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Invertebrate Paleontology and Evolution CLARKSON

Invertebrate Paleontology and Evolution CLARKSON

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Published by: Douglas Carvalho on Feb 20, 2013
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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

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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

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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??,?

x

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

8

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
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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

~l cI.crlinicrit.ll.~lpapers .lncc< .I zrra 4 t of t l ~ c i for lot) [""VC 1 '1lldtOll Kcy rhc re> tlicy n l AF L I~.rcrptiona1 fauna<) Inclividu.7 I c h n o l n p Clnu~ifiratinno f trace cocrils Morphologic.xiv Contents 12.11 m d prcccnratlonal rlascitication 13chnvto~~r.i (e.1rlv1cc shot1ld tdkcn r ~ I ~ C S CC SP1cldo In1111 ( hcl p ~ illi1ilt'r ~~)r(o 1 w tl~..IHCC 11 C:olln .l&.0 1 h c t all1 tcxt for General index !'car 01 Cbnptc tlOll 11.~~ fbr all .trc.uc .1 chnptc .ind qyrnposiCl(Iclluolo~y) Ind~vldu.~c~ifirntioi~ Phylngcilcric cla5sific ~tiola Urcq of ~ c l l n o l r . truatl'icc arid synlposi.~l p c r r p~ (ichrtology) Systematic index fmnnk) 1 hc f i r liqhcd I 11c\3.ly ctlvironlllent Crratigrapl~y Fossil l>chav~our Fiil~lio~~pli~~ Rook<.itrd othcs rrferenccr (cxccptioiinl Rook<. g C.

p:~ri~q IIL-U. Hicks and Eve Darntlth for thclr i r l r t ~ c u l n ~ ~ r copy4r !)l. fro111 . r ~ a t ~ o t i S~~s. 1 1 1 ~ IIIJ tI~.I I mist111~O n a wet Novcmbcr day. tr>gctht.crcstcnt.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~. Iligly. i ~ w n 7.c~ Hrigps. in a dusv &rr . 1 will havc achlcvcd \\. .c1n:ulc \ubstanti=~l h a i l ~ c stu c To Keisi~l Moral) of Unirenit). H.arid thc Trncc Fo.rics.iL3rc.111 tlw \ u p ~ ~ o.111Iil prr~t-chcd n-it11 thc ti-~urtlj cditiotl. From Inn came the quggestlon.lnk .' l F c fin1 rllr~c. rcqxctivcly. * tllt~ L>cI. ~iccc<~imtc.~chcl\T!t~nd p v t cxctllcnt take11 iiito a ttlureutrl ill my 113tivr N C W C ~ S ~ ~ P .I extend 11. Collcgc. G~lwny ~ . tL30 Irlg. Pcrur had lrvud iirdtioris of y e d s J ~ O Thcv cvc~tctl hccihl~clLl~\i~. t lo~~ Hiftory Murcum.~vid Harper.and I wns given to itnderctat~dthat they .arpcr. :Irr oT ~ I \ ~ r > ~ ~ likc.9r . 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.~ild to thc laboratory for study.c-11 a t every stage. cditiclns o f tl~ictcxtbook wcru pub. ~ p t It rt che last o 1 wish to tl~3rlk 3150 nly editol-ial colleag~r<. 2nd tllen t hnng t h c ~ n tmok !1:1111~~1011~ 111~11tioll..rrlnrly i ~ . l r tl~c hrytitlil~gc>t'tlic p r o j ~ c r. 1 ~hills on a f r u h sutrlrncr mom]l L g t .I[ thrir first mcntion in with thc N a t u r ~ l appp. ~ r t i~lo<t F their s u g ~ c ~ t i o n ~ l (I AS .ltcr ivr Ic<'.crtinttr ~ h c 112s study.uld I hope that thcqc \\ill 1 3 . I ~ F i t ) brdcl . ~c~llcaquc~. g i q r r t t f t ~ l i j ) .11\\7\1:1y~.lhovC 311. coledit~ng pmofreadlrlg.:1~tt.lio have p e r 1 jc. my 0 1 ~ 1 1 C~lLtlltry .Ib l i F L ' a t O ~ { l r l ~ l 1002. a rrlcrc. I ~ ~ C . T l l ~ r ~ZIC) \puc~Sicirldux for thctn.HI Itclnl\ a i l ~ i c r n ~ i ~ r o o f fvlssil i~~vcstrbmres. ~ t .id Cecll~a i~ndermkcn me 11or >(.~1r.I g e n t deal. and I all1 still glad to cscapc to t l . and IL.2 Iwvc bccn revised to Jane PEo\vman. ~ l l o i half L)r latl Francis ~ r l d ~t rl:c i!l~~str.111 n l y fncnds.itl1 IJ. .lKL%.IS to IIOW I T y n c .\vl-ould rlcvcr hy .111 <t.o~. h ~ * ~ r h ~ ~ i ~ I Oorhcn Ilavc . (or 112 t ~ > \ bwn trmsfcrrcd kotrl C I ~ .~:~h< trut rid st:ll\~nrrfricnd Ct-cilia Tnylor. 1 Il. If ro.>l~rt: 11rlp1ng<ttnlcnt. a t ~ do f the rncthods avallablc for ~ t s r r > Irly 0 . 'rhcrc. yratuii~lthanks for her two beautiful 1-3.1vc 111:~lly Otllcrs. \c. I am hkcwi~cpntefiil to L)av~d I ! I V \ . I \t. I!:IIL~< I I ~ L ? " l:~tnily~v!icjIIAIT y1~-11 ~ L I L .hnt 1 srt orit 2 do.ii~i~ct~rr~l . lifc. u ~ (:ji yi-~. . ? .aii which has c n n t ~ n u e d ~11inE now. to sudrch for fossils. . r o rccord r r y hc. I IT ~ l i c I I Gc~icr~il ITILL-x. and r havu casc wort two g s n t arnniniai teq (probably 7'i!r111irrs nri t l ( \ .rrltton :~rld Ii.. has bccn urldertaken In corl.iqilq rec. 1 Iiope t h ~ t h i ~ to 111 will be o f value to any 5tudcnt wislul~zto cxplorc ~ltrl>ail.: a ~ii~gi. 1) p i i r l t i n ~ thc Pnrt Title pngss...lr loll? \vIotcr c\. that the new c c i ~ r ~ o n !.n: . 11 tlic.I I ~ O ~ IVI~-. Liz H.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.d so malty ad\:arlcCs both i r ~ alld prepaxing new coilfie ~r~aterials.rr :!I:. 1 I~a\.rt . Dick JctT'i.r.rrii\.c. tvlll rct.irl~Itl1r. Lorldon.r u. . L h t ~ . c ~ l ~ t i o n .\~<>L! 111i.~irlits tiunctioll as a course 11'1ve had the tline to cornpletc chic book bcforc t h t . 1 \\:isll to th.iiinF.uvc scdra\vn . <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. N . and tcr Kc\r ~ Y C V .'ruv~: o'l". over 50 yca1-r ago. This. r :o~:I! St.rtit~ri. . ~ ~occupation all posriblc asslstailcc. I ~ i~ I ~ o back I!I.lr t~~~\v. Gl13<l\Ll 11.. co iii~l wl~ich we l ~ a v c follo\vcd up.und l~clpfl~l rt ~uq. L ~ F .t . to cscape from the min. hut Nvnnan McLeod and Paul Taylor of that ~~lstltntlon 15 I oxvt .~tik\ duc t o all. ilthrr c11~1ptc.t .m ~ : ( ' ( ~ 1 ~ A l ~ t l t .> .~rrcl I?.

I hct of i l l l d g ~ c .at flcxlhillt): in di~playing the dat.i11~1itlform~rion n stratrgrmphic rangc.(. p ~ ~ l ~ c o h ~ o ~ tcrc.~ricty other cntcaln.Srrnt cint~barc rn.. or cxport tlleill illto other .I v. CD-KOM dnve. n. o r sclcct and sort record< h gt'ogr3pl11~ I I ~ C .AEC>VlSION diptal lmnpng cvcte~n.. Muscunl'r l ~ i g rc\ol~itioi~ l~ I'AI.o The~ ~maycshavc bren captured 1ivng thc Natural Hirtow ~ ~r ~pl~~. h40 x 480 colour ~llonitc>r. hard-part rrunrmlog. atld the data arid It~iagtg rcude 111 tlic Gvrtlt~.ontology ttcnc11111g. rncvphnlo~icaltcrnls.uk .llH15tury Mtr~rtirrlLc>nd(>n dnd Dr E.K. ~ ~O IiIl L'IL K O I I IC .lsc contact: ran. and thc uwr ran print records.v.m d database. Wltllin each rrf rhr Ir1. T a v n 1 1 3 been ~clcctrtl tlicir rclcv~t~ct. r a? A qtn~~dalorlc o prntluct.ltll .lilnpcr. I hy ' i- /" - 1 I For furtllrr ~ z l t r r n ~ ~ t ianrir c~rrlcr~ny c~ t rlet.)scil< (111 UU-KC_S:\I tn.yy.co. 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. o pnlacoorolnt. ~ t r ~ t l p n p h range.I carrrbincci Itnage I~bmn.Ire a 11u111berof labcllcd irnngcc and dldgrarns to allow rhc 11cv1 to bccolnc fnm~li.~pplrcat~o~rs. E ~ c h rcroi-(1 iotltnln.ljor gn>i1p rlicrc . c o r l t ~ i i i ~ n g ~ < I recnrdr fur 1000 k r y fowl gciicra. mtiillnum t> Mh R A M .~lls. ~ ~ for to PC o r MAC:. hc urcd as A I I a d ~ u n c t 111vfbrrrhratc to Prlfoeot~rth. lift' ! U linhit. ~f :I l r l c r q J .'llitrvt.CD-ROM T h c Nnt~ir.l. .1).frar~c~s~u~black~ci. Thi\ qwtctn ~ l l o w r u w r gc.~r ke!.p1e. 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~. For example. or hy .. Earth qcicncc tcnching \vorldwidc.N. users cnn \imply broww frol~i rccnrd t o rcrorcl. pull lip fnsrllc frortr a taxorlonlic indru.

P R ONE AT Concepts .

Charmwh. an assembla e o immature ammonites from the Lower Jurassic of Black Ven. original specimen in the James Mitchell Museum. England F [ h e r Sinemurion). hartmanni (Oppel]. by University Callege. knclw IVucdl I. Galway.~IIT . died prwenting sediment penetrofing the chambers. These specimens were pro&bly ca~strophicall~ buried.I(.111 gcc Amixeras cf.. since he soft parts must hare been in when + e . Po~nf~ng Roisin Moron. Ireland.

nrc dictinpirchccl 1s).n o record of terre5trinl . of IIC'W ~ c ) ~ + I \ > II ~ 'I~~~I C F I upened up to rlic WIIOIC nrr not cor.Ma pcnndq. ~ l l t r t. of pcrl1. .ls n n t 1ir1t11coiiip~rativclv ~ t e Scdimcnt.~ ~ p p O r t.~ atld phato~y~ltlrcric . d clinncc of ~itldentnild~tlg 11lrto1-y of-invertcbratu thc .tl . ntbl~l. l l{n>\ < \ I T T I ~ ~ O .J< lj: . \v. ~ t tllrlt.ir .~hl~.II~:.(l l ~ l l t )~i11111ot1 .II-C all I I I V L = ~ C C ~ ~ .Itrnosphcrc t11.1 .~c. !iir hcc111 ~o hnvc dom~rl. tllt*cr . ~ n c.rll\ ionnlng rhc pl~ncrg c\ ncqnircd hard rhells and wcrc cap~hlt.rlc. I ~ C nrdmal? T h c ~ t n t co f presrn.. The?? In. 51r1i 1 . ~ it~rc)nern r ~ d~-nucf~tot~cs o n l but 11 I I I I ~ L ' ~ . l t ~ o i ~ (. A:<.qttcd nr tlle~t. 13un11g rh~scritical period r l ~ cpriiicipnl I.I.~l t i d tl~crt* hissnn. nlid rhe qul?iect of thi\ book.nbllc>leof P!lanrrozo~c tltrle. S ~ I ~ \ I I i l ~ .irly ~ t ~ d c c d . thc 11c~ut2ll01) Mn.!~.hc tmic i ~Lvntlng ~nlicli lntcirqt ha< f Ilk.I \.~ct-clcratrd.cry 1rh111tlic t i n t 30".~rc i . or L-ynt1ol7acrcn. Tl~ert.1 of thctr~arc i t i .lr the rsjte Ph:rnt. Tt.~ted f itcrlth c . ho\vcvcs.ln.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.iiid g.ir dirt.cl~.rc'. I 111~11 <lowly ri>t. nnd evolut~nll Lr t h .r~u!rlcr~tnl ch.ition of fos~~lf v. rrttu. For nt tlliq sta.lteci rouild \ t r b m t c ~ . vc.IF k kll0~77.d. FIIJICS. itil~cplnl~ctqIn thc Sal. ~ hv l thc diccav~.r~.n. 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 term? js. a qtr~h. buns focof Ot' .~l~lv onpnntrd hciorc cl1t.~ ~ I I I \ I I I(<l > ~ c t c rIjI ~ ~ L .rl l~ir. 1 ~ . of tl~c'iuorgL~nisrnc I ~:c. i g ~h c Earth carrlr inti) C:nmbrlan Pcriod.lry rock depo~itcd 111 the senr driritlg the ( I I L ~ I +r\u..ingc. tllc rr\cib.lrc .lrhr. Nr\~enlieless.* ' to 2 .111 rllt.roll~dedthcy drvcrq~fiedand cxpa~ldcd.~ins.lc<.ly be very ablitldnt~t in (1 r'rollrt~r>rl. ir.~r Syktrnl onlv slllzrd. ':]I ~ t l t * r . pal.wnntnlo~?. I I T C ~ * F l~i~ltm-y..~rct. of thc cnrly Irvitly lbeaii<s o t ' t h i ~ l r t l ? llarl rr. and only becausc of this can t h c r r hc any 1.1)t ? rock? 341 It 1 .ictc. Thi5 \v.:!.~l rcprociuctioi~nnpr.Somt. ~ t u r t ~rnltll rnuch I.lnr\ ~ r r a t l y .1t war previ. .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 . l 11 :~rl~1~~gc.:. 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. hlly \vll~. mcks F U C ~ F L~cy%vnc-ker I I>~\ 7tbO A nr~dcvcn coirglo1nerntr5.yrrllh~lllt fi>rriijnqf r r ~ i .larno11 m s a i i d ~ t a ~ ~ e s . I r y r p~rt~c. I L 'h111rE~rntef that rvolvcd thro~~g!r succcssivc Ii~qtnncnl . .we 110 tbrcil ~nltnals rern.c\: rrnit~lc form< r. c ivlv:~~. a .i. and with it all lirnc<tt>r~e\.l$.117ci Inort part~c~iPnslv c.rt. prcwnl.I lionrlcnr~ng ~ dlrc of den pmllfcration oidifl?sent kiilds nf lllarlttc ii~\vcrsl I I L lr.'~tcd) w.nt~aI?II. h. rth..Lhrlnc~.~. r i c h l ~ c r i t a ~ c the fossil rcunmnq of thu invcstcof c r ~ * r i ~ ..~c vcn slozv. fur they #:I: jltf LK!grn i i ~ t o .iry ironcrorlc< iilny t~avc rich fo\sil l~ir.roroic... IIIIC y~lv.yc tlrcrc \\:IF A q ~ c i t . ~ c t e r ~ a b werc I [ .111y111~ 111 thc cnvirniitllcllt.h .i\.irl LI~vtjiil ~ t \ L I dint nr~imallife eventually .~r qnlilc. ~ ~ l ~ i t f0~111ed ~~r~vcrrchrnte~ 3 wurc c5tabl1rht. their ~ t l i d vI$ the dornail~of in\rcrtchr.I n ~lr of nur u~lrlerstandl~lg t l ~ c11istc)i-y o f lifr-.n. and thcy thrn 0s ~ ~ t i ~ ~ ~ ~ lquups Il. IhIL.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.I O F tlw ~ J .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 . S v d n c c d I l k .

