n Ic c ~ ll~~lI~lrr! . , ltp!r<: 17 { tinn docs riot work like ,In cHg,lnrt.r. I t work5 likc 3 which rhcv find thr~nscl~cs.
1 1 6 4

..<

Darwin, the species and natural selection 'chiclcic-d*compoticmt. It iiiny cvrn. nftcr ii~i~c-11v o r Il~rlonaryrl~vc.rgcticc,tilrii out to hc ~ ~ s c fs ~some u l i :ral gr~rcticc makc\ riw of htccdrilr: rspencatr-prv, to ,lrl c \ ~ c i ~ t ~d ,ei~ c c n d a n r i l nrg=~;;i<rIi. +n tll.lt ~ I I C qcncr ,ire irndrnrood. ;lr t;fr ,I< ptlrTlic 'ninplilicd' ,1rnrliant of I nforrrr.lrion <.>I-ned in t h r t n ~ ~thc.~r ~.tK.ct<. h Molc~-~tl;tr ycrletrrs. u11 ~ i i u l t r ~ c t ihlriilicq mav, in \VJYP not .IF vct firll\~ c 1thl.r h.~rlti.conccrbtr.itrc 1tl~o11 ~ J C I I T C the undcrhtoud, I3c ncccFr,Iry fifrlr t h ~inniiitcn.tiicc 13fthc tlir ' , ~ n t lulw)i~l i o ~ v gcrw-~~tlc 111,1tcnalI \ 5tmcturcd complcx ftrnctit>ns a i d intcmr.trc\ri- c l f rhc cell., nf ir+1ni7c.c! N r w t h a t trn\.c r n ~ c r r d h a w < rjwtaru;ln\. I t had 11rdcud I>ccti \uyycstc=d rh.~ttlic rhe I\~IL..I~ ~.voIzrric~n tl~con.. i r wnuld scmm apprn- Cntnhnnn 'rxplos~on' o f rncLa;.o,Ill 11fc i 1 1 . i ~ IIC' :tn ctrns~clrrtlic ~ ~ n l m r r ~ I C ~~ V C C O Y C ~ I C I I 111 rr'Idt~11 tlir ~iiitialcrncrgciicc 4 1 t - quch i ~ i t i l t i ~ t . r i ~ to . ~ ~ l ,gcr~cttc.~. ~ I C T ~tl~rw I ?r W , I S arc rcvnlut~oai- Li~~~ilics'.i c t ~ Wl tnnrc i q klio\\+t)of their iiirl~tiot7c.i r , tC1c rht.m~~cal pntli\\b.lvr tl7.1t Ic.irl h n ~ n qct,cf t r ~ sri,~v pcrlrap~allo\v :h fustlicr ricvcli)pn>cnt of thi\ 11; il.loyrri, c ) r g n t ~ l ~ t ~ ~cr 5lt1ll F,~r<clv ,t c ~inl;nrr\cn. cuncept. For the Irlnrrlcnt, t n l ~ l t i y c r ~ c f;li~iilic-.; < ~ . c . 1 1 1 , I # ' rl11t11 17 t. I~,lvc. cl~.,~rc.r .i pll.turch rrf rht. rr.lar~un- tc, be pnman; organrz.~cic~t~,il ilnirh tvirliln thC .Ilr < c r ~ r \ r ( ~ d~.vc.lop~nc.nt v c ~ <h,411 snll bc fir pctlome, ,111d ,111 l ~ ~ t r ~ t parti rof ;I cIvii.irt~ic ,111d ~c I 1111~lcnt.l11r4ing \ ~c.uo111tiot1\ttorkr nt thc hn\ liltcr,lctrve ~ v > t c ofn~ c l ~ ccot~trol;~trJ ~ t~c ~-cr:rilnti<~i~. ul11.(,lnil IiCr~rc nl+r)+tt~:~clc) thr. Ic\*cl. ' ~ I I c ~ c IlCl\' CDIlCCPrS l l d ~ c~ ~I C ) ~ \ V C ) P O F \ I ~ I C ' I c L ~ ~ \ v I n t 11~1% :11r~..1(4v ~ r v i i c ~ t a ~ ~ l ~ 15 l i challt-ryec tn 11co-13nnvi1i1.;ni. l ~ cd Tlic lii.;t n Killiur.i'< >I7ui! lctr zintIcr;t.ir~~i~r.