Il.g.I Lmeat dcctroycd. t1lc. ~ c s r r ~ ~ b l . ~ . ~ 11ollou~FFBCL-.~F~r c .1.and 1% (7tir nnlx rccord o f anitnal sccry~talli7.I nlir~lr\pr.~ll\ of' c.d.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. M. .lrrn\v hntd onlv cnvirnnmcnrs.n~cly ? cig. .~gc+rlc.ll 4irll.~lltc>nlu rllc hard-<lirllcrl tlcmcnt< hc Icft.iricI ~ 0 1 1 1 kiio\vn rn Middle Ca~nPin.1twr C R C C .d . c. \ ~ t ~ d rnrk.r tvr-ll.a oripn.iftcr dlngcnctic pl1!. ~ n ~ l . spun yes. Moulds (oitcn inlcc.l.t.ilcitc prr.c.2.1litycrc ~ r i ~ n .coral<) and i~ ~ f t t ' r l~ ~ n ~ ' r t 3 l lt43z calc~trrllir~iig genetic or .l~nnitc n It'.in{[ in cxcluiu~tc tc.~cc. and rl11nn# .in. ~ ~ ~ n\ o ric. \vcrrirlq.\i\ hcr~cc l~ .~ ~ . nbradcd.lrltic i n ~ c r t r b r ~ l t c c g ~ n c s i s (clrcii~lial .icr 111 the rock ~ t t r -d c p o ~ ~ t r n n .Iir nf khclls tlf .irclv have thcrc h{-crl f i r z l i l r l ii~cludingdcc. ~tlqlit'stit>n~tL-l~ liiiitted. . \ ~ 1 t t i tlnu or$anli slirll<.~ncr lIy a ~ r and rnl1xt. i r ~itb r pn l. ~ C C C ~ I I ~C ! I E ~ J I ) L .rv he rcplaccd hv an cntirclv d~ffcrcntmln51n.Tlic\c t r ~ .rnnl ~ n intCt-tlalcurt:+cch\c)f thc f i > ~ ~ i l .11r f c\l<tr.IEI i)r tlic w$r-l-rnrlicd clr. ~ ~ \rqucucc< c o ~ ~ t a ~rluinrrmir came.~ r ~ r i i t i pornclcs. ~ ~ ~ l ict>rriccn ~ i illtdct W ~ I C tI1~1 ~ ich t I I l F t d t I C C ' .4ii.1. tllc. dcpcnciing Ilprln thc n-itliirr t o r nrt cvitllr~lst. jdl\-ti\h. I O o C 111 tlic l>ior.n ~ ~ d rci-n. I c ~ I I o ~ c ~ C ~ F 1s . ~ thc tlrnc .I:) t .111cl ivheri dl1 o l tllcqr.~ ~ r ~ rctlirncllt\ 1vit11 a high (Fig.~l i .\c ~ r rrninc.11 I-hnilpei) t3k11lg foiii~d ttl ccrr. r ~ l ~ of totnc 01 . ~mnllshrimp-like crr.:rrilq chcn. hcd~ cullrditung ~l111c .111y .I prr. I ~ C I I I I I I and she11 was ~r I IivirtG orghrnsrnk (c. I) ttlc o n g ~ n .cl dii~i-chtlFcd ~nlnt.cd. Even (hit k-~hcI1c.I T ] \ V ~ I I L -t~l r ~ . w ~ r n c .e d~.left i n thc caviry from \1. of t h r ~ ncrivltv 111 tlrc f i l r r l l o(tr.ricnl c n ~ ~ d i t i o r ~ < die wriirnciit . ~. {I) iron. whctcas cxcerhinct.ulitri:~l< of qtiitc ~ i l ~ k n r ) ~ v n nFtii1itic. t l i e ~ c ( arc clcccribcrl in ccctirrn and thC h i v l .nic othcr than rcc. with rcGrct~cc rt> h r ~ ~ l ~ i c ~ p r > c l \.~vc y T h c olclc<t qurh f ~ i i j I~ . I . :lrc .halamcc i r uilrtnblc.l~-couc i i n n n ~ l runJli.i'r For il~sc. . r What \te.~ltvrril r cxprctt-d trilc~bitc~ ~ ' ~ . ica floor is !lot al\vny+a rcgroll r d ct-mt~r~~tous scdirnetlt dcpour~on. I ~ hc.~TIO. ar.L J L ~ I ~ C ~ 1iii!i~101vi1(C'linpter 11). tn. ~ Llse I'l-cc~rrthnan n=c.IIIVof tlir ~ ) ~ ' ~ ~ .I r l ~ h c w ~ t~ rhcll\ nlnv h c sct. > l ~ l l 1 1 ~11ketviw lll~lll2lTl~rill~.d. I I I L ~ ~ ~ 6 1 d: ~ I nle11t. d ~ d Trot calc. T h r \cdirncrrt 1i1111 bc in r t w l n ~ cal cquil~briuni(>lily.~ppeiiq i l f o ~ ~ i prc\r-rvrd In iroii<tnnc\. r.C . c ~ I C J ~ S L l c f i ~ rt~ r \e~lliilri~t r ..r. I f L d 1 ( .I\V:IY inlr~plctclr.li.hrt>wn tc>g~-thcr c!~eiilical 111~11tr1t1 t l ~ c t. ~preqeiit 111 . 'l'llr.7. rhcis b i u l o p li-r~~-rit l l c ~ r 1. -+-.1t1nti i1 M >TI lifc b c f n r c t l ~ c C:niiil~ri. .Ire ~niitlar'windo~\~'i' crtlicr luz-rlc m rhc s i l rcdimcnt(: r11ay clis\olvc . I ~ ~ T .~ll~d the ord 111Ri' JIICICIII I I ~ L .c fo~<ik. (h) Cnlcarcnn~ 5kclcton~ prcqcnbcd in rrlorrn SAIIL?Y or ~vnul(l~ i t l l r r \ \ ' ~L~ tc . ~ pr~-rcn. ~ rllc.q. r dctsil. Any ~ 1 1 ~ 1011 ~ t h r SC.ll altcr-.iinctl in 1 1 1 ~ .nun.11~-rc In nr(ditlot~ tn hic nonnnlly onc nf tlitrr proccwcc tllc fowrl~I I I .ind tlic ~~i-o\\~tli 11cw ~ n i i i t .~inr.I. fowl LOI cncloccs . dnm. chow rridy b~ V C ~~O C \ S ( ~ r l c ~ J C ' ~ ~ ~ f C ~~ J.I fc\V IC 11 ~ ~ V ~ rock i c crnckcd ilpcil.I bivalve ur br. ihc. casts) T h r tils\ll rrrr~rtF . .prcrciit in t h t rnck nlny tht* rturly of cucli ~-rlould\ yc< If llavc tlccrl ~ T J I I F ~ O I . \\.311cli lr slrcll.11cnrcous c-(>ntr. i n .citIlcr ~ l i c h 'xvindow. rllanv a p p a r t n t l ~ corznliu.c 0f-~2~].~~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. rn of nftcr its indurntion.rt.111d hnvc lcfr littlc with rclntivcly littlc altcrntioei. thc c v o l u t i ~ n 1.scrvc hcst.liirl if thc wdinlcnt is tint c. C " I ~ C . ~ ~ C O I511~ll~ 1 ~ 1 ~ F W I > \vltkl~ll.ll\ .ctc~nc. ~ n c o C01trinhi.r inuch ~ h o u t U -CV U h ~ t dpart\ uf t t w ~ l.cict7clrciilii ~ i f t u . co~lrllttorif ~ttl ~ c . ~ ~ d i l ~ l c ~ i t re for core t f prcrmn.c~d d c ~ t r t y c d .c.111 c ~ r i ~ i n .'i \ (authigencsis) ns \vc11 d'i C C I T I C H ~ .~c i ~ .c n r b o n . Sliullc zhnr nrc nriqin. rn. tllr nature .I hchvccn the p.~Ttcrt l ~ crcdilncnt 11. . f i w ~ m t c I J I T ~ rhurc irlrni.I . ~ ] rerlurcltr. JF h. c11cIlq ~ 1 1 d t). ~ t ' d i ~ l ~ r n t .d \I it11 eleinellcc nf n t l ~ c r d fizln~r. kno\vri JF tfipllonnrny. ( v > ~ p .i gu~Ic.Ire the o l l k tracc of n rllvcrrc Kilt~iid \vhirh dlngciic~i< d i ~ o l v c d core. ~ 1.c k i .nJI-. ~ i -fti i l r ~ itrncc inqctlc) nsc pre~cn. I n n l w t s:wc only t l ~ c hlc tn )nF~..rilkl plivsir.~ T C~ ' ~CC ~ II ccwnc I ~ Tnr clr uilty t l ~ c l ~ c1 0 b r f0~~1117td.nt.in~yort. tliir qpnrr may citlicr hc lcfi rrr~pr).~tlilnll\ pod. i ~ r c r ipr~-\c. .m$inor1n d A < c d ~ t r ~ coftcii concirt\ (h'roriipui~cntc nt cjcnvcd ran scc In t h r rock< i c t l ~ c s c h r ~ a n. \u t F ~ ~t 1% porrit Thr qtuilv o f tlir procrcscs I e b ~ d r ~ l shrriliznt~nn nhniidnt~tly.n~t-il pl. .in i-(ir-k5 holn Rrrticli pactlon of tllc rock ilithification).ib ~ Therr .d tlicrc Figure I tr f~ l railer o f <ot't-h~>~l~r. old.c .~ficr.c ~ i lv ~ i t c r ~ ~ . nnlv r.$ ~ t d 3 1 n n ~ ot' C .I rrnl. rhc cofi-llodrrd r l c r ~ ~ in n rllc EILIIJJ.rt>rii.n at liardcned nr during n~catlici-i~1g t l ~ crock lorig ol' gc@loqc.i\ . h tilrn intetnal mould of thc Lwil shcll.~d 'irdlonce Ilviiig.111j1n.ctn~mnf the o r ~ ~ t i i < i that \ ~ ~ c r cfrntn v n r i i .or Lwcomc f>und.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.ivc. Thiu hcnrh upon ~ t c l ~ r t .from 11n~r.~rltrn.~tiur~ or] i n s s u d ~ n ~ r ~ ~clicr~~iial tlir t .~contc\lr*.nal moulds art. ti111-r of br tr..~in ~nally of ~ i ~~l turer .ltn rc~cbtypes 11. t i n ~ rtci~li~-. tjllcd w r h ~tclvnicnt. patchy and o f thc eurc. In rx-c cirruitistnncr< the r-orc nr t l ~ c ti both.In thC 1. ~ ~c r c t c dn cc. ra1plrll'rlirl.~tioi~c t:tkcn pl.it. t11c i ~ l c o n ~ p l r tU\u.t~c>uph~ dctail~ r.~IIF 13(1n11.~ving r g ' a ~ ~ i s ~ r ~ \ . nr ut15 r e d i r n r r ~ t .y rnay not br p r c ~ r ~ \ ' r 111 C . <hellpre whizli . ~~iivolvr \ ciF .1not21~r thc n d c q n n q of thc fnwrl tr.

tIc~.rcld..11d to hc perminetalized. 5.~\ ~ i\~lrl~:. witl?in u ~ h t c h thr.~tur~*q.1rc 0ft~11prcrrr:!l Ilkc rhrr.tlong \v~th 111. (g) rim of i e I ~ r ~ s e r wIn lmud w ~ t h r orrginol shell moterral leached away.~ o fctllr: few[ and. na r ~ ~ ~ ~ ~ o g o nrerytnILzd to m i c i t e rhlrh dertmyr the fine ~twcture. u ~rlud. Tuch siliccou< corvq rctnin cvccllcrit tc. 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~. prc\crv1ng the ~ntcrnnl ~ ~ l n r p h n l o gof t11c sIlull. 111. Corcs ninv ~ ..icr~ tlw ~ h c I l i~nprryintcdtv~i-I1 111 .I\'.Occurrence of inyedebrote !ossils in Phanerozoic rocks recrystallrzea c a l c l l ~ 5 " .I[i l <t l l ~ I -\!l~ijrcoc rht +hell 1s replacement. The zii5idc of 3 qt. alk.lrcm I ~ I I ~ c ~ I1i-I ~t~. 111d1vidu. n.. h.\ of p ~ r i t c .IF rhcll ovcr tllc w r hcc of d fosil ns a rvsult of wrnc crrr~lplcus~irKitc GI 1~~1.~liiic tlic I~tr~cqtor~ct h hydrochloric .51llcn could prccipitntc nr a gel. CUP F I possible rocerses OF foss~lrzat~on a bivalve shell ( a ) original shell. I fonttane re. ITOM t h of ~ C Cuid. l 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 . original ~ I C I C &dl retainJsurroundangc dim now ~ t e . srrlubll~t\.. ~ n ~ r o r h ~ c 1 1 . (dl i .1iiJ 1 i \ ~ c 1 ~ 1 r r 1 1 ~ ~ d ~ i i o f cnlcalc. I* IIIIL tilt.ll-tollt~\ IN. lew~n an external ond on ~nternol mould. 1 ~ 1 1 1 rcleawd rIcr. SL~ITICtl>rn ~ o C S t U I S I ~ of 311 of ~ ~ C / ' . w h l c l ~ c c t c t e d m c c .llt. ~ t ~ : l r i i t . I : -1rrltcrl dririiw.:ri \nIpl~~de.IICIIC very t & ~ n . ! growrng round the shell and Inside I~f tie orlglnol cavity was empty).P C core 0 SIIICU. wlh p k h e s of pyrite rn ploces.~rlcrl TIICY 111 o ' it~~it cvprefsiot~c)f the ~ u r .tl7' ~ ~ i i n ~ r . (f) tectonrc d~stortion o shell preserved in mudstone.~~tr-l-ial ~ICC.~.ryirig undcr dl6icrcnt c a n d ~ t ~ o rwould trap juqt i\ cuch an intern21 ~mcrocnvironrnc~it. buried In mud [leh) or carbonate (right). 11 r~-. . ~ c ~ d r r ~ v e throng11 thc rock nild precipltatc ill l L\~IICTCVC~ r l l t pI I i q lowcr. 1 (el u s~lrco growing o n the ovtslde 01 the shell.r. svrround~ng rnwdsl~ne a ore I o coicoreous concret~on . i3n chc ntllcr I l a ~ ~ r~licn y d.of thrrc d11iir11~1011$.lqer ~rlarkctil!.(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.>locernentof core ond port of shell B q?.~ urchrn cieu. (c) shdl *rig~. nnd thc s~hca G O s c l c .ll In w \II.111v y v n g c c .1~ r p l a c ~ r C. rhc.111 i ~ n ~ l t ~ r alc~tc i = d clicll T h r ~ linppencd 11nc '1~ IlrrLTli 9 1 \tr\ 5 rllc ulica 1ncre.

111 ~ l i cf n r ~ qtttidmt~n poi~ndswhich IIIJV b r drrccrcd gcnr.1 p.11 <kc-lctc~ii. ~ form .IS t:lr . prncr.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 .lrr A rclar~vclv ~ I T I L .Inn) I3c r ~ p l a ~ c d ovcrgrolvn bv .11 lriirlr tlic ~l . ~lt~ n ~ o g.~te pnlacon mlrrh? : biostratigraphy.> trcr t n ~ vrciii tbrcc :it> chemical fossils.lrtx nf thr $coEogir.~l.1)..I pnrt1rul.il h.Itlatoirlv.rivlty dirrctlv gn)~ip111gs.111~ o \ p l ~ ~ ~\licll.shorrlv .~l . \ ~ l + t l 1 .murt io drr (Cl~aptcrs ~ n t2).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.c brrn yrt-\c.c\tc. . ~ ~ C C I S C I Y r c p l ~ c ~ ~ t c d . tflc f G p c C t i v ~o f p .~.i 1111111oiivearq.ll r t ~ .<unrr of i i . nrganit coinor1~111. ~ t oi1 tl cxtcrrl. br. . rrlany n ~ i l l m unf.iltirnc. l t c tlir tract'tlg of t.INI: i ol~ * Forr11.~l t t ous o r silirCouc i i ~ n ~ c c -hrn~c. p \ ~ c ~ \ p h : r t i ? .inccc\oft-brsciicd ursanlsrlls thc . \vE~vrc . ~ l t111l~.lr~t~--d~clltdthc oiilv q ~ i d c thc fi>riilcr prr.lrc o i t r i ~f i ~ .I< pclrsihl~. ~ ' appczlda~e~ 1r)t:lct cmbwchogy. t ~tllc .iftcr ~ r sd r it11 .lt h.. ~ lti i . . .I.~crtohrgv t hr-r~.bgniiict the b a c k ~ r o u i ~of gcoln~ic. l ~ i C.1ppI1carioil of n~vcrtrl-rr.lnin it'lvurtcl3r. l of the patrcrnc of cvolurt'oi~ rhnt .wr\*cd nlco.IF th. ~ r ~ .rc li.~nullo~iites.1n n ~ l ~ c r . s . tr. 1 . ln il v.11 ~c unlts.r h cn Chnptrr 1'.~rt~rular .rcf.aii cirraugc-din n 11wr~rcll:alcyqtcm 111 xvhirh t h r ~ rr.itcti topic^ (Fig.lvc \cvcral ~ zontls 11\+111g .I Iiirr. ~ l t t k t C 1 ~ C ~ C ' ~ I C . I ~ Ccipliiit. 1 ~hc JIFTIIEsc.. rnli bL. i i ~ ~ r bc ilnrncd .r.~rtruildtlic f o ~ r ~ lprrdictione of evolutionary ccirticc to hc t.". rl n l ~ slit-I1 a\ rill?. ~ ~ ~ l i ~ .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~.nl\.l. v j l i y i ~ .d r .rl rciilaln\ u i r n m c part.>r ~ i r ~ d c r p ~ i h r r l (:1 ) body fossils. t ~:I\c l ~rlqvrp 1 .H vcr ~ ~ ih lI I t! ~jrrd~.tcn.ri.~nc l a% i \t nftcr t h c dcnrli nt'rlic.iofinf arc.lncccclit< \lcn\. ot' \ v l i i ~ ! i IlavC .il.r d r l ~ r . 2 ~ 111 \.t ~ LOT thc.6 Principles of palaeontology tr~lolxrt.~~.i!/v~ l~ subdivi~inno f qr'olop. 67s r.lior gcc3ulapcal t i ~ n eurlttr {prnndc). yrc-king o u t r~ltcrn.llc. pcrinlr7 d u ~ l d c r C ' C ' T ~ .lnd ..I ht'nrq lip 1i7tn I~~vcn-tchr~~tc.r~cii \ V A ~ C T~ t l r d i the.~lct) rci.ppl-cwn-cd Ir f i ) ~ < i I h . geolo9cal cnhuil~llh a i been c I I ~ ~ F C J C ~ . curnpnrnrivc . art.n~cd b!. ~liicc-l~ving urganlsm: (2) trace fossils.~ bral.cry r:ltc clrcilni*. chit-li ot p~l non~i. At t l ~ r l~ Ileart OF in\-crtcbntc pnl:~cniitc>Io~~ ct.il t\'pcbut {species) 11.~rsoc~. hut much largrr fu<\~lq 3 d illay Ilc hc cccn I l r evnln~ttonnry pcrspcctn0c prt. 1i~unlly ~ h c l of r k c l c t n ~ 0 1.t. . 111 rhr rtjck~.~l ititcrrc1. u ~ o j ~ ~ taxa.lls 11. ~nnd < II~ I ~ c 11111~t~. SiL. I < i t iq t ~ i r l ~ n Enrtl~\cit. tliilet .ll-thv nf ~ n ~ n l l t h ~ it< w n t c c t . Uslrly the qcqucllrc o f fr~\nl K ~ l i i l .lr.irr1\.I r-orr.1tuml p(111pi11p.i thin bvdicd anirrlals 111 .2nd t11c. T ~ I C W i l '~~ I proL. ir.< .~11jy1..!-l-lurcm lirr the oldcct 2nd 111mi gcncsal . a n d .i > t I. :F~C'CI.llurrcrws or uthvr cvldci~c-c i t h c .11.volvcil I L I Y ~ varicd through t ~ ~ r ~ c .C ~ T T I C ~ I I ~t~hLc-rF.~llit% of ordv .I-TU~C' blolov.rnq ~ i i c . LC.~liter. gi-11ct1t-s. ~ pafiiculnr rc. Ilnvc b c c t ~FLYL-J p.~\ ~ l l r fos~115 lin\. i~ furthcr d~vidcdnltc3 .rt pclints to this rcl.1lI'ri~i~.I bt'ologt'c~l u h l c r t I x ~ tlilc rh. ~ n d rl~csc prrwidr J fiatricwurk t.cilcc nf'.lt~'i~ C FOII cllcnllc.lti~i1F \ A ~ I I ~ I ~~~l i tCvorrlcrrd and 11.~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. lof Uuc difTcrriit h n d s of (05~11q 11. whir-h . ~ i iId~ concretiuns: c ~ l c . mnrt .1111111.I LprCsrw.I r ~ ~ ~ r n loe r p a ~ ~ l u c n l tt.rnt'm.f~ciiiir.>t Irvir~g.irtunllv 11:lt.~ltaxonomy: thc clarrlfic~tionof f I > > ~ i 1 v . t i~ dnvumentcd qcqucncc r ~h5cll.. I I ~ c ~ l ~ Canil-ri. 211 hcnring 0 1 1 the othcrc . r c l l c ~nf b~clgcil~c 47f t .111rI rnodcrn stnicturr5 1t1 rcmnrknldr* ~ % ~ t .1\'r 113A a geolcrq.c<c*. r(7vks.1~ i ~ r ~ ~ r n n ~ n(31 ~ t : or c ct s h c c ~ p1lo~pli. ntlwr ~vordkthc actual s w i i ~ l i n ~ g c o l n g ~ record in tcr1in of rr. r.V Ih IC I ~ I C t h d t ' v ~ r ~ lof i~lt'tal ~ ~ r ot ~ ty ".:.I rccvrd are nlorc.I~) * I E n p r c ~ ~ ' r ~ ~ ~i~ t 'phosphatizatio~l.ir.rrcd wrth Il.rctlot~~ fLs111 . W1tl1111 tllr ~ l t ~ r l ~ .ivc diftcrrnt <tr. ~ I I FC I I I I ~ I ~ I C I o111\. to nnilnnE~. ~ c n r ~ tI \r ~ l r ~ ~ ~ 5 t i l ~ ~ JS d of m. urcumed. othcrs v c y slowly (CEI. ~ n dt ~ n p ~ i l .i\ thc uiirqur pcr~ .rst~rcsd.hiopcd< ~ r c LIIO\I Froill ~nntc.rt~"ll I qpccicq corltillcd to iliir parr of thc ~ucccc5ionnnlr and rcprcscrltirlg tlir timc ~ v l i c ~ l spccicq \V. tllc Altcriintivcly J F E I I > T ~ ~ .O ~ I ~ I I but I IC ~ T L ~ I I I F I T C1 n o ~ 1 ~ ~ track<.~ils.i . T h c ratcr at which at'li~11.1nil4 thr \lir. Tlrr \\ 11rrlc b ~ < fur tl115 li~rtoncnl t r It l 111 t)ca cSlroiinlop .ncc \vhicl. ~ t ito ~ ~ l ~ l i ~ p s rn:~declcnr.II~V thin r>r~.ahl\.~llyi ) pli tit~ ~ ~ r t thc csrcmnl iO17ii r r f t h C I.I~~L-T A 2).j .~ni-l1711n.rl .1vc.ycan mny rht. closclv wl~divlded tf1. C(rin~~tirnc+~ c it? ot' n r i a n i r u d (71' FOTIIICI.c mcnt. I ~ I L hllit~yof tlic i i i ~ ~ r i ot' r . Sornc nt^tliece rvnlvcrE rapirll~'.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 . ~ r ~ d CCrtJlIl p. M .lttrtc Tl~rrctnnin sLitc.

y. ~ 'j ~+Rc~ d ~ C > Z C ~C ~ J I I ~ I C I Iknon. sils.ln I pur. lncilts.r~a5 synecology.~rtli'~ L: rot'it~oil(Chapter 5 ) . Thic topic ha5 proved valuable botli in ~ -111 ~ l rlln+ tEcvrlapcd pnrtict~I.s in tenns of thcir original f i ~ ~ l c t i o n .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.c l : ~ r ~ i ~ g t l l ~ ~ ~. therc arc 7111~ ~-TI.c binlc. ~ l w .both as ~ ~ i d ~ v i d n a l s {autccology) nrld i11 the faunal co~nmuuitics In r.iI~urn I ? The wrlovs sukdiriplines of palaeontology l Y 5 h .1\ tlct. infornumorphology. r I r r ~ impnrt.~.~i\l>l[llr rlt' peochronolwp.i r ~ ~ti l ~ c s . 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. Ichnology i? thc sti~dy tract. tl~: c~~:c.lI ~ nppl~catinrl< p. howcvcr.h m l c ti1rc. is concerned 1 ~ 1 t h rclatioi~ship~ fosd thr of aniln~lsto therr r~ivironmt. rI C IC ~~.111d ti I~avc 1111~ 1h.n f l ~ ~ i a lprc%crvc.-I\frr..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 .l. sfracc the c ~ r l v i ~ r ~ d c r ~ t n n d ithe bcIrai4c)ur of t l ~ canilnals th. .tron r i pcnnds o f ~ n i l l i o l ~ ~ c m ) r.ilrl~ou:l+<trat~qnphl.~pltic. . .~ol gcochronotnetry'. iiiorc h i n l o d ~ ~ ldh: thc t s n c k ~ . J s r ~ ~ a lhtwt.1~ Il~. 1 s Sincu thc <oft pans of furfil anininlr are not nnrE.~thcr oty th.:~ n l v ~ i ~ ~lr.~riyr. rcqtnutcd in scope though thcrc ci-dcavours may ~ I I I .*FIIII ilor~zoi~~. I.the basrc nE tlic pnnlnr).~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 . L1v counting daily Railits car] br ~ i ~ ~ d e r s t o o d . Palaeoccologv.~locontnlng nccrsqarily be.:.nlayc. which dcal with the Inter: r ~ ! n l1. 1his Ir o i ~ l y pnnihlc a t . pur15 . surface trails.~-!t.d.~tn.~~lt.~IT-~LIcJI ~ t ~ l h i 3 hmnng ou wlint liiay he ~~1. lv but only thelr hard FIICUS.lt iig I I .~rr.COGaf of 11 lr.~rl\. Studicc ill functional :tslirfE~ nrlp 111 cxtil-rct corals and hivalvcr. \ ~ ~ v rand for p r ~ c t ~ c a l Iio r.1tivcIyC Wm-~ys n wllich thcir biology and Ilft d3 ~ ~ i ~rrr.

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

new.~trrniiral~ C A ~ L I ofC tlit' sptcic\ ns sp. the c ~ ' . .. t l i ~ q follnwing thc gcncnu rlntnc. 1 1 1 ~ to.lt~orl ~ \r. \\. In 1917 F.lronotnic t ~ o m c ~ ~ c l a ~ i i rn~lthnr'l.~rc*nthcwr. I~ro.lrAtc.l.~tinn<>Fa n. rlnr~t &mi\ a$ thc nclv qpccic.Silurian hrach~apodr thc Girvan clrcrricr. r.lrcd for cnch ncni I onc pnrnci11. rc? h r 1dr.~ri. . 1.coin: ~ p.ltld the dLltco f pllL7l1~. ~ C O I I ~ O ) O C Jtll'l7l)~tlll~ dl5tll1If ~ F I t0 Hv EX. Srotl:find.I %4nrrt>vrr.~rt'fi\I~ Y J ~ T I I I I .111d fc\v ~ilrqultr.~nnll tlicrc i q ccntliry hr~chiopods\vcrc poorlv kiinnn . fnrn~al ~ o r n e i i ~ I . ~ ~ h C \pccic.i.r j l l l r i ! l t i l n t i t r (Schlothcin~j. ~ In rnrc nn cs~'iting genuc.r l onsidrr. . Hr.r.l~crcbvt-nc11 cpcclrs is drfined hy n ~ ~ klntl that would hc c ~ p c c t c di r l a lclcnl v.. nnmcd by 0 . I C C L ~ ln t1w ~ v ~ r h 11. is. I V h a p hc clqht such potjps.il cntei-I.~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 ~. ~ n d de~crihcdonc o f ttlc mnvt abunhe Lutinii~~ ~ln~iri~ pL tihop. I C Solllr of tllt'w ~ C J I I hiluro-13cvom3n genu< of thc Cl.b l:herr~i~tn. : ~ I I F ~ rhe qpc.pcClc. Fiqchcr dc W d l d l ~ ~P i~~O~~l C I S C ~ t FEII hrai111~1~0ds.rcctcd ~vhcrrJ 11nl01ypc h n ~ n ~ 'hr ~ C ~ I I I ~ I C > ~ I c>i<prcirc 111 pnl. 'To apprcciatc the mctliod Ict ur con~idcr of the follom-ingh~storicaltalc.~clunpctd a ~ a n a ~ slrlulnr ayc.tdv i ~urc l 11.lrc irlrtrx ( L i t ~ n a c u177s) (rcc hclow thc refereilcc spccimcn and fully 11luc. 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. 1.R. b.~tic~n. 111 fulF t. ~ c l c r t c dnr l ~ <.1i)tn13l Art= p r e w ~ c d .~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~~.iy hc ~ C W snd if rcl a fill1 rcchnical drqunption .~llyerected. von Srl~lothcinl fint srudicrl . nov. all cars. I-. lit>il . In the early ninceccnth LII In rhv crcctmn nf h y h c r t . pzcviouqlv . F . a s~.1t e ~ .d r 1 clcarly rll<tlnct h 1 1 o m nnothrr. rlir tndi<)IT n so t r:du.jnoiic. 1101. thc o n p i ~ n nutl~or's in.~ztcrl cornp. 3114 much 11rr. .11-scand < i i i a l l . rherc.ih nrk tit‘ gt. taxonomst< Knllu\v the biolcrpc-nl fi'ftrrli of dcscri\>cd spccics.I 11cwYv iliccoucrcd Ounn. bc hccn Inst. ~ ~ I C I T I nritlior. ~ r i l n n r ? ~ ~ c rntecr With~n anv ~ntrrbrecdint: pnp~il. L L I I J ~ \vIlcthcr n poplil.lr .1~ should frc pzihli\l~cdin a pdncontoaoa pt. S n t r ~ c i thc n lit.~~~i~-phologc. rcq-cc.i *. prcp. W h c n E. . Thew arc v a n o w othcr kind\ c..ir ~ ~ t w i p heen ~ I . 01 l lll\t~ncr~c>ir 1.~ E I ~ ~ > J ? c ' ~ ~TIlT \ ~ '. nnrl s~itllcienrlyrlifl2refit to 7nc rcgardcd a 6 A <pccir. it \ I ~ J ~ dc a o d~ l rr s w hut :trlhc~l I~r.5 T e ~ ~ ~ b r ..Divisions of invertebrate poloeonto!ogy 9 4omenclature and identification of fossil species conlparing the cpccicc point by point.rl~ennto.11ly D e i ~ o n i m ' I-rll~hed.Eneotype mnt.4iid leopard.~llcd paratypes. however. a< In rnosr s17c~1rs h n j : rcdt-ccnhed In fullcr and 1ttorc 11pIlnnr: spec1ilreil<. rt. Oiie of thcsc w. u c h srud~cqnLn7 be vcrv vynificant. s.rwon nf thiq b\.ltl0tl 31.ly of trmchinpods.tnhlichir~gthnt 1 I~IILI P C C I C FI \ rlifl>rtwt irnrn . hnt11 g r o g ~ h i r : ~ ltlic living Trrrhmrrrln. t h ~ it bccdinc cor111ri i ~ E ) ~ ~ . Miillcr in 1 c ~rratlgaphtcvan.70. C)nc of ~ ~ C F had ~ ~ ~ .ihlv lew. A i~tw gtnur wtll ht' d c f ~ p ~ t as d c gcn.tratcd.illy .]I1 tllc an.~cednIml-r<rciitircly on ro-dntc tcmis whcn 110 type spccirncn ha. t .tl1l cl:~tn1 ~q ~.i n n g o r sprrlrs m3y prnvt to bc ~dcntical u~it11 nlrc. niid n ncw suhgccics .I ~ P..r..\ i r ~th? CIUIIJ ncc: tllc generic nnA specific (or trivial) ndint's. fix c x . rcctly desi~ma tcd Chonf8re.ncd upnn type specimens. fnmlr From North Gcmiany in 1870 11c it~llvdncutncntccl I < thc rpcrlrs. illu5trationc rnlict hc.i c c u ~ ~ i n n ) .p.ll for c.< 14t'/. C)t the.I< FCP. In r r c ~ ~ p i ~that come of thr shclfr wcrr b r ~ F ed chlopod<.I rclnted qpccic~. I:t. e.~rliirl~ of Iogic~Ejournal rlr ~ ~ i o r ~ o y s n p l i . dcrcnptintl 15 This i nniiicr I: rrltri.\ \ ' l l i < I ~ 1111lqt th~11 Cowpcr Rccd.:~cc r.itlti-it~rrlt~rr:.~tcd. 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.ln:+tlon.lElt-r r d l r t h r r r iiiav 11t.7wd05 r~~cwpllolugi-~~l v.r.I ~ h n t wnq rcprndtictivr-Iy tsolntrd o r not. o r rhow only rnlnnr \.lmn\lc dctnil 111. pnrdl~r tbrni.ari.tn Stmphomcnat. ~ of any ~pccic<. T U I ~ lu~r(.~kcrkept 111 n InuGcmm o r rc~catch~nctitutc.~nr n w t h i i l r h fame ~ ~ C C O tC c r .o nntF I . nhr~~~ A ' l l ~ 1 1d ~ ~ ..!i. 111 d i e m c . to tlic Jumertrt. fp-nups nrc naiiic i q quoted In p. iv IIC IIJ\ ti1 iep.lcu5. ~ k 1 1 C ~ ~ ~ ~ cxn~nple. t k c i ~w o r k i ~ l gnn Ordn\+Fc~an I S~ ! :~nd of ~dclmtirrl. tl1c holotype.~l h \ 1 1 ~out l r l t i l gmtipq of n~clrphnlop~c~ljY ntw penus tior thiq qpccic~. i d' ~i Ei I~ Vl ~' . which arc n1wav.~ ~ I ~ L V nrrlculnr cct o f C ~ ~ I S .lt.c uf n I of ~ I .I< Ilac provcd e<pec~. m.1y bc .i thc spc.1111511t(1 1 A c. r n ~ ~ c h nlorc waq knolvn dbout . I ~ ~ . nf ~ C Thr ncur qpccies must hc narncd and allocatccl t o 1 ~ 1has to be the maill giidc 111 c. I I I ~111 thecc tnllrt bc 177h. 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 .~.nnly rlie hard t-ccrl dcngnntcd.lentrmln=.C glVrl1 R ~ T t C l ~ U ~ u a l l yniic of thew. dirtinct gcncra had hccn crcctcd.This I F doll r bv r-oiw~ilr~ p.h. IfLlnnuc . d tElc pnlncnt~tolo? I[ c n n nc-vcr hc kiia\. incrcn5ilig urldcr a dlffcrcnt gcnrric tlnmc.~pccicc.o f n p c spccimt-ns. csnnlplc.~ddcdfrmn o t l ~ c . ~ ~ Nore thnr whcrc a FPCCIPF W A Sunpnally drscril7t.. inuqt b-. C ~ ~ .r p c c i ~ ~ ~ c n r s ~rthcl.F. o r ivhci1 . thc \hell.clrf 111.trh n pnrhriilar mor. !r. ~ ~ n p l. a n d G. 17): it\ ~pinrof' t11c t'mcll . a nvw and I!lx~rment.

Hatprr I ')ti". ~ p ) ~ lt ~ c .T.ruth as tlriq Irlilht foUow a clgy). 1: d t .o u p i n ~ .llf~ ma\* l ~ c tlir c n w nhi. Tlicrc drr only .rmiilt forrr~~ 19ofi I'~'t0). I Ylh cat !.r ~ 211d un11l. I n tllc moxt rtcrrrt trc.rr~dCracrnf genus).vnlutio~ian !>Ills i t i\ d: lcvcl . 111 I-larl?cr's ( 1 08") ~ ~ l l o ~ ~ u p a 111 hwhich thc aurhor of thc taxnn : ~ n drlic ycnr nt .r.c.~tiorl.lv that onlv a fr6~g-n.~ll\ l . schnoh of ( I ) c. 17 Class Stropho~ncnat. d u e tc9 ~ndrtYrrt-nt prcscn.). 11.IIf y .c' 1vct.3n phyla 111 tlic ~t 114. Fint thcrc i~ morpho.g.~qeI. hc i d r ~ i t ~ f i c dV I ~ ~ I ccrt.~r of n~les.il).t6nh.ire \i7n trc11 ~ i o d ~ ~ ~ t~ it d r~c ~ r t t t ~~~ t t c di 1 17 (dcc. cct whrch h a v t bccn woskcd out s11r ulnrity rcrults fror~luonvcrgcnt cvdution . 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. .ln onlv terit. at CSrdnvics.tt'. t.nny fn.c.c ILccd ~prlIly L J ki~own TL-IIII~.31 .0 I I rLY<lr cslfring .111ri tnnc.!vi \vns c l c ~ .J ~pccieccanriot S t ~ b o r ~ iStropi~onicnidir~.lxonolni~tshavc uwd diffcrcnt m c t l i o d ~ .or cc.l i ~ \1s nrnrtr-n U/rt~~icrr~ r (Eorl1trtrrrr.i olvcd. accepted rlrttllod of E ~ c i l ~ t n t i n g thccc cndc .pnbl~cntiotiarc quc)ted.I~~I-ILX7r .~ 7/)rr/l. ~ .7~ clcs~q~areci Si~pt. taxunonly. i\ Il1c. p .iiic( Iilnny nthcr rc1:lrcd hmcliiopodr. Pl Joncc 1 Q F l : 1 I1 I l .r) Thcrc nlay hc vnriuuc ~ t ~ b d ~ ~ rof ftil~ ~ ~ r i ln l t > ~ i r : t lf c IL)17. hicr. and nnlv ahont n do7cn of tlirrc.~n hracl~iopnrl Fnriroli~ta ndru.rl ~ t arhcr l e i <4?11lc'rnr ~ d Et~pl~trfaritv.~sirbl~rd .i. ~I O ]low do ta?ronorni\ts nctunrl y yo . ihlc ~ i m i h i ~ ? a t i u n trlrcnn. i u n ~ p n r r d with). ~ l they shanlcl . l S ~ 1 ~ .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. t~o!~!~vnnrrr).: lilt :I 11-1 . i . ~ b n l . ~.ly rrferrcd thc taxonomic liicrarchy. i p p I ~ ~vtgr ii similar speclrq .xsting ~ ~ l i o o I 11avc :Ill. ~imilari tie< rcflcct phvloscnct~c ntfinit).rc~~cc+i cl In rcfcrrcd tn carlicr.rrtucl ~11tllnn rT refrr to rlic I.II r to samr c s t r n t art~ficlal 2nd ~~~hjecltivc. 1 n t T (rclntcd tn) nr i f (rnav br. or 111 d l ~ h r ~ v i ~ t ~ac i .\rrnh wrrr tnln grnnpc thcrc if pvun n cnrc for rrccrinc pl~ylnycn addressd b\ tJ.171 c-xamplc. as Et~clivx~r~tcsSubphylum Khvr~clioncllifonncaWilliam~ . I3urrlcril 1806 1'1 I L L .~~11 Ill( I1 . ~ ~.c. miqts. ! : . I to . (.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.111 full. At a .iu ( p l r ~ r i ~ l 1s sp.. Illc*.~ c.ns crcctcti to .A. empl t.rni Where.h tllc.c I o r d c r ~ rlasscs artd clnrsrc I I F tilt Iargrst C11~1siut1 m ~i ilasqical tasonornlsts lircd a tr. hut L-. 1l 1 ct 1 !t-~:k c ~ l ~ f .ncnr ~l~llgcnnis. E !I'TITI\ I n Full gcrius. 11~1. 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:'.mJ. ycncr"in families. p ~ i ~ ii .111 I t : r. accnrrling to 3 tnsnnnrnlc qchcn~r fi l \ ~ ~ t ~ ~ > n r o r p h n l o ~ Thccc.-nrr . i I ~ . I / .I ~UII".mrl I c. circh t1i~v. ~ 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.IS f ~ ns ~ O F F I ~reflect evoluly r ~ C Ullril the 197115 rnoft palncorirolopsts. ~ t c d t l ~ c to mnk ut' a runlains. .itlncnt.~c rhc -/'t.I1~ 1 C'\Cl I'lt.til~ If the cprcimt.IS < ' l Thc bacrc principle tq that niorp11ologic.< ~ K ~ d l~~q~c: clar ig7ahnc R r e d ' ~o n p ~ l n lma teri. o morv rvtrrilic c.lcc.~~~~bonrt~~ct-~ \!.i<c. rtccnt ~ r i . distribut < Althntr~h 13 tnsonanuc catcgorlrq ahove tl~c 3 spccj~*s c n ~ c r ~ thrcc arc rht.-nrcnt prcscn~ccl. E tnxoriunlv h i ~ h c LIXJ Arc t i r ~ ~ .~ Willinm~ #.c.o r lack of CJrdcr Strop11r)nicnid:i a p i k 19.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.hftlllt. Lc !ding r lt n s r q nnd d ~ s t ~ n p l ~ r h chv rllliinr rlltFc. ) . an Famil!. I n t l c ~ . fiomol.v ( c q.~ll\ l'he tllirr d Gcolng~c~l Socicw iri A~iirnrn). rr apik 1 111~1 CI<I# ot. t 1t>\v l>cqt t pnrncuI. crprcl.~t~vt. In Taxonomic hierarchy ycnn th ruc f l ~ ~ r p l y cnntr.lrlul~cs 111 orders.rc prr-<t.Icivr. ~ ~ l r / ~ * r ~ ~ l r r.lr f. 1n crccting .dr . t ~ ~ clrlgcc rl nn up-tir-darc 111onopap1i~~. 15 ( ~p.rtI~rnilx.T.'8~lt~it. ~ t r . Mollurca and Rmrliinpod.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.. n n c wnl~lciIVTYCC e .rvntion 1% pilor or ~f Subspccic\ <crrlrrli< H ~ r p c r 108') i lr. l t ~ ~ 7 l ~ l n o n a y relation\hrp..Soowcrbyclhdnc Opik 1 ) 7 4. c l . ilic ~l~fTi'.9.c t t a n ~ b o n i t a c c ~ hc J.c WCII thc d ~ v l n n i ~ ~ ~ l-tllstcr . hut in dcfinlng thc5r be * ~3t1ldbt...>snnnrn~c ~vtrrk . -3. In n 111 tlic abovc scctlon wc Ilnt.~rter.. l>. I l l i I Inter st.~. p r n h l e ~ ~ ofs C/rtvt~.I T l r t ' t t Tlncc All t. thc AF oi(lcr o t d~qtnc-tof Scnrl. ( 2 ) t ~ u n ~ c r i ctaxonomy and (3) cladlqn~ FTS tcmatic al ~ d e . l ~ ~ ~ . wcrc cvolutinnnry tasonn. ~ ~ r o l n m t ? . ? . 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 ..~d~tinnnl very flex. d a t arlitn~lkingdom.\ l~lritlpnptrts cf. € 1 1 p and ~ thc an~mnl klngdn:~~: phyla ( a ~ y ~ l a r phylum).11.~ l 'U-1 b.Ire ~. Hut I n I l ? > V flcrcl t t by a SCIYCC of 111~ P ~ I S ~ CC.I.roupcd III genera (rinp~bdr rallcctcd in thc firld. . tl~osc wurkins with fowil liiatcrinl bric h tlicy h ~ v c ..lnt rllc r d r ~ ~ i !. for othcr rcnwnr. i l r l r ~ i ~ r r E r t ' t o n ~ c \ v l ~ ditfcrcnr r a.tlOr~~rt. Si~bfi~nuly Soxvcshycllinac (3pik 1030 1 l'. o. it rn. -ca.~rthicnlclnwificntiou.ilkr1irj rrr~plitbr uccd. . ~ of r7dt~1ril Itct'd 6 [ t a t 02 8 . ~ n i ! > q t . ~ ? K I I ~ O [ ~SF. the nrlv S~rpcrtirnily 'rupcrphyl~'.pccit. Tlik is always so trnlun.t..~>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.lrlr pi?rf/lc .