~ J~ithcrrtrclu\ir.c a5pcctq (1 ')7C))'ncutl-rh rhcory of e ~ , o l ~ l r i ~TIII~ r n p i ) ~ ~ . i I l i ~ ' .p ~ ~ ~ l i tllcurv, n1;)r i t illrltvr r h , ~ trhc f:cc~~omt.t h ~ p o p u l ~ t l o n ~ irluclr niorc dl\ clxc t h ~ \~'Oiilrl ~ t ~ ) t arc r~ irrllrtir .~pl.mr.~tu+) .I t ~ ~ i t - l1 i1 1 1 1 ~ l~~ i ~nild hc euprcted from natural ~electie)!~.h c lc\.cl cnt' i~ t d T ~ I I + Y + ~.< T I t11.1n f i v ~ n ~ r , \ i p p ~ ~ lvi I l ~ II~ lt ~ t , dl(vnn.ltinli slio~ild much lower I ~ ~ T , ~ L S W diKbc 'ucrf~il' I I o I ~ ~ I L - I L .c~rnpkc.urt\ ( c . ~ . t'crcncrc chould qprrad rIrroug1-1tlit. p c ~ p ~ ~ l , ~ t;rnJ . ion lplll~r ! #I/.,l L ) S 2 ) . t 'hnnnh11' v.lt-~ants h n u l i lhc cliini~ ~ ~.rred. 5111ic thcrc ,rI~i!,+ tllv tntlst iinport.~ttt ~IIC'IC O ~ ~ C U ~ V ; I ~ iIc ~~IrIt ~ ~ n l l yhrgl~d e ~ r c - rof cvidcirt ~ . n r i , ~ h ~ l ut ~ ,w t l i v I 11 I I I ; ~ I I Y C L ~ ~ A W O ~ C E c,~n-vriit~rh morc 13Nh o i i r r G n n t l ~ ~ n g do w ~ t h ro naturdl qr1t.ctro11. [t .inccr, ! zhri nrrd tbt C O ~ ~ pror(~i~i\. I T ~ TIIIC ' s i ~ t.I< tin111ch;ltlrc d ~ f f r r c t ~ c c ~ l ~ tn~cdert~l,ir ~ q at t Icvc.1. ,trl~i I < I I I L ~of C ~ J ~ I C LIF L I W Y:IITIC y ~ ~ 1 1 r <rt\ of + or mnqr o f the v;lri;ltion ~ ~ c ~ tah e r i i e c 1-rtarcl\rill d~m lior ril <t.ncr. Such rnult~plc.rchpc.hnon\+ccrn rn i ~ c i n d ~ v n ~ - I u ~ l ' ~ s~lwlval. It 15 pt~relt, ~leutr,ll, ~ n d 11:llqle~r. rspvi-idly ~ i 1 - thcre sccr~lcto hc* t ~ o whcdicr gciics for partrrular chnractcr~ i ~ ~ perci5t or <Its: rtl lt~r)nfli~p t ~ v r c ntlic .ltnr>ilrlt t9f crrccrs br appcrir if hv rhanrr alont. T h l I~F 3 d ~ h ~ t d b 1ptl111t. .e l ,111dthe uze or cuiiiplcsrty ( ~ f c l l nri~rti~~l; c ca1.1- but cotivcscly it could he a r p c d thnr even ifvthn1 ~krrdnli p r i n i t ~ v c l t \ l w ~ C ~ I I I L -m o w ~ rnniiy differences arc ~lcutml. rlioqe ieiv tll,~tdu curl1tl:ln 111o\t n1~irnni.13~ 5i1cli tlncwcr~tralgc.nc,trc fcr n fclective advnntnjic arc smilable to ljt' ~ - I ~ u ~ r l ~ l c r i n 11 11;15 brcn c:illcd ' r c ~ l ~ ~ t ~ c i , i n t' ur )unkS DNA. hv tlic redonbra blc pow-cr of n;atucll k r l r c t ~ o n . o r ~ rcinsistr of rnattv rcpctitiolis crf thc ~ a m c t Ant>tlrcr c h a l l c n ~ c rlenviilg Crmn the cur~crptn E , tult nwht ~ t ' t i i rcxccq\ 1JN.4 r \ . ~ < t s ,is multi- multigcne E7;Trnrlles I S Dovcr'r (li)H3) tl~enw ii~olcnf r ~ l i ~ ~ l i t1.e. F v u y r at' rcpcatcd cctlurrlcrs or .<, cular drive. It is hnrrd on tJ;c undentailtiin~th,it I ~ I I ~ C cuprrs ol' chc w n c gc-rlc. Suc.11 mult~plc indivichlal membrr+ of a mu1tigcnc E~~iiilv sl>ow. y - r ~ t :- r ' ~ ~ o n - r o rsrcliirllccs cvict too. 311d 111111n lr~~~ ~jrnilnrity within a spccles. Thc iiidrvrdu,il ~ n t l n b r n rlll>:t~of JU ~SPCC OF ~ C I WC C ~ ~ ~ I U I I ~I)c Cn f n species do nor cr~nlve~ n d e p ~ r ~ d r n but drc CJ11 . C tly 1t,.,11 qulrc ~ ~ ~ ~ cx~ri,bblr.7uYrl1at doc.< thiq 1;1rgclvIiurno~cnizcd In n ph+nnnienui~krlon-n a5 itr lclv IjNA, which ~t tint \igllt ~ p p c ~ n hc 'concerted evolution'. Accordtnq tu Uovrr t t n ~ to 1s , I > . l l r ~ t .rrr~i,iIly . dn? Sor~lcf i ~ ~ ~ c ,ire trcbnucrn- due r o fiua tlon of n~~~rnciorlqt l i ~ a a popu1.i rmn t~ ~t~\ wi ~ ( 1 4 1 1 . .4 n~iiltlsrnt-hlrl~l\., for rsaniplc. cdn t,cc.~~~<c mnlrculnr n~cchn111~1115 of w~tllin tlir llllnl r l o s t l ~ rclatcd prntrlii\ ,lt onc .und thc' gennlnc. Thir hc c-nllc molccl~lardrive. A n u n ~ b c r t1111c.rrtlirr than p i h i O ~ I CM ~ ~ c u \ c ~ , o f ' ~ i ~ r molcc~ilnrn i c c h a n ~ m i c . ~ . . tlic' .h ( gnrr ur loss of !\ t ~ a n l u t n i l r gcnc, iq.lt~cl~ t rrthrnviw ~ r ~ l ~hcr t ~ct-tionr gncrmc hv rmiirpnciti on) (nn oprr.lte to l of irn,jrlt, n7iII 1r b l i i f C ~ c L ~ v t11r otllcr q c n c 7 nlluw gcnrs rn rnultipIv o u t nfphnw i v ~ t h orgmthr 111~.rloc: n o 11arn1 hut s t ~ v sIn thc Sytcrn ,IS ,I ikrn thnt hc;lrq thcm. Tire gent- roiltrolIinq clitkrcllt II
oleculur genetics and evolution
I,' 11 m m
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d
c

~t

11-

1'

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38

Evolution and the Fossil record

X I I C ~ h~~n7otit.il

\pcclEll In( ch;~rrcttbn<- I r I tlllt~ ~TZSI.CL~CC i e q t i e r ~ ~ v e to j i ~ ~ r w r x e in wclurnct.. Fmt. maternal genes crtabit1 f du q I T ~ ~ ~ ~ ~ ~ ~ ~ I .~ ~ i- ~ d~ I to T,lllhr eEect4 h5h the antero-posrenor ,)lid d u r < ~ 1 - ~ r ~ 1XYC, i l pmucw-c. L nilt D I , A tr~ o f mirur,~l~rlct.