~ t p r ~ r . h ~ c hd t c l l o t o ~ n o ~ ~ t~~s in ir~tificdand cnniputcd .incec.3).I fprc~fic C t ~ ) . on< .rc~I ii~c~ ~~nport:iiit.I IIIJ bc uced n\ .3 Is I F ~ ~ C ) \ V I IA< dadism or phylagenetic rntematics .kiiip .tcnnincd) in Hcnirip war of rllc aysinlnn that rccciicy of coluI l r 11 .iigwith phc. l 4:!.tllrrd ~ c h n o r ? l l e ~ l rlporl phyloqciict~c criten.ilq nrr a c t ~ ~ . o f COLITCT. rlir sl~nrrd llv ~ i~ t poctrnnv nf rcccncv o f c c m i t n o t ~~3rigi11. r.1rnttcr. chnmctrr [or all vertt. I l ~IIC . TIILIY vcrtchsatcr hnvc lurk211 ! lac. c n a >rc irnpartnncc (ag.c much ci~~p1in~i.\ct~cccl the I I ~ C hlallrhing points Arc arr..~diitlcc srr. J cladogram (Fig. rel.corlc~dc.71111~1.lxonnmv.11 for n Fni19 tlnlc I ~ P T ~ backhcli~c the nf p a l ~ ~ o r ~ r o phic) or 'dcrivcd'. S l ~ ~ i t l r 1994) c ~ ~ l ~ n of nrladist~c ( '~ r n ~ ~ r r 1 ~ i ( .i~ldWJ+ \ O C ~ .t n~crhnd strntnphenrtir pnl~ctr~~tr>logi~tr.. ~rclntcd to C A C othvr. W A F tIici~ rlrrivcd ~Ilarnctcrs. ~11. I QS4. sister groups.~ctcrr' d (synapomorphics) ~~pllicnl di<trihutlon niag play an inlp\~rt.lir~ l ~ a r ~ ~ c t cfclrn~ing cladistic aii. anti ir 5t111 C C I I I F I C ~ ~ *tnChc rhc bone<.rrlcr intn ~ ~ ~ l l ~ i t i i~~ IrS O I I~ \ ~Thr .Irr ~lr.jr~tlvit7: l ~ l arc~ ~v c l nlnicxt m c r c . char~cters. thlq is commc.lnd I\ rralirtic h~iidc.11. nor possessed by taxor1 C. ~ . cniph. \vIii~hCIF w ~ snil of cvolu tiom? novc-lnc~ clr 'drrlvcil r urw yr. ip TIIL.111~ ~ylageacticrelationship? . R and C:.catinn.rf cP:i<~it.!. ~~ .41 11~~11r 1 3 u p i n g . iiinrlc.1111 s ~ ~ L ~ J L ' c ~ L Y ~ I rlrclcr to in ~ ) assurncq that d. ~ n d rrprc.bmtus ~ ~ i rl not. h~ whn d :.~rlotlicr Sccnnd.ittic.lartrr staticnc\.I bnckhnnc t~ n pnirltrtve ~ '~ . cvidcnt 111 d~~trihtitinii U I I R ~ I I C C C ~ h ~ l r tnrnonpt o r p n of ~ . Inon o r ~ g i coulcd b c ~ hc choliln 11). i O ~I I ~ Thus in clorcll.vl~otomou~ splitting had occurred t.i t t ! t l r 111 st. ~ p ~ t ~ l c in any c v n ~ p l e s i n m o ~ hthilr dclinc. n< dtd zllr J ~ r ~ n t ~ t i nf tllc piwL~lrm. 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. . By c u rhc 1v. I .1ditij: rdanonshipf 7'Ills pr.lln rneanlnghl rccultr. Now C i q the w t c r * ~ ~inct. 1.$rl rccord In tcnnq of pre<rnratinn.nlcthodology i ~ c rn L-c>mprrhci~sivc that mJv hrlef rrc r ~ ~ r c r n h ~ .'Ir.ic bern fuilrld very uqrf~ll hnth ~ h a r c n evoluliorlary novclq clr qyi+i. but < ~ r h l t .lrc ci thcr 'prim1 tivc' (symplesiomor1. 2nd D Ir rlir d the t ljc~tto nnnlyc the ulraqurcmcnt. i r t ~ u l a rcharnctcrc wirh rcqpcct tn . chcrcfnrr. nnccstor. ~lrcd. a h l c r a r c h ~ ~ l pattc-rr~.lnuu. Eldrrdgc v 111 Cmcmft. What is a pninitivc chnractrr l l'h1) fnr all vertebrates i r ~ 3 C O I I ~ Co dcnved c h a r . They nrc +did tu bt.iynrnnrn nlmc inrt.nini<t<rncd tn cscnpc Trpm t l ~ i hprahlcnl by rncthodcrlom for dctcnnining rcccLncvof corninno nlig fur rlt~.q I havc tn a m m thc tpresrtlt ni~thor).115 ~ L Y C ~ I ~ J ~ C .m i t r n ~ g l .1~(2s h r .~ ~ h t ' glr'irlg c-t'rt.~.brrd g r o ~ i p ~ \vould wc srr~i~r ' i l l ~ ~ c ~ ~ 111 1C7 ~ C k r ~ c o r d.rppronch tn Ficnr~ip's csr~tral ct>ncupt war that in a11j7 g o n p lTrlnnmv. ~ l l h c t n n into ronq~dcr. thc nrcthod itrclf <IF .~s~zing icaturcr rliarcci hy c\rganisms that miftct.g. .~tinn.+iigcd in r ~ c ~ r c d bv hiernrrlucq. Thr): I e ~rn~erical taxonomy 11.Divisions of invertebrate paloeontology 11 logical (nr phenetic) resenlblance. opcmtor l ~ i su clloosc (~l~l?jcctivclv)group o f tllr conlbincd taxa A and R.t.1)). i i n h ecot~ipdrrd to i~~vcrtrhratcc u~l13potnorph-y nnd t~. dnd slrtcr %qorlp E c o t n b i ~ ~ c d A.r. o tdXa nwrl to ' ~ i t .S i ~ c hr c l : ~ s ~ o n ~ h i y ~ to ~ It . thc d~ctanc-csbct\~ccii clu~tcse Hrre tasa A and I3 ~l1~1i-c uniquc cornmor . Hcncc the olljrctivity nf In cnch lincngc 2nd c o ~ r ~ p i l cnn (tinwc~gh:hccd) r ~ncrical taunnorny 1% less th. Lj~lt. 111 t l ~ c t.. 1 r . gmup o f vcrtcbrate.. rlls pus<cssion of .I\ c.whi~-h took . ~ i r r ~ c n .it~riI?erI pllcrlct~crcsrr~lblancc ac tlic nnk4n In rvlated tnxa. In pel. l Qt.\nt pdrt in tvhiub would d i s t i n q ~ i ~th ~ i x grnup fi-0111 othcrs. ~ .drlv d a y of cl.~l .~ I I I I I ~ .~t~is v IJ rliil~r<iliration P C I I ~ C ~ be particularly acute o f p . Gir~gcrir-h.nts .alysi<.trc ~ ~ v hcrc.Ire csprcwrd rv that BC cnc11igl1 ~ 1 1 .~rl bc d~.irucal r.n SCIFQO~ t'o~~ndrri tllc C'T C M ~ R I ~ hy sfl>rnolopstWilli Hcilrug ( I W C I ) . I r ? n . in narure. 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 . ~ c t ~ r f ~i Itlr toillr t n x ~ ~ t n n i i \I~owcvt'r.btc thc rclntlvr xt.Thc nurricricnl 1-lcnnig endcnvo~ircd to providc . i ~ \vcrC r n c n ~ ~ ~ r c d .Irl olyectivc .toron indic~tive of any c l o ~ t rcl.~nlnl. i i i c a ~ i ~ ru f t h c ~ rdiKrrcnccr.rlllrd v r ~ m r o u ~ lto pnlncoutc~lnp(c.oi.. bascrl upan pritnitrve .c. wcrc iiinnv d n u h t e r ~(ir~clitding. tJrc ~ I! %t1h. A tax-otlorniqt thenl. ~ n dgcnOV i0 rot ' ~ l ~ a r cderived c'l~ar.ition\hip bctivcrrl a n y i cndniir rclarinn~liiy~ : (Hcnn. c t r cannot bc clirniv~~ phy.

nated.lc the larger the o f nxa in ~tratigrapliy databnsc.ilrt h ~ hasls nf ~11ascd-rlcrivcdhotnolcr~ur.?cd in txxnnrock record 1s not taker) into . from n c o m n o n aricestor (u~uallynow cstinct and rrrcncy of conlrnan o n ~ n but hnm convergeilcc . T h e PAU?' (Phvlogcnctic Analvsiq W i n g bonar)' ~ t ~ i d j ~ ~t.111 Fig.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. 3 major god of p . for cx.1tionship5 in ccms o f recency borc them.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). 1. 'Time i q a fiindarttc~ltal ~ r n e ~ ~ in~ cvuluJ r on gtanis. paraphyletic groupr arc derccndcd chatactcr S ~ J ~ Cmay have nothmg to do w i t l . R. 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.lnpmsh ( I 'ju0) conia5 sprcdrd up by u u n g oollc uf rcvcral cornpurer pro. and tlic. For iiiqtnncc.~rrrplr. partlcul~r o m p l i i a t ~ o ~ that c is of conirnon or1Si11. to be a h t callv hozv orgmuslns Arc rclatcd. Monnphyletic g o u p s contain the cnm.crgence'. pl~vloqenrtirtrees.for all tasnunmlc methodology..o l o p . rcsultq nf the avo.Polymorphic species. ynlacot~tologi. coupled 1 ~ 1 t h bjostmt~gr~phv srerri.lrrrt t~ cannot bc denied ar an eqrencharacter stat<-T tl~cre JTC availnl. and coulp~rcd with .mcn ts.thcre i~ at1 csccllcnt correcpniidrnce hr.3).12 principles of palaeontology character datn tnktns. 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. r nv . T h u ~ C. character present In a vari. Kescm blanccs 111 c h ~ r a c t c ro7 F in Fip.lctrr s t a t c ~ ?T h e most ~ l s c f i ~ l w3y I r thic. 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~. the order of <l~cccqsion the cnusc troublc. arc t l ~ c and WiIln~er{I 997) i l l corlsldcril~g rclat~onrhip\ the result of ion\7ergent cvolu hon. fur C S Z I I I I ~ ~ C 'is . hnuing can~tructednn JpprOFnate ~ . minimize and thtis ca~lccalconr. it iq an important than has genernllv been believed. T h e r31urc c l ~ n r ~ c andr ~ opporcd by k d l r y . 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. phlc characten arc only prrxent in thc i n g n u p .gmup cornparnun'.AF S L I C ~ I t xc 3 valt~ahlcand ccrtainlv . ~ l a e o ~ i t c h o ~ e ~ d Pars~n~nnv) program.In '~ngnup' (ofwhich The result is n phyloge~~etic trrs. blii gouprnp.7t41. hnwcver itnprrfect tlic I ~ W I ~ ~ ~ O C C F S JC ~I the conctnlctitm nf a uladognm ~ record nctlially 15. ~ifrcn routtncly tnl qource of d. Very co~lunrn~lrclated 'outg~ol~p'.'r%_S). C . 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. tlic succcssjvc ay-pear. apnnmr. T l ~ r r e I( 111ct11nda n a l y v ~ rrlatto~lslupr. Even so. arid ~t forccs taxon. wlicrc sexual dimorphism i~ stroll. A vergent evolution. 'E . 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 .tic analyqis bawd on paslrrlcrny w.I errpntenttd fur re.tratlgraplllc r i d r ~ .d ol-ycrtivity.~plittingof I1 neapcs tl~rouglltime and n rii2ctivclip I rally de<ign.h\~r. and large dLirabaqcc arc. although these may look superfici~lly <imilat. Smith ( 1 944) d i s r ~ ~ ~rsne \t h n d o l o p 111 J c t ~ l l 'out.. clad~stlr of i rigorour wav of \\-orking otlt and rhowing grayhi. fronl con~ mon ancestor and all cif it\ rlcsccndantq (D.3).I tlmc JX<O.1 technique vhould ccmtiilue to be t h r sti~dv rhe diurrtiiii~-at~o~~ . not ordy for ~!ddl~111. Likr\virc.ill tend tlm hencc. Here . 1. result-.I closely 'a"be~tertimntc" of thc tree of Ilk'. There art. T1111sW i l l n ~ e r I Wr 1) and lWoor< polyphyletic grnups cln t l ~ catliur hand.I problem here.qi~rl~cd rhrtrc kind< nf cladiqric T h c ultimate problenl. 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 .Fo: (shared prinntrve) from ~vi~aponlorpliit (shard hine this i n h ~ m a t i o l w ~ r h c dcnved) cl7ar.~ccuunt (though only can d o little to distingir~hsibling species implic~tly l a d o g ~ r It~~ ~ t.but although it record of the stnictwrc and hfc of ~ h c pom-avs t ~ x o n o t n i c rel. \ l Wh11~t3 few 'tmu~f~xnicd cladiqtr' n c q t c the mEuc specie$. Her~ruq diqtln. .hesr way fonvnrd.the tomhincd cladiqtic and hiostratipapluc appm~cts may thmw 11p uncxperted pattcms. \vhich look nlikt n r ~ d live 111 the sdrrle arca but canSo wherc doc? t h ~ lerlve thc potential contrihut not ~ntcrbreed.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. dlld 3n t'q~elltldprcrcquisitc iiv omiaq tn be expl~cir about pattern\ nild ~ o u p s Thc drawing up nienilin@i~l . 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 .

Hrenchlev (1975) and amended by Luuklcy ( 1 qX3) i? I I I .11ttlllcvlltC"~: Clllptt'r S). ~. An assemblage refers to a ~ ~ t ~snnlple €rotrl a glc ~ ~ l v c a ti mlcrocoinputrrs nnd thc pron patticular horizori. :isot wluch scquircs wrne d~scussion. Instertd ~ a k ~ ~ n t o l o g i ~ t 5 !.!irrIl F ~ L ' ~ C F . i n rc1.A sttnilar qCdtfeT fronr n poppnsitioi~ qtrwcture and orgamzation af palaeocom. r ph. The animals themselves are a11 dead and their soft parts havc gone. lLj()i). 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. u rncct thc nerck uf pnl.nnt..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.i dcqigned 7.houph palneoecolo~is obvrously related to l r ~ L l l ~ q ~ r r i < rntdt from lot. 'undcform~ng' in nssocianons inferred to represent a oncc-distinc.r<lhl~t. ture. I qr)C): Ry. nt'llev iit ( e . ~ i n c e soft-hodicd animal$ are not the preserved.. nf . Each .. palatclr~ulclg(hew diwus~ed with pnlnro\~icr::qnpl~v).~lt i t E ~ ~ T I A V11clr to p ~ t 'hiyhrr drgrec of tionqhips of the nrun~nlswith e ~ c h In t .LL rcolog 111.~nr>rr+ dcfttmicd fclc. thc padicnt a i d I. chcmjcal and biolocjcal asp-cts. it 11.lann to the qtudv of Pataeoecolo~y must alwayi remain a pnrtial and incomplete science.nnr. A plnced on energy flow throusi~die svstcni. in t l ~ r t ~ n ~111l a r ~ appcnr. n d r e UI w l ~ i c it~feed. anti t11c rrdncrd major nsns drawn linkage patterns In food w e b and in investigting 1.rl mntennl.palac-occolohy i r the shdy of animal!. for so much of the infomaoon available for the study of modern ecosystems is runply not preserved in ancient ones. msv hc iricludcd In palacohiulla.~tnplr. A n association refers to a group of assenlblagm. 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 . .tinrny p. I A I-.f ~ I I P F C i if r. I \ IT h.. ~ n d past ecosystems (ScI~Sfer. Rn~rrtli .. c:ln he iircd whcn .liarl the true si ru1 :IF \~ l~t of-it% physical. 11 I ( 1' 11.ilv xr~d of cornpanme them nvc biological entity. Thi5 givcr a simple Ing 1 ~ 1 t h dead commu~lit~es. n ~ thcn I>c conrparcd ct.ind tllc~r environments. it is not and cannot bc thc s.I\ .I ivholr r r n c ~ f pix~ihle v o tern.11 111- \ es.!nil fcrr~nlcr d the onc spccics m ~ br 50 r v ancierl t ~r~rnnisrns their cnvironrnent~l context. lW2) T'IIIIII ~ ~ I C l ~r ~ r t. .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 .. ~f ground for iiiterprenng anginal comn~unityscruu.I wries o f present . tnr c.I k9n~ltt'rt.jrc.*( b. pcci~lt\t.I I ~ W ~ V C T .I< a fcattcr dlngmta.mri tl.lh~tlcnlly.tlicrr arc a number of qtaI adaptanon$ 113 fossil orpnitms and ahnut thr rclnt~r::\.lr~. If n rnllectlon of F Al.I. fu~lctln~lnI o r p h o l u ~ n ~rlinology m ~ d . This definitton cerresponds more or less exactly to that of Kauffman and Scoa (1 976). \\ dtlr r l t l l l + nr trmlr nrhcr appropriatr parameters rn modern commui~itvecology much ernphacis i~ 1ri.Divisions of invertebrate palceonblw 13 C r011 ctc [I.t Iir.~l. me o i n u r n c n c ~ l tlie nnnlvsir 1s becnn~lng 3.llli~dtngctlrcr. 111- :nilur cnreynrlcr. wftw. The scheme proporcd bv Pickerill and . 11 lull 1c:11 Palaeoecalogy rl y. rnctliod~: I ~ P likewise enrntial refers to an awernblnge. Normally tills only repre:. to providc the backI ~ c p r t l u d bring of tlir same SPPCICF. 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. the l e n g h / Ir ecosystem ecology. and breeds. h~sanlrng ~ ~ncrcavngl\Important In LIXlliLt~'llIt. \I:. ~ \ > l t ~ tt ~~~~d ~ ~~q ~ ~ v i t .I otlier . sents the prcsenrable parts o f an ongitltlal biological cotnnlunlty. ~t involver the e~ploraaonrlf both IT. A palaeacomrnunity (or f s i community) osl ~ ~ ~ ~ Sr~ti~tic. assoc~ationor group of d.u.~ctlcornrnr~nitil. plllrtcd on a p p h . I ~ I I + I ! I I ~ . = .~rrI tn d ~ r ~ n t a n ~(a? with rpecies I S prrclsclv adapted to a particular ecologic~l lc . Thc rclatlvc s l o p c ~ and <c. ~ rnorc clnhoratc t t ' c l i n i q ~ r ~ f niilltiv3n~te used lirre: o :#).I!\$ 1 .~tontoli)pstq all ~ h o w n g sindar recurrent parterns of spccrm ''1LC.n the.~lity A. 11'11lon rollc~tcrlfronr locality R may hc plnrtcd on munities. . al I .c.u'hcrt.pnratlans of the autecolow of ~ n d ~ v i d uspccics. but t h ~ g IIIIL.lr IIF '"S r~nlronrnrrit.~nrr composition. ppnckngr. I .lrprr. 111~lcx. ~ .r + q~p11.d d i r ~ l . yxcicq different. thr perfarce murt concentrate oti ertnbhshmg thc cornwtcrtcpt on tlic y ax!?. in attrinptii~g h r n here to work out.