~inii Molel-ul.ir clnvr c;ln thlis f x J the egg, before ttrtiIizntto11. Scc~md, i segmentatinn s.in he esr.1 v s r i ~ n t .I tnult~yrtlc f > t r ~ l l v,tiid cuc.li tisrd varintrts. genes come illto opcrariot~ it1 I , aftr-r thc c ~ 15 Kprtiliz~d. ul~dcrctcloc ? c t ) n ~ t ~ r ~tt~ vn l ~ .nrrcr In I breed in^ p~up111~t1on. ll il . give a i d tiley define the numbrr drlrl pocltlon of <cy- n q . .ind at cvt ruclr I l ~ ) n ~ n r r . n ~ ~ t v ..i L-oncertcclp-irtem of t ~ s n - mcntc 111 the hody. T h e rhrrd ~ c l ~ c'oii<ict\ 111 n~liiilw Such ip tin11 dt*tjnr*\ hlolfipc.il i l l ~ c n i ~ n l l u ~ nhnrn n n t ef homeotic gcnps: tl~ekecorltrol the unlcluc idcrlt~n~ (Jf cq11. F ~ C ' C I ~t ( > t h ~ ~ I C Y T.ltld ni,ly it~glc'cil ~OIIC mc.lnf of C ' I c o f a segolrnt. 1.c. lio\i~it I + ~011strii~ .111d \vl~,lt frr>111tlic t ted ir nni$r, at h i n I ~ > ! ~ c .l~ l ~ + c l h ' . ~ r~ MCIICCIII.T~ in coil- looks I ~ k e T h c ~ c drive h:~vc. varic.~ . srru~ work 011 aiid iiitt.h~ate to tll,, ~ c i ~ ~ i u m:1v c . ~ ~ l n t h c r ~ l c Ilc w.\y i r l wliich qprci~q~ ~ 1 1 1 qiqinlc produccd by thc qcpnenmtmon grnrr. PI I1;rs 17ci-11 ~ r i \ c . .~r-r-i~lt.nt.~ll\.,d IIV non-Mcndrli;n~ causc.5. nlllK,lllOll In .-I homeotic g?ilr corr~plcs .~n 111:iv pmducl I~c'n vcw Thuk \ V ~ I ! ( . r ~ , i t l l r ; l l \ ~ I ~ c t is otrL~rfi~onall\r i ~~ under- n dlipE~cntrw g i l e n t , re~ultinp.for rc,a~rrplc.r r a f l ~ r v c n over l I-.ire rhnt E~CILWI .'< i ~ p c ~ ~.it \ lt ~ Ic'vc'!~ ~of thr n r ~ G ~ n i s r n 1% 1wt11 four pcrfcctlv Conned w i r l q ~irlsteaduC hvo. ;r ir ~ mid lilnd,~n~~nt;lllv l . ~ ~ r i vmolt-cuI,~r dnvc rs non~c c, Tlic-sctlircc group^ of penes ~ c nil easlt tltlirr 11, ~ltlc\not 1 t . ~ d a p t i \ ,17irl c+pu.\tfi.it ttlc t c ~ c l tlic g c n n n ~ I t . s r c l ~ e i i c ~ pre~umahlyc n d e for proteiilc tli;lt u ~ of ~ \t,1rt\. %?TI nnd m.~y a h Itc cr>r~tr,~stcd with t:cr~ctic drift, w l i ~ c h turn Iinvc t l i e ~ rc%ct oil other griier, u i c l ~ i ~ l i l ~l:u t ~ o i lr i r t opcr.irl-r I C t h It-vcl octhc orprusrn I w t is ncm-nday~ rie~llvd ~ v thost. cnn trolling stnicturc. Snmc qcrlec (1313 tmlal ,~J.lptc*il ti) ttvc. hlr>Yc.citl,~r 4irtt.c m.iv \vcll prrlvc to bc rhc third seyinrntnl and crpeclnlly hntneotic) c o i i t ~ i i\ ~sl-tun , c+liw cd\pcr ~ . t t i o n ;hut I I I < h.?rd to fcc Ilnw z.ipifiIi~glily cnnscnrative D N A wcltierlcr (or ~ t l o t ~ l ' radiation! CRllt .icI;~pt,~tinl~ ~llj!:I~r .ITICC thcrcfrorn. known 35 the hornenbox. It 15 1XO bare palm Iunc rvolutlcm N r ~ t l l c ru tlri. 'tic-~~rral' o r t f ~ c'inolcc-ulnr drivr' , ~ n d C tiyr~p1 rnnstnnt in vzc a n d &ape. 'l'lie 11t~uenhnu w.lL ~tr.3 thcnncc F[:I-IOM~!J. I I C ~ ~ ~ T ~ I H C F II nco-nanvinicrn. 6nt dircovercd in homcotic gene? 111 IJlnco~rlriE~i \olllllj. rllc to They 110,Jjni~,c.t'ci-.TAICCt l q~irstiurlo f llow ni~icfi IOX4, but hnc nnw her11 shnwii to hc hnr~~nlnq+:tlu$ tl1u cuu ~ of thc \ ~ ~ r i . ~ l i n.?SCL' I C ~ I I C ~ C~ C C J L ~ of ( . h ~ n c - ~ % varint~?i n ~ e r t c b r ~poupr. including prii-n~tki llatLlrc d nii W in tr r ~1011r~ 1 1 d liio~v [I\LI~-II , \ C C I I I I I ~ ~ ~ I l~ C Cc~ c ~ it ~ mct~7n,lnx. h u t n l ~ o ti vcrtrhmtcs, ~ i l c l ~ ~ d i ~ ~ c : e d ,ti11 11.15 ~ ~ i~c d lt f l l l l , 7cc. 1 1 act!r,~llvcon li-rr r \r.lcctivc .~civ.inragc. h u r n n n ~ Hoinrohns-coiit:\i~iii~p . ge~iec nlwt be vrr I ~ ~ r t h k Zi i . ; i ~ \ . ~ r i c k.I[ t l ~ u o l c c . ~ ~ lcvcl iliclucic nnclcnt. at l r n ~ 5 0 0 Ma old. cincc they nrr found c m l.~r Thew r trar?$pnwix, J I I L ? c:1111.~ tivrlv tliic nc.w ~txL>rnm,~t~or~v c n c l v .and tlicy alrvny F~ii~ctiir~i wpneilttl ~Eetail111 th d~ in r31e outce n ~ ~ ~ iA V C t hc.~ri~ir: I ~ 3 lipon m p d but coordin~tcd anim:lE< in conrrrrlling t h r r p o t ~ ~ l cxpiyrccxinn nf tl:, ~ Y O ~ U T ~ C I I I Jc l C .~ ~ l t '.!irrl the ~1r1qi11 ~ I Z W r p u i T~ c-, ut' n body axis. Thcy arc. hon,rv~*s. prrqcnt 111 nnn-rrg- gct~eticicts ~ n t r o n n pJrtrrfiE l clmcntcd phyla ;and cvidcntly thcy d o riot nlw.~\ I'he c l i , ~ \ C I C C ~ ~ V1 C npcratc in tllc snme way. It qecrnc t h ~ three high1 t tdkr Il1JI1\ con~crvativr h o l ~ ~ r n h o gcr,c~ can t-rrol,t orljc r s hlncrions, snd w. in thc cvolunon nf dc\~clnpnicnr,~l rcpreelciltel Gene ragulati;on during development l.c.~ltl~ng p PTC>CC\SPF 'tlir r;.lnIc rcpilator). dcmcnn arc rccnnlK q ~ c ,in ,~du)r t~ cvF:,llil<rn ( l c i ~ c l o hont n frrtl11 l~~ zcd I>ittcd into ncw dcvdoptncntal n~;~ciiincc* (Rnh cted to r 3 c r y . it clucq FII , I ~ :~~.cftrig .1 prrdete~-r~?ined L WI pnch- f 0%). Clearly thr hnrncnbox ccv~ccpth n ~ n c v - cvnlut~nn o Micme! \V,IV, corrtrnllctl IIL \pr.r.~fic pcrlcs Thrrtl yerres nsc dcnt hearing on unc o f t h t m q u r nrcnr of cvclluriru>r\x*~tch~cl : ~ t l r l uffLl r i r e q ~ t a rrcqlrenrc. and nny a r v yalaeontc,lop?.;h o w nc\v body plan? origlrlltl on ~ ~ ~ l t h51% ll crror in this prticcs, r uulil hc Ictli.