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

c

o

l

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 ~

I

~:!IL4:3tly

7-1

i ni ~ % ~

rhq-

!'
(-8

rill- tlir~ct
(-'

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
Ilft n t

p.11~ rcflcctlnn

OF

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tacloting \cdirncnt~ wrrc

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~ l p~ ccvrcl..~\v-l < I N c w Yirtk.n l ~ ~ ~ l c hy i E~TII.r ~ ~ c~ d l n c ~ hullal prtlvof nt P ~ i ! ~ ~ ~ ~ + I I tI X t tJ i~. \ I L I $ C I C I ~ ~ T : Isrrilti. M. p-vGlr (I{ Pftlt~i~trrolaqy.~lSLICL w i ( w ..t) r i c s c.. 2nd Harprbr.hicli c.nrrn\. T .!? nc5.ltl I I CMI~ Hall.!~lri. .A vrrtic. ~ fE. (l'n111. London. thcrcti>rt.~l ~ ~ .tlr.C.I~ C ~ Y C C L ~ I .rt~l?q).rutnln.~ y ~ ~ i vC i~lu i-~ a~ct 1 t sraprcl~ .iI ILI t:l~rt~ar i r dt ~ t l ~ e . N r w Ynrk.il sr.\. ~ l i r y ~ ~ 1 pni~llllr. I).itin: lvill Fnlrhndpr. CJxlrrrLl.111~1 in .I I r ~ g p J. i ~ ~ ~ l l mvcs ~ ~ ~ .-irrl!. .ln ~II ~uall!' I>rzbsccl cjn . ~.!YII to 17vtl1.tn bc p ~ p ~ idlr-lu?wcr. (1053) Z ~ ( ! ~ V I . I h~rirrr :!:rm< dciinrtf all ihcllv t&il'. IIII.nd.l~lrlr~fiirl( i . (ctls) (11)7')1 '1111. } :1 a 8 .~r)rc .nprol~tcrare r.iv. u r o n r n l o ~ ~ ~ .rcliwl~.111dfa~l11.d ~ .nrc usunlly F ~ I c I ~ c L ~ ~N.rrrhr. and I'owrll.III. l c ~ r ~ c ~ ~ t c .~ l ~l ~ i F~L. ~ l ~ ~ ~ r~f~ i ~r.n1~11"'1 a n d i .I l n ~ ~ i ~ c lof r siir'l~ ~ i~% H n ~ t o ~ i [>.tblr rctrrcl~cc) II...n ft7crilikmur seditncnts.~vingp r .dr(r 1il'r:I~ ( .lrv o t str.1lly thc h .lI. 1 I i I M C< . A .rl\.Iilllcal. l t itg i ~ rc\. (lnv.S.11nsrl. ~ rertair~points t J.~c.r o ~ m .ct~r-tx. r. ( I . I I ~ c ~ ~l .iin vcr! fcrv r . 1 ~l n t e r c r ~ ~ r r i n c ~ ~ t : ~ l rr)rrt. :tiid NNJ Evr~l~rrioritiry rllr Pro[-css. I : l ~ ~ ~ ~ ~ ~ ~ ~ ~. cdrr.sL~rriplrs Ll~rringc>n.C u l u n ~ b i Unuvcrc~tyI'resc. tl~cu ?I.~ 111 ~ T ~I~I Cuwcrr.arc I ~ ~ ~ c ~ * r / q j~ p t * ~. I ? tlic prl111'1q.Am<ttrJanl..clrnc uhl q i ~ t t c m ~ \rorld\vidc trwils. V l l + TJtt*L't~yc!ryi. t31t. Cr~ndon.~tivcr ' ~ 1 i l c n ~ c c n l r ~. ~ .I+?~-tci:l~ia p ~ ocl i znijCs ~ . . I M .ucd ..1 if Acntlcll~ic blc Io~91lr v r a rcl.vcr approprialc r o c k rxcnr. ( 1 '1'5') k ~ ~ .11 p t ~ 1 2ih.Sptrhcsi~.N c ~ v 2ntI York. tilt..n.I.~ l~icaI ( )xfi>nl. ) n t v d r l ~ . p n l > nk 1 l a ~ t / .~ .iiblc. utl thc t ~ r l l c rhniid.T 11f:dI) J ~ ~ I p o~c I I ~ ~ I . Inmt 11seh11 4 r l ~ ~ r u r ~ t o lC ~ ~ IC~~ I~ R . T tht.rn~~c. A .\tcJ ~ v i t ho ~ c i l l a r r i ~ ~ r c ~hu !i*\.J. (Nc\.cl\irn'l-r sincr.~lil.*!.ir-. .tr>~~lphv stage. H u t c h ~ r ~ r r >a~~i r d I G l c s . (1Wl) F~uit. 5 . 1 2 ~LIOII.~blcr j c > l ~ )trcut) y K !II-I.ant1 Crc~\vtIlrr~ .\c nlrl clcl~'5 Huctu. J. (Adv:i~~rcdc s t .5( 1 ~ 1 1 .r.~ I \ C 1'.T.ltion. . H (11mc3 i. ~ I I C t:1\111~\ dtn . A J.I\ b(=~i-krtcd hrt\wc.: :hc t.1 I I I.J.A 1-1. rrf cq11iv. .I.at SUIIEC Ic\:c~c~t i 5 f ~ l l i ~ill ipl r ~ c t i c ~ ~ tli:lt diutrillur~nn) . 'Tlic~c.rrrd . I ~ ~ O I I S lr EI~cvicr. i'cnn.1r1d ~.!/' ~ 5 J .~V.. I .~iliat~i..~~~rn.~d.r. Ill ~ . ij . two .o J * / ~ ) ~ .iy collt.ti~~. ) i ~ 1Sidpvlck .~ r r i .. ~ l CC)LII\C. L T ! I W . C ~ O V I tril0hit~--br~i~lii0. (IQC)O) Ti.ljy~licrtrii)~.71 hr~t.c 2 approacl~) !. I ~~vt~nliclcrc. l t p r d c t ~ c a lu w .1nd J~hlnrlski. treatises and s y m p o s i a it nt' T ~ ~ ~ I > I I I I F ~ ~ . F .~ph~c. r rSrr Ifrrrc~r. BlacL~~cll. ( 1 {)74) ilf.:. ~ Cll~ ~ 3~ :#.:t~r~I. and Smntnrl.. ~ ~ .r~. C:hectl~. rc~ Ill runlc RTC.E.I< '5.irlw~. it] m ~ r ~ g ~ i t ~ ~t .mi. 1 L)LjJ.1r11~ d ~ rk*trirv.(All o l r l c ~but still uzcftll text 0 1 1 tl-iannr lifc 7 0 r i e ~.! '):*rc. is cuppI~.~l Ivriri!lcvr. l l r n ~ . t l i r 5ircs of rlepourioii of Socrrry of 1 nrrclotr S~L'CI. 583 py.~) ul'. i p t < ~ .Ire .it/(/ /~~'i>I~~tiotr..(111trrc.I T C L > .i.Irc.liwrll. Prccs.Sr. z p x n c dijtiiulticu in t ~ ~ n l i i nprcii5r zo~r. prc~crvrd 1 1 t h c siltstuilcc 1 4 limt-..11u. . ~ Efirt/~ l i ~ i Veil..r.111 hc .C.~ ~ ~ \ i ~ r gv . V ~ ~ ~ ~ .l-l.W ( ~~ g ~ [ .4ll r.W. ~ n d '['F~crcnr:~! fro. Hi$loly '!I' L+.1l11r rhl>~lyll prccicc 21111~~-r0-201 C C ) S T C ~ .Drntntr. (Invdluablc rurnrn. 1 r i 1 . ~n .p~tt-t d 11%tl -~ .thlc. Ar nloir m .)~L..) (14'45) Tlrc Few! Rt. Ill. I)odrl.IF.111L.n~ild. W.'I:'I~ ibTCLCrCCtlcc I\ 1 1 1 ~.rrrchvntc~. ~ g r r h ~ TO&< 1 3 c u ' I C ~ C I ~ It'v~*l t Ilrll. aild tlic Do.?. o u r c d by rhv hhcB1y fo\siIs.II L O T T ~ 'ti011.~nt 111 1 1 t 1 < m v-h~cli the p P ~ ~ t c r l itvrrc l w t yrcscrvrd. IU. I ~ C I I I . .A. A.-. ~ r ~ ~ ~ .iltrr113t~' f 111 p .t ~ l t ~ ~ 3 l e l 1 .>rr l i ~ r dt ~ 1C .11 ~ L I V ~ I I C C F ) ~ . (Exp1.rd I..~ ~ I ~ ~ ~ I ~~~ . ~ L * < I . 13. (cd.rarrt'. A i :In t.r ~ . r l r .i v Strnudslm-g.ldinnwtrii cl. i'~ '.>n bc c-orrc-lntcrl. (I t3tJO) f'L71L~r~~/jr+ll(!tf)~: i 3 ir .thlc) ~tlj: 1'1t.111 c l ~ ~ c . L ~ O I ~ I I J I I ~ C T \ 111 p ~ l : ~ c o ~ ~ t o l o. L a t ~ Cir ~ c r n t t1 ( I W I j) PIiyltyt.irprr+ f ) .lrrowest t h .C O ~ I ~ C 11z3t lillrnll~t I~ ! the st:~p.ti~ f t t r ~ t q c. LTPC11v \ i ~ ~ ci t i<11orn1. I ~ I~I ~ ~~I l.tcst) U I F i II! fn'ni~il xrcIur.Ill~~o~ . Gvrrt'pts . P n l ~ r .~rivt. r l ~ ~ r .111ri cc S.~llv .llr .i~f:rc:\t hillla1 p r o v i ~ r ~ .11 proviilcc\) !.!~!+l*ncilc l .. rn.I-. 1 t J . -$:.v. ~ tc.~r~dysix. t p hori~ans ~ . L O I I ~ I I I .~lu. (cdr) tv ( 3 'W7) I Irrt. t 1 CEt.I\ y I I J I : ~ n~cillatctlto .. ~ f l t h l i .c-r-5$ l>f lFTLi t . ~ t E1nt. r ~1 ~ h d i ~ r i t l u t i n l ~ kntnvn hnil genrr.p~ ~ r ~ ~ r i i ~ rqtt/' i ~I//(-> ~ f '. ( E l r l i r ~ ~ ~rc.R.~rrStnrrrrrrr ntrd ~.t i ~\ -1 ~ ( # pqwrs) 3 ~ ~ ~ I I / 370. ~ r ~ r e lvorking tirlic Baoks.I! T't~hl~cntlnrl No.d c r l ~ r . n r n ~ ~ t r t ) ~ ~ tI iI ~~~I t ~~ ~ I~ A l ~LISFI~S ~ I _{rt3111 ~ ~ I. L I ~ .~ticc ~ t a ..ill. I* :I p-oup A <ucccq~ir:c zones Ir.is r r t i t * ~ ~ it h:~td to inrrclntc p r c c ~ r ~ l y f!t-ol(y]> . IIC)LVC\~~. (I>cllnlitntion of G~ln.itis~lxt.. n~ethnils) #:. ~ ~ll\rllutc datci \\.lcr:t \imtipap!.'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. .i~. Ncw Yr~rk. NC1<tui.cM .rrfhir. I<.n. W ~ l c y .L o t ~ i i n r (I. Z .it~i-) .J. ~ I I C ~. i\ m o s t tt Nc\v Ynrk.I rcLjotl. (U\rLll bntlc tcs~.~OICI~LCOI .e i ~ tbl<C.~tii. \vull d o u ~ n r l c n t c i\r'itli l i i$ricgs+D. d i c y hcltrr~gtn tin~ 1 . . Tlic 11s~ c~f'.:.rfi.IIILI I rc.1lcnt 4 1 ~ r 'ill \ v i ~ l ~ ~ l y srl-r>ratc~l arc:)$. R..jsSli. 7r1d call.p r i ~ c c w i t i g G n i c I r i ~ ~I<. o r ~ r ~ b ~ ~ ~ .lrdtri~n. l H : t ~ k ~ ~ v I l .::J . (1 9')4!) Micm.irrr! rill.. (13c)ll) l'tiEnrar.i o m t l ~ I v s t ~ v i ~ l wilt11 I>rdChjopt>d ~ t tc ~ IErjuc-ut. (.<i ~ t I/I~-FICIL: !1~fi1r1~.~ t t ~ r ~ Ir~~I l~ ~ l ~ III. ~ .l i r t ~ n l h o\ v l ~ i c lr. ucu. k ~ 1 COIIIIII~III.n tr 11ot ilnpcrc.r.T. IIy nr t11r ~ t . I ~ ~ < ~ J thc ~ . (-Tlir...rt~gr.in'1r~ ~ ~ ~ x ~ i .~ ~ ~ ~ tI1c~ l ~ ~ ~ sLlc.-.. ~ ( .l.lt tcr.I ~ I : I ~ I1I~I. ~ s r.wi\ ( % < ~-l1ri3tjt1strat1g~~pl1y.. (1 9% I ) I'ritrriplrs q{ Rr. ( Ii.

f t c m r r . (Sundard test. (The Frerich 'Trcanw'.irrrl C ) ~ ! ~ T C ~ I P C L ~ ~ F ~ I~ . Lawrmcc.l~vtr. C~nlhriiidjic. Wridenf~ld and Nicolwn.K.K c t m \ vS .-IC.) ( I O N 1 ) P~fcntrtolngy irnd p~lt.n p l a n .D.\rir7n for rlie (Exccllcnt phocopapllic coverage of ~ n ~ puuprj in bun.) (1 % .<. and 5wnlrv. ~ l a e n n r o l u g x ~ ~ l Assoc~ation.ls it1 Pr~l~ol7iolrrq Frccrrlat~.rjr1. -/TI< WiFcy. and Row. tci~ccc. Frecinn~i.4~~ocintio11. (1 1 nnginal papers) lowcv.C:nnikrid_ee U n t v e n ~ t y Prec~. (eds) F.C.S.J.F . I . +i r d L . (13avc M ~ . J I ~2.) (I"7h) '17rr. J. ~ r l d .4 (197h) Stmrt~irrarrd nwrl Stmml Corrr.I< ~ ~ Rtlct~rd1n2 nlanv q g ~ ~ f i c a n z ) . K 1 and West. rb7. ~ rrting) Il.colcyy4 ..fi?rc~pfrii.M.I ~ ~ ~ l 11.~wrrnd-c. + I ffLtl/iicy.lrln)l[ E Irt~r~rrt~rtrncrtts (trant.+II.11 Skinner.~mfi*q\~/( Lonsv~.*: Rmdi~l~gs. t1111i.ln Frnnclsrn I'l~trc Ffltt>. W H (cd. (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 ~ . C (Exrcllcnt tcatl~ookcmpha>inng apprudchcc 3 r d rnn353 pp.~l ~ i i l cto \tr..Knti.f (:li~.11lc) Crolugc-af S o r t r w oKhrlerica. I<.111. 13. A ~ n d ~ n ~ i i Nt-w York. Harlow. L I r~ > ~ ~Ii3I . ( I [MO) T/w >\Tnrt~rd Hi<totj# Fvs~r/s 1-1. Cll~vrr 3 r d Uoyd.E. o l ~ ~ n l e s ) .Wl~ckw~ll.n~l.1..T. Gcologcd .t>i~~tralrr~r). Rohlqnn 1i.rrrri .illi) F . ~ t ~ d~ I I I F T ~K.~tic-r.1rt. ( I cJt){~\ .vrisr ~ I (1h r i t ~ r e l~.1lk11~i. ?I 1 I 1 1 ~ of Lu pmccdurc) (C:lcar trcnmtcnt o i dtfkrcnt .4rw~siron S C I P Irr ~ Kalr fnlann.C.~ ~ u a l i ~ t ~ clawir work i>n phwlcrl lnwr determtniny c .17f Anicnca.1nrd0 1 1 gloh. ~TI~~?I~~I/ . Moore. L l r $ 1 1 Rrt-t~rd . Tr~rherr.~rlnnd. P r w . C:m!g). (lW5) i<i~td~~tiort p Rtfiin~rnt~t~lr C l n r l i * ~ ~ ~ . F ~ r r r l r ar~ti Pro/~ir.l cnvirormlct~t'. G. Londnn. ~ = o l o ~ ~Souiim i$< Ain~.C.~fr i .i\) l'.) ( 1972) Rhlrit.C.I W I ~ ) 'rr~~rti j)ti!C[~t!f~llo?rr~ . (21 cclllcctcd papers) Sttiith.\ppronchcs tn cla\\ific~I I c ~ ~ q ~ j tW. Calif.Ccrmbnrlqc. J-tpp. (IqH?) I'~clhl[. 2nd F r ~ d ~ v .11 n~ap\) O I ~ t IIIS\IY\.I'C'ZCIIC \1orI11 n1.~tihnplGcal lG4lcy. Kan.U . S.r 'Thomprot~.')4[ .) ( 1 Y 5 h ) 77ic Sperie'cr Colrit.(1 ~ d\l. I'. 1ir1 n o t i n o r p h n l n ~ c a l <trati~mpl-r~cal t or dct. I ) .!<.l\v+nn. M~ddlclii~rs.M (cd.-It1 4. C work nf r c v c n l papcrq on pdaculltological tas@narriv~ Tcicl~crt.~ r t i c < S ~~ cn .ppr rrl! rrr P I r b t ~ t ~ t ~ r l ~ /y ~ tq y .<ifirt~rfr?r. p + ~ r s ~ I I C B U ~ I I I ~I'~~.K ( e d ~ ) C i n j ~ i f i r ~ ~ tofr r ~ ~ ~ ~ o c o t n t ? r ~ tDowdcn. I I .tn tllc A n ~ c n c a nTrtaticr or1 Itrr~rrf~brtr?r. A.~$ivr cditorz frorrl 1453) Londnn and P .M.latcd c d i ) (cd. dto M. 13cnn.1 f-. R .<vci/< ! r r / ~ > n t t ~ r. ritr~ ~m C3?rl'nrLi (Lucid rxplanatiun o f rladiitlc mcthodulop) Iluckwnrth.111d NCU.Plt.Ytr. London. MLKKIX .111tl. Lnr~clnr~ (E:nit~rnr~n~ty n i ? ~ r n i t ~ t ~ n n r ) 5ylve~tcr-Hradley.Grti~lr.III? KJCIIC~.ltv Kans. (cd.s L>: A ~~ / 1) Pr~lcotrrt~ll?yy.~rr. S. U..t~ini~m. (cd. <I~ghtly cllrlc* 13d~Errltrfitliy~' L~s\ilo r d c n nnd ch.h r d . A. ( d )(IXi5) Arlhi L I \irr~t~rtc/*rm.rn.rrtr of I)(t)d~4pcmr~rirt Rt~tt~~r~~rrrrtwrr.I~I. but of vcry h ~ g h qualiry) tliro~rglitimc) F1.\rw r ~ .indard work on Kercr~t North Sc. N F ( l t l S t ~ )l.rr. San Fcqilci~cn.lc~l Sorich. hutch ins or^ u P nitrn. Carnbndgr (Ind~vldq ( n i.(10 ont$nal pdpen. (cJcI (IC)C)O) Palaeo701c. Ncw York.iart. M .ll nnd up-m-dm) icpn. Cyqtrni~rrc\ Atrorratinn Speci. d'Arc7: W. (1 97K) Prirrrrj~k. ( I c 5 7 ) 'li-.W. 7 'rratrrt.1rl.f l a r l r ~ ~ ~ ~ . SCC?~CCC. ~( ~ l ~ r r lL)7(1)I ~ I ~ ~ > W I .A. (lL)94) Syrtrrrratirs arrd r l r ~ -fossil rt~orti D a r ~ ~ ~ t t c . G.I.b~il~~n L-rlrr~rqtrt.fionr .t!aeoecnloyv) S~=hoyf'. (cd. l w r ~ n n i L I U ~ ~ .rica No.with applicatio~nfor p. I1nrt W (Suppl. 2nd rdn. C. R. (~?fFici.1j. Coopcr.~tln.1.t7tdt)qiin! liaup.i c h c * w l r l ~ drucr\~wfucru..H.1nr. 7i. a r r i l m d ~ rPJruvcnin.o I ' i v c t c ~ ~ i1.~zt~ir l?i~~i-tlrrc~p/qj. (14)77) ' r o h ~ y rritii I~alorclr. 011 It~t~~ft~I~rat~ t i Ccolog~c~l Socicty r ~ i Atnrlrican ~ r l dthe LJti~vrn?. ~ r ~ g i ~Iprrql C ~ LI I I I ~ I d I~ < t n h i ~ . (1917) Un Grorrlth nrlri Ftrnll P r m .wrrr~~r~~r~rirrn~rt. 1-05 I J Vt~lurt~c Nn. W . W . (S~rnplrbut IntcrClaibrd C T n ~ v c n ~ r v Prcs~.J. Edinbursli (St.. (FC)71) i y i t r ~ ~ r rPr. I1. > \ . (1075) 'Trticltrsr o ~ rIrrr~cn~bro!u Paleattr(~lq~y.lol!'.A .. IJalnrontolr>pcnl As'. I I I \ ~~11.~rlv~ls t11.v I'rc<<. 1t. Altos. Carr~hnclpeUniversity Scott.I <1. ~ d dJ.?. Schsfer. TErrw. ~l t~rrri'lirtll. ~ IC :.r. h4c111nirror thr VL~L I . K.Y. ( 1 1 Prra\. 1E. r ~rt. Lswrcncc K.C. Stroudshcrg. Xcc. (l:wenn. A.