~l. r p r o d ~ ~ of d n d how they can bc ~nodificd. In thtr ctrtitc\' k01~1 Tll cr one unc grii~, IS Iikclv ti, I~avr.n st.qucnrl~ll ctlicct oil p , ~ l ; ~ t = o n t o lcupplic~rhc tirnc dir~\cil<ion. l o~y wh~l the cltl~c utltrr rrcrwc, . ~ t r t l Ii~nt:-tr.mi rlc.i~rlnl>intnt v can can be ir~tcyrntcd ~ r lvith scnctlc and n~olcclilnri1.1t.1 cprclc\ lulr clrv1r;lzc A 'c..I\c , I L ~ CIC !:r.iit'hc C L ~ I ~ O I C . TI~IIE o gcne ~ ' i t l iaddp r w i t ~ r m . o r r~~rrlt-d ,it , 1 1 7 ~n n r rleurlopn~cnIicd oil; rnoJe\ W ~ I tal rf.llrc c o l ~ r r n h the. cxyrcccion {lirxhcr genes nt n <ubserlut.nt rtacc. Tllc cr~cl rccult 15 n fi~llv drvt.lopcd orptmcrrl 111 \vhicl~"tlirr~-ir ,I p l , ~ c c v c y t l i ~ n c . fnr 2nd t.i~~~rl;thir~r: p l ; l ~ ~1C;t.t.. 1')'$(1). In kt% .' TI~crc.arc. rhrcr ~ ~ i > u t11'rr-~l;itorvgencq rhnt We linvc alreadv s r r n that palaco~rtology.firith I I pc 1~1vr rlitfcrrilt f~lr~c-trans r l ~ ~ r i n y h r y o ~ c n c x ~ c cm 2nd unique persprct~vr af gcolugcnl tlrne. wn pi1
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111

Fossil record and modes af evolution

121 I I I F I ~ ~ I ~ ~ l t z t tS
C\ICY.

39

the rl.iturc o cvnlunnnsn. ar~inlal K and its ct~r'lronmcnt,arc pl>cr;rllv ,~ttrihutcd III partic~r drqccrlr c)t' e\bi)li.ir?l: Itiic.yc5 to lrilcrocvolutivn. Althr~ugli ycnc-tlu ' ~ n dl w l n t 1.1r. thc be rrt.~hl~~hcd t h c I ~ i ~ t l of l i thro11qI1 ~ I I J I P itit: proccsse? bv wf~icliqpccler d1I1;'rciitiatc art' well and vv ~ cnrood. -l-li lr I:, 311 Iliipnrrant c : ~ kt;)r ~.IIDCOIISOIcit~dcrstond, it is a n l r : rompnr~ti\-clurcccnrly that , ,\nd .it p t t ~ c n both pllylctic c\,ulut~on t ,ind cnm- fill1 confim3nrior1 of how tliey 1 3 3 ~ ~ dct~1:111y npcr::111ri cvnlunon art. rccclvins nluclh .itScrltlrrn. ~ t c d h r o ~ ~ gtlnie h a $ bccn clrarly cccn Sirnprnn t li Of crliinl Ilnpolr.incc i- thc pcrcrption. glcnl~rcl :1',44, 1953). working ~rnainly \v?th vcrtcbratrr. ,-1111 tllc ~ C F I I rrc0rt1 .I~IIIIC'. l i , ~ tratc'r t cvoltlt~on 11nkc.d gcnctic- rhcorv a n d p,llncontnIoyc'al d ~ t a n in ,<'etaricrl tl~rnuqhtirnc 111 rolne rnrt;lrlccq rhnllgc tlliicly rnnnncs. a$ dlcl Cartrr (1954). I t carrlr tn h r 1 1 hccn vcnr r;rp~rl,hut 111 nt1ii.r caws t11c.r~Iln* ~ ~ i ~ d c r ~ f o nl d q t i t r ~ l *1l1.1t spcrtr.; could anus ,it t i r \.cn. l~trlr e v u l u i ~ c ~ r ~ mnditicatii311 nt all, cithcr by litienge spl~rting .~n. (5pc:-int~on) 7 I)\' rr.111dor~ r >rlovcr long prn.l~ld~t rilnc. I t wrmr to I r rhc o 7 ltlntron of thc ~vholc pnptilnrlm~ ( a ~ t n ~ c r l c ~ f l q ) . :Int L T ~ E I ~ ~ I D I I , I W C ] I ; I I I ~ L . 111 1 1 1 ~ 1 1 1~ I I C ~ . I I I ( - C ~ ~ Unt'ortur~atcTv. howcvcr, tht-rc wrrc \*c.ry few < nut prclirrd . ~ ; rcg~il;fr tl pnrc hist In f tr d i ~ d Instance\ whrrc fixst1 populations were ncttlnllv \ccn t 5on1cririic\ wc ran di.;ccn~ . ~ pIiI dI ~ I Cof CVO~ I to chow 3 pndtial ~ n o d i f i c n t i ~ n tlrrougli tirnc. ~ I I I I I111 thC t i w ~ i lrervrcl, n singlr. qtclcli, 1 1 1 ~ ~ cudlnclcccl, many of dw clarsic- c.l~c$ which bvcrr cup14 divrnlty , ~ n d Jcsccrl~l.~r~t\ qlllc!dv bt.ct>rnc pnsed t o rxhihit quch unbmkrn 3ent.c h a v c nor It17ti-d tor 1 i In vnnuilr 11.1h1t.rtr. \i~cIi adaptive 1 C stuocl up well to mt-~dCni ~ i l n l y s iS m p n i i .ind ~ ~. 'intions JTC' CIIIC t l t ' c 1 1 ~ 'I T I O S ~~ - \ * ~ d c nd t t r r ~ ~ athcn .;.~sgi~cd, I ) ~ n v i n ,that the saritv of ti)*~11 p t of a7 11ad : u n n r ~rCvt.:llcd h. p.il,;cntitc-rln~.A l t h c ~ z ~ l ~ .cp.lcclnlens rntcnntldlatt. in mo~plroro,q S).bct\~c<'ll ! rhc ~ I ~ T ~ I ~ I I F C ~ ~ ~ I C ~ CI irliainp~ttcrt ,till ~ V L = E I L I C C C \ F ~ V I I fnrni.; w a s due tn thc ~ n ~ ' o m p l e t r r nf ~ ~ I I C C ;\ ic ud, t1ir1~1gh w m r 111Ft,lIliCC t??gmieiit~rv,guidc the rock rccnrd. Tllerc i.;. hc~u,cvel-,i n a l t e r i ~ a t i v ~ rn : hr. counrn~ , F c l ~ o l ~ ~ t i uch.lncc. A l t h c ~ ~ tlic~ v1t.w. cl~unciatcdby E 1 J r t . d ~,111d GC-UI~CIIC171. llnn~ yl ~ 111 IT ( t i t h v pattrn,\ seen miy1it Ii.tr,e bc.t.11 prlhthat the rciclrri r n n b t takcn d t CIL-C V,IILTC, ,lncI t11v rd, tliey tvnulrl he utllikrlv trl 1-w iclut~rrncrl vcr? I A Cthat rhrrc arc so fi-w tmnsitioi>nl writ< ~ 1111 c71.1~rnl~tior1~1 0 1 :n d111n131~ 011 1 1 1 1r done know11 ha* .;rliiicth~nghndarr1cnt.11 tn tcll r l s ahnut Ilhccr mrioli\ I ~ C I I\\'id1 b r cvplnred 111 pcatcr P~ the nsturc. uf ni~crnevulunnr~arv praccssrs. 111 111 the hlldlw~np tcsr Ir 1, i t n p t m , ~ nrtr nclic . ~ t t Accnrtlitlg tn E l d r ~ d ~ c Crru~Jd, ,ir~d mart p , ~ l a ~ r ~ r i outret. l~rrwrvcr, that p a l . t c . r > r ~ t n l r ~ ~ i,~ t s ~ d c h n ~ ~ c h i n k i n s hnd bccn Jorrlinatccl by the idc,~ ~r r td l>t~rich w t ~ r k i ~ i y q i ~ i t r %Ire .ir ~iitl>rrtit I T I ~ G 11c-v. ~ Cthat evaIu t~oriaryc h n i ~ ? ;niuss incvi t a l ~ l yhdve bcPn .