f~ i (Fwcn t ~ rt. {1"?-1) T l ~ p a l . it~ ~i Locklcy.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. 47.IF. (1'177) Ev(b1utit)n . ( l ~ ~ r .u t rcirrlrrulllir> r c o l o q ./opr. G. of bmclilopnd doiilzn.\..~t~c-rn P!r.7-1 0 . K. . I. kI.ln . tix coluililt~nir\) struuturc. Jovsc-y . 1 . : 1'17) C:~mvcrgcnr c\.<c:IIICI I~rnlt.\Jtlrll1~. F.~qi~r. 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. I.~frrt~/i~itr srl~~r. ln~-allna rlcLlr Kilsytlr. \Innnyaphs (tnnnrrmnlr.111dI I U I ~~111~1islict-I 111 by (. ~ l . of oh~ 511:d~b {Lo\vcr Carbonlicrnr~<)t J. ~ n f h a n i l F . I-I.<orla. r t d t ~ r t k c n r y .. Ii1. nl' ILlmclnll Syrr. . .idlmi) ablc. R . J>r~rrl~il ~ ' d r l r t > ~ ~ t39.itccl) w : [(lt-- .Aucllr\--CI~.t311. . prcjduc( ) r i l ~ ~ r ~ r ! y j ~ ~ thr t C . Sr~rrir~rn.C. aiul U r c ~ ~ c t ~ l 1'.~. 574.A. ! ~ V ..M. pclint LIL.~rcd y ~ l . \<. (Si 11 I ~ I L * .ltinn< f o L ~ w r c . (1"'X 1 ) Iihvlog~-:iy .. Harpcr. ~ l y \ c c-I. pp.( I 984) TIlc hjclnlerc pmlllctl~. ( T C F I I ~ I I I2nd R P~~~ E C ~ ~ ~ C I I I ) :H P~O ~ (Whcrc r. 10.)nil Scrltt. ~ l ~ i ( ~ i ~ / ~ i i +lirrrr~1>cor1 111 P ~ LT ! ~ ~ ~ ~ ~Rr$ dl : Cr~iiycrt~~!i~ 1 I> 13.-nc.I-53.i r t a!/' Pitkrrill. a17d R ~ t ~ h ~ c hK K . I ! U I C ~ > I I ) F ~ I I J I I ~ J<n\<.text: ~~iri~ls Strcwdrbrrg.c\~c)lution r. ~l.A T. A. (1061) TIIVli'cding I I I L * C ~ I ~ I ~ ~f Ithe ~~ II Fii~ii. .Ocftlsd.n c rK. M.cnltl.rn!ogcal F n c ~ c b of Plrtlcnrn.~rk\vcll. A C J L ~ C I ~ U C NCIY Yurk. l k > i i ~ ~ r rI ' W B I .rdr~rrrrrirI I . O k l . (llJOS) 1310tnrrc.ding) I!! 147. (d.r.U. R.(-i.3) A rc\-ictt. 3-' Publicadun NII.r. 1 . ~(1OH I) ~ ~ ~ L I I .~fi~ci>t~ttdoqy 1 1 1-47. n r l t l I<v. Perin. { l n f i ~ n n a t i vrllort p3pCr) 226. LA rlrttt. C.~d~. (Clnrr~c pnl.I ~~ ~ * I .F. I F S L ~ ~ . nnd Willtncr. t l ] i ~ ~ U p p r r Jlir. i?f ~ r ~ . ( T C ~ I ~ U I I O \ L T # p>het>2h. P Sc~ntt L! / ) ~ I I ~J T Univcruitv 3 'rc. K W (147(1) t l . 5tr~1ti>t1l~t~j1cr1c>. (. of I'r.<. (. (I'lir pp.lxb~t N n .) 'rh0r.I roinpr!nt?n<c . Val. ~ / tuon of fishfnod.~~qic) ~ n r i l u r I'crrnl~n hrac h~opnd Prmrlr t l ~ u / ~ r ~ i rJol. (14X-I) A t ~ .rl I<r. (lt177) C o r .~llllii.rr! rC1~-i/t>~lri. I5nl.'tl?: t l nbrtdqcd r d r t ~ o nn f this honk nr.llg.ii.~n) rccf cun~plcxcr m i nurth-raqt tnglnnd.~i/s* I P I t11r L J p p t -/~I l(>l-R.12. Stit*rrttfit . IR.A!. .tvpr C)rrlnulcinn.I<M. S Ernnotrnc l ' ~ l ~ . ~ n dA. 1-02 ( T h e tint drtinlrivr 5urnrnni-y niljrnFortcy.t : u~\t Hl.ock\.? n y 1. 1' j11 i11\~crtel3r~t~<. lrrdtb (cd. R.~lSr)cucrv of tl:jiv. ~ ~ t ~ ~ ~J t nI ~ ItI ~ I !q!!/' t s . ~ ~ I ~hI m l E11~1. . in> ~ l u . c (1'175) Ttrc appl~c.\nd I I ~ N w r ~ i l ~ ~ ~ d y 43L-7 I . 1 J. M.C.3 IWYV k113cio *trLltlA f yt-*~pI~i~.) M n t ~ r c . rl.')) i~r.rijrr (rdc K A .K.-lr. 1 ~ 1 ~ .. pp.It.C.d C~~t. G .I. Toonlev). \~)irr. Rqlr~nq of t l ~ c T>at~i\h R1oloyrrll 111. I 4'1-53 Individual p a p e r s and other references P ~ l ~ l i c)\. ctnllicprr qnr\-. [i ~ n.ti~lur c Trilob~tcs:~ : t l t A n t c r ~ c n No.irit(ly) Statton 25. well i l l u ~ t r ~ t ~ ~ T ) 1 I 1 ~ I I . v .. Thoma<. i ~ = c ~ ~ c n l r ) ~\? ~ * r t)1' t r Top Hucic . 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 . 1y-IC]-3. A . 111 Tri. -30.( I . I ~ ~ I I L ' I I I ) tint iiinjor work a n Lnwcr IJ.~tFir~~i/. I ~ ~~4>c ~ S ~ ~ l~ t lD.r 1. fll7ii P t ~ ~ ~ r .~ i i t t l f P~:/~~c[>t~i~>ft~yy ~ ~ -{011nid 39. a n d Jcifctrcc. !. t y bcntinr .). prtlr-ccl~~rr rr+trcd to it1 text) 34 1 -60.F. . C r ~ n v t ! > c r ) .rr~w nr >vst<-~n r ~ ~ .r1~ l s t r i c l .il>ram.OIIL~OII. 1'. C. I I ~ J tl~rc o n i r n u r ~ ~ h r ~ ) c w < ~ c ! n ~ r i-s .1 3 .\di<tlcr1% ~ x c 1s not t ~ h ~ t i t l l l ktudwick. l 17.13. I'ctcncn.In idcn. .. I C~ ( ~ ~ I-I JCI~I.G. c m t u l o ~ ~ *u.~~~!ic~ttorrT ~ ~ ~ : ~ ~ O I ~ I I (~ c Ih~ II t . pp. ~ c o r r r r ~ i n ~ u n t i r n ~ i > t l ~ t .'3111l~i~!i~. ~ l Hcdgscth).i~11r) Cirf1..~rlc~. M4. p (I')t.iil L~lnr'itonc (Upycbr brc.I.t)' I~*rrrla*tr ~ 3 5 .er~lr.. m r c d u r c ..~1~ \ h I~I ~ ~ I I ~ #~ 72. 1-77. (Aott!qii (11 h T I l ~ i s tltr .111tl 1 t . ~ ~ ~ + I l l .ry t~fl~vtiio/r110.IW. (I1l7H) A gtlidC t t i \tr:~t~g.~rrri~8t-i-tltrsrnrr. T (I1+*.. l .-ltrir~rr~~~rr 5447.. c t f thu ccr~nrniin~h' cnnccpt I n pnlncontolov.r~liyi[td .p t ~ i ~~ ~ i g n cEl~oErir~or~ cl . (lnvnIr~.-L.1n2 . ~ l ~ c ~ tSEPM. dip~~ P ~ ~ ~20. \l.llvc c r 7 ~ i l r ~ l ~ l n ~ t i ~ * ~ i Pcnm. M c r n o ~ n of chc ( lct>lnp-. ( I ( W \ )Tow:trtl~:I ~ t a t i s c i c ~ l T l i n m o t ~ . M.UJor!d U ~ i v ~ ~ k i t y . Ilou.17+1ilnti1~r Frld~y).K.I .rlitcrl I."~rh.~cal . ~ * I K (!.isc. r P ~ l i ~ i r r . ( 1 1j71)Tlic cvolurlon uf rccfr. I f .C. (Standard \vnrk.rlarn7uic r o n ~ m u n r t l c \ .i. -O( Ncwcll.ET. Tl~lcs. ( ~~ ) ~~ ~ ~ ~. . (1957) I-Jnttnm conlnlurIittr\.E. h / i . vuC. tir P.3 .ii/r~ty. n /'rtdj/(3~if.rlrir. ~<~pproacl~) I+ . 1. I>.K .7.t Licylcr.qiprr.. 337-s5. IVC-I1 i l l ~ ~ ~ r r . 1 3 t i ~ p:and P. very wrll illuw-.wtt.:ab. 112-47 T'FC'IF. Silurian rnannc coninlunltrcr. .~qc..lcj ) r ~ ~ ~ l r m3.p.I ) . K LVrstl.llllrrntlrlcrt~C S C ' . i. 67.S. J.. ~ ~ 1 tC'.in. rn 1 .Y. ( 1 L):l 1 ILtfi?i t t t l ~ SI>J..1.~ ~ Ttie Magncsi.5. l r i o l < ~ p ~ ~ . (I"X. pp.. h o c ~ r vn i Spti/fcs:> ( L ~ D. N.i>lutint~ J. .tl Sncicn. 1'. ( ) ! .S4lrulrtrb 196. I<.~ dM u l ~ r ~ ~ l Spcrial ~ . (L.~ardp : ~ l . 7'1. Er17lty). ~ c n e r o l (ill. ( l V 1 H ) T h e rea bottom and it. -!(.11.

i l . long ago.l u7ar pubh n r m tvl Ilrhcd 3 q Ft. ILidlcy ( 1 c)Oh) and t v or \ v ~ y <TI . Campbcll and Schopf (10?-1).I n TI<-cdud licrt ~it~rquc per\pccrivc of ~colo@cnl urilc. Inrcrtw~~ irnrnenwly rich qc3urcc of pnrn'ln d. l l l t l ~ it~g tosether iiifnmi. i l ~ ii y l inuludc~ Foinc ~rlfonr~at~crn \vt-rc.tncl.\tnrc of rvoIutio!~nr)' c EX.lnvlbl I l l 1 (c. I j..~cvol\jtinn. othcm 1iirtt.~t. Copc . I t rcmainq .1i4diiig ltu and Thc rrcc thi~ ~ynthc+i+.iirncd p. #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.changc. Ifttt rcwnt devcl(>prncntc in thcnr).inyi tig our wlit~lc conccptinii ot tlliclltlv d1y Amon<-t . o f tlir nncl o 1 1 wlf-rtylc.in a$wwng rate\ rlf cvolurinn. ( 1 9771..<ccnt. o n c v o l u t ~ n ntlicory.lpliv. fivt--tingc.~rticul.ln.llllc. ~11dto iiitcrprrt i t can in \ornc. in inrc.Cyrt~/tr.-~tcd .r it< ilarne .lrll\rnF t3I. Ctanluy ( 1 979).rc g v t ' . whilc Valentit~c r1T rrlariric T h c ~hzdetltof p . Icd rrcord cannot wvc n~lirli ~nform.t r F>rniip tllcny ~ ~ o l ~ z c t = ~ cyccpr ~ ~ l innlcculnr h i o l o p Arc tranxfornii~~g .oild part of. I T1 1 the ( I15 1912).1 1 ~ ~ it1 \\ i r ~ . Llr thc irnnirncc t i ~ l t \ r 3 ! r tl to . ~ ! ltot~kplarc. From tl~ic higl-rlv r u c c c ~ r t i ~ l attempt a t lvrld. niolcculnr b l n l n q and .~coiitolog-yttn or) thc impact oi ncw i n h r n ~ ~ r i o n thc from mc~lcc~~lnr tll.i ant oil>c>\v their cngincq ~ r ~ o r k ' . cvcnt+.otliurI Skclton {lC1X5). N n n . nclv v i r w ~ n tlic Iiaturc.I <tib5equrrit a ~ i ~ e t istarting pnnit.leinncl . and can Hofinantl ( 1 9H9).r trl for tllc ~tutlvnf cvnlonon.73. L71 111~1 cytolnl nr!:.d In the biblioyr. Fullcr t r r d t t ~ ) c narc readily avnilablt.siv{Huslcli.. Lcv~nton{I'IXX) and S k c l t n ~ ~ (1'3Q31 J ~ L . hullding upon thr nriyrlal F P C C ~ C F had contcptloilr of C~IJTICF r)an171n. This section i r not Einq thc i i t > i ~ prL. For tlic fos~il rccord can tcll uq about thc n.-c ~ to^ o f iik. elsr=t~ .l and i\ undcrp~nneilhy a ~ndic. cx-oluhr>i~it is n n l ~palncontolnp thnt har thc T thc following tcst. It i q thc rock qunplificd account of classic neo-Dar~vinl. . thi >llyllWh 'Ilke trying to rc. nlaiiy source.grnctic?. Crrc)] fint rcal n~ulttdi~crplinnry nrnnlgnm of dat. Dawkii~c (1oSh).\nd lirrcc rlf cic.ln7 wilth h ~ l thc pagc-5 13oh7haiisky rt al.11rprcrlrrc o f cvo. rhangc. IOSX).l ~ t ~ t l ~niiy ~de. Maynard Smith (1'375. M a y i i ~ r d Srrlitli and Szathtnary ( l c ~ 0 S ) .11. i t ' o n t o l n ~ ~ nrcds to harrc some (1073).~tiou from v:\nnus sources.ind rhe hoof5 ( bnckqnunrl in hinlogc.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 . in<tnnccq hc 1C)8?).lor r lunnnnry cvcntq t h ~ ~ i t u .c v r o l o ~ .rLldt. I present fintly . this has to hc progroup "f bl iriiled by gci~ctlcq.~di.g.1r rhc For~il record i< incomwhcrc v i r .rpristlrrG p. Ac~-orrlin~Iy. f.~11tt~totr: tllr . tlliq chuptcr thc G.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\ . . n o t n Frovrn f i t . Hdllaln (1977). input of pal.rrt .I .>i ~iioclrni h d q tn he ~dinirtcclt11.rkt. Garnlirl and Vilics (IOX7). Yct it <till rcr~lnins nri Ronner ( 1 9881.ill the sricllcCF ~ 0 1 7 c ~ m~sirllnrg.11liqrn of rvnlutintlnr).I 1x0-Tl_ranwniansvn t h e i t t.inr>rl the xncch.ar%AFcl the ct1clc'. it cnvcricr.~l pn population dyn=irn~c<. Mnr~v yt. t lir>nt rhc 11 m~qcing' (Shrlilon. . tlicrcfurc.itn. adnptix-c r. thc of hioln@i_calphenomei1. 111 t 1 1 ~ rrc. Simp~nu (151431. nu d~vcrrrty in .F u m y n a h r a t c ~ .!.111rl St. ifrcqcnlvcd fincly c t i o u ~ l i a f. M ~ y r (lOli3. I~owcvcr ( 19961.inlc ho\\' organisms rv01ve. tn bc a guidc to all rccvllc dcvcloplt~cntsnnd Jir\ ~ ~ I I ~L ~ 0 I~ 1 1 I ~tnpnrtantly.1p.1111~~1+ of tiiric cvnlurinti t h c o ~ hccn In unrfc-r~cni~rlrng hi$.li I c .~ n td ~ i qig t l ~ c l evidcnuc of thc gcoIo~c.ir\.irly ~t xtudents nf thr.T h c thcnry o f cvnlurmn liiikq togcthcr J nlliltipltc~t (Y-F.I di.lttcrrls of ci-oltttrori irlrcrlcicd tn be o o m p r r h r n ~ ~ v[lor does it pretend c. ~d .ln evoluAltllnuyh rccord nlonc th3r providcc nn Ilisrot-icnl pcrspcctivc tionnry thcory.irht.. . 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 . t Q7h).lrl . ~ gciic Fcun to bc r h. Ertdlcr ~ t l d McLclEarl (18)XX). origii. Stricklwrger (1 906).. thco-rr). .ilrecord. FrlttlltBVC J plete.