~ . I 11ariqr\ t l ~ a t,I scntbnclst ~.:ln prtxl~rcr'rhrouyll <In\\, nnd qtcadv. Thir 11-todrl of ~ J ~ u , E JCSCCIIC I ~:l\.r brertling ovrhr 3 r l k7 lf <cncr,ltiolls may nr~th~nodific~cion .1 qcclnccl to bc in t u n r \i,\.ith tlrc . ; i l a r y vc.lrr Yct this i d i n ( . ttmcqp,!n rrl.ly hc n~njrsricallvl~nfoldiilg.loiig-tcnr~ cbpcr,lhnll of tl~rh ;r\(.ntcd g~01uycdIFvhy no rrlrprc than J cl11g1:lc procrcw5 of rlntur~lw!cct~oila? onfinally ~nt.isn~~:c.d I ~ I I pl,~nc'Hr.~nrigr h l q 111 riiitld. w c i.m) pruIS by 13ann1i. W , Ii~ i ~ ~aftrr d .CI.IT<~C I )nrtv~rlirtnj t t , l I! 71) c x a ~ n j r ~tlic nzro ~ n a i n~ T ~ ~ C I C0 '~ I C J L ~ ~ S Tlrc ide'r of phyletic gradualism implies ,111 rvcn, c F1 OF m i ~ a h Ornicroevoltl tion :111drnacro~volution. .;lo\r,. rncthodlcal c h a n i y o f tlrc rvlrc>lc yeiic pool ~l: :l~rrocvrolut~ot~ rctcr$ ti) s n ~ ~ l l - \ c - . ~ l ch a n ~ e ~and hence of the ch:lr:~ctcr~ thc er~tirc o p t ~ l ~ t ~ o u . c of p hlrl species. and chpcrra!ly tlic rr.r11riorrr1,1tlc~1 T ~ t m l l ~ ~ r i lti1111 \ilolsld ~ I ~ V C J ~large niitnhcn, F ld VC "I Ihnr ~ p e r to c ~I L Y V nrtc. M a c r r ~ r v t ~ l u t ~ con , ueunlly rhe rntirt anccl;trdl. populatltrrl, snd noulri ~ ,I I rii klther hand, conccrn? C V ~ I L ~I I C I I I ;ll1tx-c the OSCIIT nvrrall as 3 large part of tllu ancc~tr.11spcciec , , , r lrvrl. 2 ~ 1 t hthc clrlpns o f - ~ n . ? ~ ~ O L I ~ S 1 ~ 1 J T ~r ~ 1 g(*.npapl~ical rdngc. It srcrns a rcast~t~ahlc conccpr. .td,tpr~vrr.iC)i.~ti(I I C T'IICSC t l I I~ ~ ~ ( ~ l ~ ~ t l and ~ x r v wlnc populntlon.5 may Ilnvc chalrgrtl ~ o ~ I-crtnidv I:\\v111 bc C O I I F ~ I ~ C Ti C>~turn. t In J U S ~ h i ~ x v . Yct, ar ~TC'IICILICIV r~utril.C C I I I ~ ~ I ~ L I t n I mi$ < C T ~ C Fof y : ~ J r d tcnnedia rc fortnc , ~ r c C I ~ O I I I ln F \t-cti in <trnt~cr.~phf~xl qcqzictice - \vh,ii n f c ~ ~ n r i i i a u v ccc i\ a Eequcncc n l sr,lblc f o s ~ popul;~tint~< ~l wparatcd by abrupt rnorpliolo~cal brcakr. T n xonte d c p c c t111s tnav indccd be due to ~n~ol-nplercner~ ~ t ~ bins 111 the fockil recurd, hut t t t Elclrcdf~c. r t 2nd
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