riciitlv . ' ~ C I TIl Ii~ . LirrangcdS C I I ~ Ss l i a \ v ~ ~ ~ g t with ~nocIitic.ui.ld .g-v.~tir)ticl~ipc hrtu7ccn c. ~ v c w h c t o n rc.urJ modificd t. I.I clinvrr timc ~c:rlc.it must a fcw .~pl~ic.~lc.lncc of arlapt.irc rhr ~ ' lll~~ntlv CI1sparc11cd h j 1.li only \i.tttL~ptc-d II\ ctlwrontn kryn. cnrrcct.I\ plant5 I Z . ~ I I C g r i ~ ~ p i th~iirl l q ~f'prhtn.Irc.~. n p r d l d Iicrc. lcd Itirrl t o 5. I_ikt. tr~drnccIn .il wltlcm~prc.tee-ord~ ~ v i t h l ~ r v ~ ~ l c oicr>inp~rLitivcbrredinp of ai11tllnl.~nckv s.~n~. n 'vltnl ~ p . t c ~~cc ~~lrrnmr.rrolnrnc-nt niciri o f hcrcdiq rs rindrrstuod 111 term\ o t . t l i o t i ~ lprob.u r ~ l e l e c t ~ o n-l'li~rsbruudmg cspcrimcnt\ .llncontologv d 4ltliou!:F1 Cliarlrc Llanvij~I < y c t i c r .I Irr.llcrC c~r-I111nd l .ibout thrn~igli n l.. rhc mcclin111tl Yt.iIl.elution t h c t r y to undcrnn c. hiuclrrmlrrrv . li\t Ilr~tr'r.it~*k..~t~oti. IVLIT~CTI .ltcd \pccit. and ~t I T ctltlc. I ~ J C I ~c ~ ~ i I1. concept oKwrll adaptation c Thc . 'Notliij~git1 i>ir.r)itllrlc-ln tl1c. ~ l l ~ t~ Ihnnltl h~nlcclf ~ J c i i t i t j c ~I l I A I I Y dl tT~.ltiou i.mv~nctl I T I < cornlp.1q hrrri i i ~ i ~ c l ~ i in d v.lnlev ( I c ) X ? ) t h ~ trlo f~rrt11c.~ll\ ~ n c v nf I tl1. animal\ 11.11 p o u c ~ kof Iv~*tlc111 tli~q :~rcli~pcl.an . \<-ho propn~crl I O I I ~ h c h r c i c .imilar.~r~'riticli~.t>\I.~pprciintrd ~rPi:tt IT In of ~ ill? G . I I I ~~\ol.iHv nil .lnz*in as~lirn~cl the inrpcrfcctioiir i ~ i rock that tl~c~ ' n n i ~ n \.l w11.-finp-rrr\.I? irnpnrXV.chrrc ot thrw ficldc I. ti7r cxnrnplc. prrcc~vctl b\.ill:.inti:iI.rtu1*r: n ] . of h{>r<ti.r~ri.~gcr.lrticI:rJ . \\ h t c l ~n.r ~ F ~ r o~n l i i c lc~icoinp.f i n d nnd ~ t 1 J ~~ tCl I T . the species and natural selection 27 ' v r r n'l~rsl~ rvolutio~ ~ r v li. Thu \xitIi C3f tllr prc-1Janviui.lvc t. He I1.ccrlt Llcllr .I r.lnt{ ~ntcrprctcd hriilrlr up j ~ ~t wltli nlodificat~on' irlttst in all .~1111 rrstncnnn nf n ~ v l i ~ c h made nni1nL~l< bccnrnc iilorrnC ~ I T I P I C E .~tclv 111 m related r .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 ~ 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 ~ .tnd clrlwl!.LII .1t lic I1.ilJ.< liot penllrt thcit ~lircctc~hsrrv.md plant5 ur1dc. Hc cancltided t h ~ t.111 1uc)L di\.~ cnilceprs 110 1o11pr bclirvcd to Ilc sciiablc. ~~i:~.it . ~ t i n r I~I ~ rmlrrrcur5 they inhabtt.~ptcd I i v ~ iin ~ h l iilr n p. ~ r k ' ~ri'wquc~~t popul.J hi5 1n\isllt \virh w m r Tliu\ tl~crulrtencr o f ' r r l ~ r r ' .111 luv~nqclrgatllsiil<!1. I l t c .p r c > ~ n i rt~ r ~ i the Frcllc h i i ~ t ~ ~ r .imlr had bcct~p r o d l ~ r e d111 only . i l l r9rhc. cv~dciiccfor thc o n ~ n I n Lct.iiwrrrnrr.~c l I i n ~ l ~ higlicr vrrtr-. .of ~ C I I C ~ ~ CH r rt-al~zcd I [h.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.rcrlrq Ii. Id C ) ~ I to tllc 1.cnrn~ srnirtlirr i r l c r cln.rct~t I point.d Iic1pct4 IO find c v l d r r ~ c r :lilt J I I I ~ T I .rLc11 . r prilt. . in all the phy51cal and blnlugcal pr.iir <t.r difirr. I I ~ L ! p h v ~ i o l o p ..r rhporv of tioriii~J.~kcriplncc CCIV qliecles to EIII. q~ichad~pt.Inwil 'ir~tinitely i ~ ~ i i n c r o utr. I r t a qeiihe. sccr3rd 2nd tllc lirrlitntianr of ktrnwlcdpc .ho po~tul.IF While npprcclatlllg the iinpoi-t.~tinn. ~ ~ n i i i i . ~ d . ' proccsscf.orcls 'il. Ki1~I1t.u. whcrt. ~ n p -olrCr A~F p h.11t pl~yrnloycnly aitd trt~pl~icallv . l ~ i . I~ll\vect.. clrcun~st. ttiort i~l.ltn I!\. ~ l rcgarrlcd . 185'3). n1 ! i < ~ u p h ~ r d \ .il hpr.ln f r a n ~ I the‘ oldy one o r . IIIL' Gcng. I7ccome addptcd to i t 5 utvn cr~\:irnnrr~cnt. ~ n i t n ~ l ~pccicc.r melit. F .~rintninicnlly.~blr o r ~ < c y u ~ ~ ~ l c c .Ire acccpterl tr~d.ibfc.~iid l n ~ i t ~ L l l . il w~t ~ c ~vtri'mnnv prr-Dnnvj 111..itlo11\ how c..clf-wled *crc. He . ~ Ille rc.~y.. 4 1 tllcqc .ilir.111 vnrlnntr nrl .111~1 morc m.1rf cspcttcd..~:tcrnptc r o Jiccrcdit c v u l ~ ~ t ~ n11nd npcr..ly 'Seeing tlric q . 1~ cjt' ~nodrrn EPCCICF jj-or~l prr-cYi\tIIiy .lii rvnlut~otii~t:.inct'str.anlarck 1inkt. 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 ~ .ltcd oriqin.lLrlrllv.lrJrlvc aii.1t (ion1 . frlt HC 1 1 .~rict\r Lnrl~arck (1 77&1 XllcJ).~ I C ( T_.111~. :I snmc kuid OF iiltrrnal d n v ~ n q forcc.e cpreiicd up ld wch ~ r t ~ t l l ~ i ~ l r)rl!vl~. r l ~ tnr cvolutlnnnsv rhnuyht 111 clthcr x v . hut to this pniiit wc \l1.lu~t(1 OS?).Il{r.hil~f.nnd crqiil#nl.rt tlnr prcscllt g c ~ varicry nf r l c r ~ w* t i ~ c f t . c and . ~ t \ tnn ~ however.ll t o cx. Tllcrc 14.~li k ~ '~ 1 r 1 1 1 ~ c t i i i c ~ d s l~i ~ !tnnl prk.itr' IIAVC' I>CC-II \ O ~10. t d ~ p t a t ~ ahaci cnlnr* . o ut11t.orlc rr~ightrt*. l y t ~ d ~I ? to L*~~(rluiltcrcd diirir~g thr I ~ C ) Y J ~ ot' thc Brnqk. of I\ of tirr~c n tlint n c n t\pc\ crt '\prcic\' h.ttlcv . ~ .rli~'ftl~nt ~ 1 1 7 ) i l .ltrr.nd an .~ted11) t h r \r~ildstarc.r rfc>~nc.i17sc.in t h o u ~ h t . I \ ~ ~ the II~'< ~t the lirnrr of cvt>llttivn. ~ < ve.r1I W 1 ) foundnnotl nf cl. ~ ' n r l d wrplv behrvcd tli.~~tor~'.111 oripriatcrl from (I' nlinltc ninnlm.~rwin nrdcr to livc.r.~t~oei >rrc. ~ lunyii~l~) i l ~ i l i v n t I11 pnrn o r tht. I'lnnmii n r a F also r o ~ ~ u c r n ew ~ t hp.J\ irldcurl part~~lllv I< .-culftlnt: O ~ L ~ ST IIC'\C T ~ I T ~ ~ .i.ilow plnccce oi :Ilr hoof.r \\.c\trnl typc. ~ that rlrnc ~c 1 t tllwr~chr l . 1 I F 'dl adnputinli': i t ha% t o 1lc . .111 \ L lcnrl\t\ \\.nt' gradntint~nl chnllqc h r h ~ .s~ral yrncnc5.irlgcr Ii.11rrc to bc.\r~v1~1. ~ Y L ~ ~ I O I IWVh ~ r r . L > ~ .rcl. ~ y n w ~ I L .isccc$0 n r ~ L>nnv~t~ p. of<-clunc.3nti ~l~ rlr .~ r cct'rcct~. itlrirri. 'clrqcrn 1v41cl1 cil~~lil rrqnrrlr-cl '1..~rti~-\rlarlv \vas it~tcrccrcd i l l ~clcctrvc rrllltli ai-rtl .. l JC~~~1-l%ipti<tr n ~ .111 rctrirt> I.pon.Darwin. r c n.ld Llecr~ r.i ~ t l fl~ppmc I>an\?ll th..I g c a t pstcrlrial firr 'tic\c-rnt ~ I l c brolyht toqcthcr .ircd tic>n:ly changu.rnr.lci ~ l Ion? pcn.td c vrolr~. tlic lvie1yr of h ~ r d c.m tlrc c ~ i \ ~ i . ht did not I. tl~c. I U ~ I I O I T~I v o .~l d~~ri-ihlitiun + c r n . i l .ant-l t r f dit..lvc to hc rtt.111~4 h t ~ : pnrlllhvt niicestors ~17d that i t 1 t h ~ ..I fc\\ thnu.ivi~c the ~il3r i i v i r o ~ ~ i n r n t ~ ..lnp ~rnnkr. b11t ~ c change< h ~ c bcconlc ad.it . oc 1111%.i~~rl.nr~ qtructurc ancl ti~ilc't~on tllr hunk\ n i nnpnatcrl w ~ t lLnmarck. f i u IV.J r l " \ ~ d \ ' ~ .cricnnot~. 11.lm~-tc.

ill euknrvntes (a11 O ~ ~ : I ~ ~ ~ F I Iin:S ' vlnicee.n105r S C * ~ ~ I I I I~C ' c ~ k fi>r ~ivw tlic r h c t ~ r y . 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. it'thc ch:lrdctfr\ tli. ~ r t c n < tth. lio\i~c\~cr. I3) fir1 ~!ri!qqlt~)or !$>.\ ' 1J~nv111.111d of rllc 1. 50 that l i i c views oli tlli5 wcrc ~prcul.11 ~ t i d rc\r.M l v ~ r for t actually accumulated. rough endoplasrnic reticulum Eukoryote cell structure.or .e< ~ntluc~rccd tl-tc I?): En~lisllmnn M. 2nd th~u nll varl:ltlon Ir ~rllicri rcrl.tcr<' tvrrc inl~t. u .ntcri T ~ n n ~ i tlvnq thnr Figure 2.rl n ditkrcrlt th-otir.1 Fr. A~~imal cpcrin rc1.1111tllcir nurnbcn.Ire it~hrritcdby tiirurr yvilcrG~tionr. 'The t11c.irarr~r~ I I ) ~ Ir I 111~41ly rrg*~r~ll-d w a d . 4.I ' n c d ' tc reach Tlic pioneering work of thc Au<triai~tncd Icavcs lip un thc trct. Ir pr~. thc t ~ t h c rhand. T h c othcrk Arc wc.)c.nn) <IFiiihc.cbdticc.111rl deosyribou~irlri~acid (DNA). ~ y ~ T. ~ ) ~ C IC ()I! t / 1 t a [ l t Y T f ? l O f ~ ~ l l ' i / l ' < /I)) 1 1 I t ' ~ l F P LI/ / 1 ~ 7 f f l l ' i d . 7. T11c hvour. had 11ut rually c l ~ c r \ v k ~ u ~ vL ~ ~ ~ C ~ ~~ r ~ ~ C wcr:' ~ ~~ < ~ I MU Z ITJ > All thi\ ccchrn< luq.~ ~ r s t ~ ~ ~ of~l~crc~llc t t I \ ild ~ iirig the rnvlrmunli>nt npcrntlny to prtducc.+s \tr>nci thc tcct a( time .linst.ud the founno . food.Tl~ic tvh.illlr.il'/<'ltitlfl.\ kind of' ~ c u ~ t kcdhack fro111 t l ~ *i~ipplicd l ~ c I C ' C C ' s u~ .28 Evolution and the fossil record clrpnrlq ttruct iri\tv horn lrciv I I C ~ C L \ and t h ~ a thu nnturc of' v ~ r i a t ~ r ~ r l and hCrrdity wal I~scc there '~url~ritc~d c.11zJ if gc~~cr.I of b\ l.IT L ' I I ~ It-iiic fhdt h. It1 rlte ~tnlgglc r lifc t l ~ c > w h ~ndividt~.id-IrkL r~iilg thr ~ P L .~r.<c.lrlins: pc)int fhr I. coiiir evi~lrncc.h. t h ~ iirw t I ~ I ~ L13. .111d thu accuintrlatic.iint.~bler h . . .ir thc 1 1 ~ .il~lctlico?' f i ~ cvnlunoiir . hodleq. ) t~cI t cratlon..cdecl out in tlic intcliw rr>talpctltir>ii. thnligli flr~ctu. particular ciiviroii~nciit ntc thc i (>TIC$ tu IIVL.ltiorl o f ~ p c c ~ c sv c l l p ~ .rncc o f Grcgnr Mcndcl i l l 1 Xh5.1ter schc~nl nm ac~luircd 1-1l. J / ~ l I r nrc T h c 1narn prjlrltq of clic dicr>n.trtcd. t l l ~ .jt indiv ~ d u : ~ h .~bIcc t ~ ~ r a u C trm Imds to tlic ~ c Ir. the chromosomes. 111 hlc unknown.unrivc. ( ~ i l t l i o u ~ hl ~ c r c i u d ~ t i o n gcrlctic cxpcrinlc~lt t for .tq: 17rlt ~ii1~1>und.rnd t h c u n ~ It \ V a l rhl\ .i vacuole .>rIl~irrgILL^ :~trd(~vitl~lil c l t l b ~ r < nl oftlie qamc qpccicq) tbr IrlJrcs.trf tltc s~mffc had ~ t l d t i ~ i l f f ~ ch~r~tlir ttllenry to hc fccn to v.r in rc\ponuch t t ~.rl~rtc~nvnrd.1nvin'\ I L ~ I carli inqk~ncc. Anlrnnl popilt smooth endoplasmic reticulum lntioni.lorl: i r~ Ilt.rcxlucr ariorrx r'lpidlv than ii ~ r r r d r dn m. 3. t c ~ ~ c c~ r 111 for k '.lt r). ctn p m v ~ d c d. ~ ~ ~ ~ ~ tn~i tlic' timt gcnbcdr c .ltcr d c ~ c l o p n l ~ i i t ~ .comc lo11gc.1) c ~ ~ i i t ~ ~ elotlgzted t11rc. f. OY ~ nucleus (Fig.H. tvhn I ~ a d w r i t ~ c noil t11i.lt ~makr >llch sliwivfil ~c porsible . 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 ~ ~ .I lopc...~tins. ddaptcd rtl p ~ r t i c r r l ~ r chnvircl?lrrlcnt*.iEth~i\.inirrl:~I\v~n'.lblr (Hc-rc hc ii. wl~icli t1-iaJc of prntc~n .t. 1. 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~. ~ 1 . / mitachondrfon subgcrt <clrrlcyc. tcnd to rcn~. . I .lt~lr. Tlrcrc.tlly ~irlrc.arr cnrlicr. bactrna dnd C ~ ~ I I O ~ J CtJ1crc~ I . M o r p n which b c p n in 1310.i\sic prvrtlilntc th. 'qtru~glc c x ~ . with orgonellel. ).y c d thdt n ~ n v i n . I11 t l ~ i ~ Cnlcd c ~ t they \~c.lncitr 19 c~llt-d'n.1nt1 ~~': Inheritance and the source of variation p r n v d d c .nnri tlr tc-pl.ric.\rc c t ) 1 t r . must thurcfi~rc cnirlprtiriun \ ~ i r h i nand bc hchvcct~ ~pccics rhc.~ls tittcd host to currivc 111 . Wir111n s p c c ~ c ~ .Y ~ V I J ~ o/~ ~L i~ ~I o z t r t d ~ 1 t ~.tit. \ r .11 \clccrlon0.) 7. arc rtniq. ~ C T I ~ I C F .alcllo~rgh.lt~l n\ cl.

~ .~Afound ~n l i v ~ n gnrpni?rn< . r>t~rnl-r.) r the pn1nar-y Ilnlti of I~c~rr~din. TIIISF :I neudy ttncPerI 1. chrorno~nrric.ntidch..~ttacI ~ L % to t l ~ cpcil tow sug. 14 t~jnnerl.x <ye(-itjcprnt~iiis. nia! Ilt.n~.h c g e n o m e t $ . 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.l\:t pl. ~liviii~rrc~ c d 1i1. O I > 1 1 1 1 .cr n t the ~ u r f i s e01-n yncric rode.md ~ b .4rhrtri try iir tlic nzht orrlr-r m iiinkr. ~ d ~ i ~ ~ i i c .~ ~ ~ J ~ F T I I S tls*~~~\r..lrts of . \\ htllC ~lrqnnlsrn. art1 i.~II 1 Lrnd of ~lyl~.1~ . of.r cllromoro~n~.I. rnrntd 117 rht. t ~'d ll .~ t ~ r nf l~r f thp x ~ r ~nnrl . \ gu.sc. but l 111 pT17~ I I I I + dnri 111 illrcr. docs nor ~ l \ r ~ a \ .: r11r I C I I ~ T ~ O C the chrnmn~liroii~li cnnip.t iil\tarc14.b l 1 3 ~ .r r x t together 111 rvntl~cslzl~lg . I t h .w rhc protein. r part~cu o r f u l. a'l~chc can b ~ . '1. cndrrrg h r cets v~l Ill . h lrcttirnlc lurl~. . . . .lrt a t ~ t n c ~rhc ~~'11111.dtld a t prrwnt ~ ~ %':pvrmrnt. t m v g e n e \\'FE?illr 111.lrc . ~t. 0rgs1112c the c t ~ u ~ p n ~ ~ i ~c iddirrct thrir iiuildiny.ltinn~ th 111 rn. illlt in rhr iinrlrtls.murription'.~\c\ . IlaX 13ern chorr~ii It rrccn tly Ilnwover.11. < -Tli~s tnkc placc 1 7 ~ r i AJcnu~c c.ivhir 1 mcwcr oiit of' rhc t ~ ~ ~ c l e i i c . rhc.ir \vl~ichcndr For ~ ~ O I C I I I Fnild introns.uimc 11111:' togrtI~cr .I~ Ir>ily.mrl%Lmvur.i n c Irc r\ntt1c~i7cd. 1 ~ . Thew gcucc q~rctrc-csfor cvcrlut~ont1icor-y.tily the cic~c-lnprn~.ut-sk r ~ o ~as operons.r-vr~pl~lsin. zvhich arr rlicsc arc tiiilctronal.iry t KNA. but \vllrii nuclcnr ILNA is rcprcrc~.1t g c i m d~emcslvrsarc inatlc ut' hvo lut~ds of 1:1lcrnf \r.lkr protrinr.lnd when tl~rs romplcx proccw 1s coir~plrtc. Scn~ctuml3cliz5 c.. ~ t ~ d prolcL. .1non h .h rlo .I wqlc-\m. ~ I . rcldcd h r by thc 11~1 what I\ Inipnrt.ire Invr 2nd rhc c ~ o t l c u ~ l l t t.I ~ rr. ir conlplcr nlc)lcculc~. i:lngcd in discrcrc LIrilrq. w-h~rh tontrul dncl yrs'tc ti.I fu~rthrr ~ r ~ ~ l ~ c u l c (mcwcngcr wnsnr gciie receive? a n JpFropnatc 7 tllnnl~r c.111bv nctivatcd a i d cle~ctivated K iA artll 11r3c. ~ c ~111. i infim11.l-uc .ise< ITIA cl-mrartcr:: arc v c cnlr~ples. ~ ' h ~ .icrd'i. genes.cn(l+ 1.\ rhc vhcltlc.tcr~t~c R ~ I I O I ~ tlic Fcnctic c o i i ~ p l t .~c-id.I J N A . ~t%clt' thc nlwconir.ynthc<i~.un tarkq for ~ r ~ r h c c t ihlolo. ~ r hrcn F I I ~ ~ V I . the species and natural selection 29 Tltc DNA ~ric\lct-~tlc ~ r r .Dutwin.f n~hichtli:.ly. In ~ O I I Iklildt in .~r-c. transfcr I l l n l n g ~ ) rht.Ire spIiccd r n ~ e r l i ~ ~ r . l i ~ ~ k i r ~ g rocctlicr In up .lnd 111 thc nngn. : ~ n d rhcrc : ~ r c tnnrrl. .iiw N.lr wqurslce. whcrcar tflc 'nri~gs'arc rn~tc-hrlip ~ 1 t s nor. p i r ~ l fi ~\ ~ t11. (TI rn 116 I C 1101 a a p a r t ~ c u l ~ tl~ctaholic path\v. DNA fint pro..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.cc~mhin.hacl~tllc. thrv coding !I.nbow. .n specific o t h c ~ r Are 'regllarnr' Srilrs. All tlic nccc<s.lr D N A I F rrarl~cn 1 bcd t o r c l . snd o111crcon)~l :rlIlntb nr.Ire switcIlrrE on nnd otf oi tllc ~ ~ u c l ct~ rthew slrcc c ~protclu .ilo~ay its I r n ~ ~ h . li )In(. In Iugllci.111il thc pcnrovc sug. \vhic. g e l w ~vhicllror~tsold~tlircritp.1 4t inyc~Iv~'C1 ~ ~ C ~ ~ I I C I I I 11l)oqorne it does not ~ . ic cular p n t I ~ w ~ h ~ lead frcm ge11t. .Irk.ir:itivel\' fexv ~ ~ i s rllutflallg -:~'\~cilt~.I Iprotchin dirertl~.I platcln 5eqrierlcr.hi] t n l q li.il 13NA.hlch st. t m ~ r tn m ~ k c rhc htnci. \ .lcrioilf. ~tw. the teclilt I \ .ivc rlrlportLlntcuiiwT ~. c ~ ~ ~. Though r t ~ c ~ c tn thc cvroplncnl irlg I~'. \' c Nltk4 of pl~orpllor~c .. thdi~ tiinlre..~E ni-dcl-.iltet-r~atc cnimponciitq arrsnged in series.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 .l~ ~ ~ ~ ~ r i d TII~'>C nuclear R N A it r-ctnln? thc orpniz:ttion r d t l l c on?L . : r-. PNA.c c . TIIUV.111 tli~. .5 ln thc nuclear u~crnlir~ir~e. Some p r o q e u has alrcndv hccn 111adc.~ l i ~ k r l n wH a. tn\ilic.t ' .cc\.i?r the Tencs r~vitclicdr r f i a n d or1 tngetlrcr.r<tcd. 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.lr I nlewrngtr R N A tlic untroiir . hon~evcr. 11 onlv Iurr w ~ t h 1 tl~vininc.irrviny rhr Whcei mcwcnprr RNA nrn\.. T h e w ~ r exons. t h r traii~ferIINA n ~ c ~ n w .rr t t ^ pnscthlr.il rrpl. 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 .l$acent grn.tr~u~~trnn li. ~ .intI< are thcinsrl\-cr IT] iiclrJ c n r r ~ ~ u n n t i n ~I ~rcn ~ ~ ~ d te ld~ t ' . n ~ ~ ~ l e .I . .!jor n i u t ~ ~ for ~ ~ ~ n l r r ub a l l ybut thc u ~ n l c pl ~ li r c ~ .dr)C( ~ 1 1 ~ 1CXOII ~ l l ~ l f 1 .r~prnducts \\.ncc 111 ~ v h r c hany nf r l ~ e 21) lnrgclv. I rcnll uccd to stor2d phcnnn~riion. ~ rtn bt. this wlrolc ho5r of c o n ~ p l c sre. cvsdciltlv thc ~ n i r ctlti~tilu~ the ~~l For ' i l t ~ I+ I I ~ drvlxvnhr>w..rnd~=d ropy o f itrclt.lc~~ay rhynlinc ntirl i-iho~e wgar xvl~cnnccdcd.ro$i~ic w1t11g t ~ ~ i i ~ ~ r ~ ~ .III ~ i i i ~ t \1.IT tli~s I for r . i t ~ l .lnt I< rhr w q ~ z r n r cnf tF~cin11r lluclrnr 1TNA.r tl~cst.lninc.I 111: Ijllr ' 1 .thc ~ n ~ c l c .l~cr arc innil~frc~l.1l(r11. a114 to i all concrnlcd orllv with the yntlwsir of rnntrnalr.~~t fitnca d ~lirtlughvet. tran~fcl-rcd of dcveloprnent the priies ..ln-unu . ntlotller ultemirdiate ~-l~ulcculu. st.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. Tlilq rc=rlilcnrc ai ilttcleonrle h.ICCS.<cd nr tn P. l . ' i ~ p r i ~ l l t comtrt of . but 5kirh gellef releaee cheiiiic.~ct~ial tv t o corp.ICIA\. r a1 nl 1'1 ''CiF. Slriiplr orcnrusrlrs such A S bactcr~a and f ~ m p have I tlir~utgh prrc.N r . o l I ~ : ~ h cnn cu~ct. ~ <III. relr~sing proclucrs wluuh F L ~ *Ill\ tr+ r.thc I3NA rrr. . m.~ l c ' l ~P. Whrn thc unclc.urnl h~inrlrcd 2nd In ~110rtp c r l o d ~nf tinie.

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

l n ~ ol the p ~ r c ~C t~ I ~ O ~ ~ ~ SIIICC T ~ C S l ( ~ I genctic nuntbcr 1s H.2 The rtnges of mitotic cell drvision in a wkorptic cell.: "' . . .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. centromere prophase metaphase telophase !c~8:?.. interphase . . and the chromosnrnc .'I "'.nt cell. the species and natural selection . 'Ir I.1nd none srr iclentl. 1983.\ornrs.) p m c t c r froni a rillglc p. arc caid to he haploid. . ' : l l .Darwin.lrc. For explonaAon see text (Modified from Kershow. Each iriarcri.. ~'lronrotornc\ arc uupnircci..-.

is producrs o 'rcprr~rt~~ct~nr~ It 1~ T T F .ari. In t h i s proccs\ of recombination . h O~C~ . -- --\. Ar.arc not ~lccccwrily d c ~ l t ~ c fur . n. (Modifiedfrom Kershow.kllclcs. i1pt711 r y r rulour.I-< ' .In.a . SUCI~ l f f U ~ t tlldv ~ ovcr a n d rcuoml?m:rtioil I \ t l ~ \.-.lticrrl \\ itliir~thc fro111thc mlnor to thc very \ u b t t . .d R 2nd r in cnnventlol~~ll notation.32 Evolution and the fossil record ~nterphase <-- -L..\Kri)~iicllifi-rc~~ t p..r ar~ctpar up.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. R coritrtll h l y brown t. I I ~ ~ I Y f I F higli. 7 -. 1983 1 hn~vever.4tt.IJV t-stqt111y C ' T I C \cieriving ~ .tllclc<o n I~orrinlogouc L-hrornocnnlcsnrc of t l ~ c \. .-- .yc rolot~r. t l ~ r r cma!' by only tn. . ill thr Fenef controlliiig Iiut11: .rs allelc~ '. d l ~ ~ I.11 C ~ I C C ~ ~ C K I I~~ C F .3 The stogas of rnerotic cell division in a eukoryot~c For explanation see text. d c ~ l c thls v a r i ~ t i o t ~ t 113 tur..tlwv 1n.111d r cnt~rrollingbiuc r1 culour. .l i ~b V[rl~cri ~ lire* icrtil~zcd cpcn n tllc I~c)riiolug~~:i.Thc ctf~hct of 110th c r o c ~ i r ~ gc h n r ~ c t r r sit CC)IIXTOIS.. " chiasmata anaphase second snaphase telophase gametes cell. 111 rhc hornozygous conditrrl ./I !. rR or rr. s q. .. rhron~usomc.lrcrl tf coin? tnger11t. prophase .. % .us c p 17v CX~FC a IILIITI~CI ill of ' C X ~ T ~ S S I O I ~ Sknucvu . -. . ~ t S o ~ ~ l c o m c s . 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..rcrrt<ourcchc. rnetaphase .dI1L' fkorn d~tlL. \ C Z s '\ . ~ ~ l t l d . /' " . Figure 1. .

~l cliatigcc in gcne I*' ir><~trgR will Iln\. may ~ n v o l v c inversion. ~ n d propornon of thr rng. Thcve t n l d to llavc larger-scale phenotype I'ld<yli~~rlv..~hy I. or a pair of base5.c.rllrltq d c < i q ~ n t e d q lmin dkcursed above in nc\v cnrnbillanonr or geno11. I~ICCFFIVC ~ C I I C nipl lit b r espcctcd to giht lmt rffcct? a d arc njorc often disadvanrageous. of thc lixir htlrn.: o i vari. ~ I tn c x ~ it n L I (0 11 ~ 1 1 1 ~ 11 ~ I~I ~ I ~ J E~ l o o d v.lrc c c ~ ~ ~ t r o l hvd ordv a ~ i n y l c Thc gr11e~ lc I nh~cli or.I 1~ii~tvol1it1on. In t i c t the proportianc of allrlwlt It. .w d l i ~ t c r r v ~ g t > \ ~ ~ . A.ce.~llrlr\. 1911 ind~uidi~.lgc cnlourr 111 p.. and It-. !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.~ . +nup U qll nr qr. and mutations )rn[rr). 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.qngc. 11r 1 kc i. But i f evolving orpnisrris arc cvtLrto +c risc radicallv ncw there muLt be a murc-c.tl. >I 33 111 thc heteto~ygouscondition rrrnt kltidc. . This i s rt~l*r. Even this proofq' ttn. t l l ~ C~ fur I ..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 ~ . ~ . cx.I. ulally plants increase thctr porcnti.I part~ciil~is ~llele.. being ~ r h ~tt r ~ t h c s . inultiplr tion of thc g c n e Mutations on a larger ~calc~ 3 1 1 afict from tcrls to thousands of base pain. it has recentlv becn &own that rnnny 1 the t ivithitl tllc popul. ~ n d 'rc 1'' ~ n d rcprcqcnt the rrlativc proportlorls of putr thcrn hack in tllc right order.r f nl. I t ~ l c h. tcr recessive gcur.1d tt'r Isri>\v~~ C S . brown c y ~ c . ~ < irc cffrci. I \ ~ I I I .lrakcen A wtr<tin~no~-rreatrangenlcnt of rlurleotidc bnsc.><ca r 1 ~2ld hc t t ~ c . tn have thcir nnlv hc ~ t i ~ f i c d whcn thcrc ir .. ctc.irnplc-r dc.I poyul. T l i i ~ ~ 1-rliun. i q no\v apparent (llrakc ' ~ n l ~ -t\h ~ r IrF 3 <\~cc~. Tllc cqt~ilibriiumso dcfined then are f-tucd in tlie rhrornnwnrc. R .II l i ~ i ~ . a ~ t h i ipnpul~trot~q l ~ ~ r not ttnr~ndlly c!l.irintinll illllcrerlt iit the Kcnc pool o f .. comparativclv rnrc cvcntq.YSO~ Evolution can thus he cnnudcrcd as haccd upon frcquenctes tlunugh i~atural '9.~ i o t l . . 4llclcr 11 2nd 11h ~ v r cclu. 1983) that Fpontancouq 'Icsiolls' arc contrnu1t1 one pfimculnr type nf nllrlr. l l + ~ advat~tagc ppotse+\. it. 1.' itrrir .~~-ttrs.il\c7 lc. vananon. This kid o f rando111 chan!gt' ic known as A gene mutation. l greater gii~lnbcrof l o c ~ L)r allclr<. wl~rrl~r c the tnoqt a inon. l spontatlenus rnutatiom. thc b ~ l . hnwevcs.111 blond po11p5 C2 to nrlvth~ng iic~n. ~ r ~~c1c1 1nltcr . rJ :. I F r.ltron . ~ A l t l ~ o i i ~tl ~ i was fnrmcrly held that mntationq are .Darwin. ~ ~ I O tloinu.igciri pn~srscing. I.~rcly tiilund 111 anlnldl~.T h u rimplcst kinds of mutatloll nnsc bv o j ~ ct~uclcotidc haw. Ir rnrnerrlrlcs happens that a getrc will huddrnlv t ~ t . ~ p17. snnlc ~ c l l c fart. l I .+lrrtrtrmcr . at' 17 will hc blllc' ~ v c d ULIZ c r j n n h ~ n ~ . rc. Natuml sclrction vpcratcs upon tIlc &?cat rtock of . 1. 2nd ~ n .. Thiir if a While 111utatin11shave bec!l tmdi tionally undertcul. I ~ I ~. If tllcrc is t ~ o tuull ally occurri~lgin s h r o m o ~ o m e sduring repl~caholl. II. This acts as n 'proofrcadcr' which picks up wrongly psired ba~cq.tnt hulnoqgotc< (KIE) and rrcesrivc heterurending systeril sonirtitncf dipq up. Ilr .31 rlnm~nancc ~vlirrras types. dupl~cationof chromorstrcrne kind. l~fuc.un a< three . L~tluvr.a ivtC1rr C L ~ T ~ I \ J I ~ I Ji~ ~COOfII OI U ~ c ~ bv Ir rruuld I yossi'tllr with ot~ty. thr Rr or r R o*iII..p r c n . Htit such niisnr tlit.cm$+inp ovcr. un :OH trol of' two gc~lc$ xvhich 111 dlffrr.mation comc ' 11(7ninzy~t~rc &711p A mav carry P 13.t.I drc~ded c l r c t ~ v erfftcts up017 the descendant.lr. addition or dulction o f long rrqucnceq nf DNA. rearralipe~ l i ~ c d ~ l o ~ l l ng r.lien t\vo nr niurr CcncF control or :Ic rc.arc all cclnqcqucncc\ of dipluidv. Where doc? this x~. lrllaqc a pnctlc cquilrhnun~ is ~rlwltair~cd in there m y be mailv pcr cell cach day.r. u. 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). III chrernosomal mutations.und rl?.icirni u ~ l Incrcasc. c t l'~\ tiition. yellow ~ c y u c i l r t .n yrcsdi~cr.rrc uontii~udllyrc~huflcd the mccE-t.\.11 \vitl~ gci~csaffcrtrng and spontnncou~lv change tto a new allele.tdun\\.lr phctlotypr 17 precenwI bv natural qclecstood as being due to DNA damage or mispmcrn.~tion. the species and natural selection n. ~ ~tlr. rc~pcrcivuly. irrt r~\amylc.I: 1 Ir uwtllinant nvcr r. C'FPCCI~II~V ~ I I O F C o n a Inr~er I.w h ~ r c tlic ~ ~ ~ T ~ < ..a dliplication or loss of a rrnall part of thc I-i Inr ni.Ire ~n~iltrpl~. ~ ~ i c r l t .I s~ndcp n ~ of rcpl~ccd anothcr b x e or base pair.or .11 vari1l11lbr1 by polyploidy.

uf .t light.will nrllv be ~1-3cto sprmd ~f (1) it ic rccurrcnt condition. lrfunlly of cmnll tcr ~C(.ire I .tllc inhnhita~its m\vn+ . ctkr-cs r l f c l ~ p l ~ ~ J ~ o r g ~ i .. LT !4?ylc-:llor l7 tlO11~ %<> t11.~chp ~ n o d c r n c ~ r r i c s t s ~ v n c ~ o gcnc pool tl~crcir clc. Tllc orig!i All aUrlrs havt rc~iiltcii&(>in rnut.dirie bcing c a n ~ i d c r r d I~igh.tlri-c. lrl. geuc. A iiiut.f si~clinatural yop~il. \\ liich drc DNA r n u t a t ~ o ~ ~ . I ~ V .lr inutntmlrs xvill rprcad v r n (1st 111 a owlv nnilicd >urnping genes'. I r l n n h i r ~ ipoplilat~on< thcrc arc.I p.o t ~ l i thcv arc dom~n. The nccurnulaticlr .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 ~ . thc 'parei~r' 2nd ' J ~ i i g h tcr' T h c ratcq of 1n~tt.IY L-ollle 41111\' . ~ I m o . wlielc gr. ~ lt t~~c 0 t0 1 ti)r .c.~hli~hed cvcn I o n e btnmly Ins!.I p h r ~ ~ o n ~ r n o n ln 1%-Enrhhas c..rtc.ltiol~r t11. 1 1 nonq ~ r c .itnr. mag bc ctio~rgli ro rnndlfv tlir dornrnnncc of .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. u r 1 cl~etrucnlnlr.v h ~ v c x-I\eil .n ' arc spccics n r t then c r ~ t ~ r e rli4titlct.ittot~ o f t l i c pdttcrll n i p r n r esprecslon bv a quch ganic>dcmcq. bccaua L Ic>catiotlq oi tllc Ilvhridv T l l r t c nr >clr~c.it I (lilt 711 ~ l l ~ f l < l l l llv <011 1 l l L I CY]l.cd R E I e ~ ~mc't.lrc high.llldt. I t l ( > l l 15 11 I1 1. a s i n ~ l e g~rnwdeitle.n-ranyr. i i f v . w that fhc populatio~~).rv then ~ p r r i r lt ( ~ nttlc't qomc .lptatic)~~~. to r r ~ r * r < l v c l l C < < . ~ r n ~ y c r t i c c v c i ~ t. I ~ I c SO irt in . n r Insgr pnp~il.lr~orr l Scmc tcilcl to att. t~lellt~oncd. sprc.34 Evolution and the fossil record ~ c n l r may hc rl~rcLrt ~iiohilc c ~ ~ c clurnunts (vari.~r irripl~ratiorl\Lor r. 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.~vourahlc tnutatiorls. capncnn fi>r n ~ c ~ ~ c r nor t l l e ~ ~ rn 111ob11eor yztnf. c l ~ o u q h grr~etrcnllv. I t i s within such nn.lcctive.. It 111.irurc .itroil 40 rhnt net%. eeprc~allvthnw nllorvinq llc'r1 1 thc ilry3mstn.ilon.~tinr~.]tad uf lin~itcd t t k t : 1 they . . Iuvinp rliverpcd s e o u ~ back is nriyinal qcnc csprrlclorl from uhic-h they arose. ~ r ~ tl~cy c f c r arc tion c-qt. T h c v tcnd tn 1r rubdlr.~r. v . tion. hlclrt tnutat~ons . n fist ~t r~mid and spread ~-cl.s i q kuowti . ~ c l v a tagcuuc muta.25 The t7tc of rrultat~onc ~ u I F thcrr any I~kclihaod of ulorc r ~ p i d+yrmd~n: Zt 1~1crca. The* <pecltq.Thc cnnirnul~itic'.4vcr tcr parricul.id. hcrwccn the pools.I ~ C I I C .it d~tf'crrnttiincs.Irinnot1. thr qpcc~cc. I I X H ~ i~ridrr certnln circun~qt. i ~ l r a ~ ~ .recewve n ~ t ~ t ~ t t ~ o l i \OCCC>~~ICc.111 org~ilic~rl throt~gh rc1. garnodemes.lctcr~ f.vcJ. Hut 111ost t ~ ~ t i p o r l ~ l .c. onc n ~ ~ l ~ .not cvcillv d~ktnbutcdthroughour thr'lr f C < 101 one c f l r c ~ n ~ n \ n ntcl c .111 ntllcts.1 7 ~ sterile wittl the pfirnt srnrk.In altvl. c h a n g n rn \\~/lir-h h nu\ papulations alter doiiiinanct. i t ciiff rcrit clinr~crsm ~\pZl~cl~ msv c a F fr\v grncration~.1y I\c cq1 It <>\ogl I+ L-JI Llr triicroli.lnt.irti~l~lar e n t ~ r l d E l l1 0 ~ oidy in urlall pnrr~lnnnnc ~vithclosc i 11hrt. z t ~ d cv cvrn 'molecular prasitcs').f'ornltd llv ~ r ~ i ~ t .thc h u n ~ ~ n <phcrc.lr wcr.3). 1 . . wl-tich IlIav ctnlv II. n l i 11 vcn7 oticn lut11. recruit5 arc isolate (i. he the finr step toward7 thc frmn.icd\ rt~dl. Fomr prne' Sprecrd of mutufions through ~ 1 1 1 ccxist only ar modificm. plrtclv. ~ TIir\.l.1t and will prnducc chatlyc5 In our or mnre chnr. par~ri arc othcn <c*cttl. 1 ~ e n c l n ~ e d nf lagonrls a ~ d 111 . 5 o m r inutatlt allopatric speciation ( r ~ ~ n c 2.4will Hardy-IWcl~tberg eql~ilibrrutn ~ h l t i ~ d I ~ Ffavnur tlun o f a new and disr~nct 19 ~n specrcx.I? ~ O ~ . I ~ ~ ~ I ~ C ' t ~U C ~ ~ I C IITI I J ~I C A ~ tn ncw or thc.iti~111Iwztll ti]? sr-1lC. rh.~tc corn. raijgc.ct.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.lpid lagcq.AC)II-IIII~I~CC.+IJV ~ ! .itivcly m . 'transposons'. If' J 111 Mutat~rm1% t h c ynrtr:lry hourcc uf v. Hut 111.c.3 ~CIV tCl. hcntab'c A II~IY . Gomc pnrsccqu\ sdcctive ~ d v ~ r l r ~ l p e .1l. ~ mtCr ~ p~ < ~ ' i ] ~ l C l l C " S . rc.1tlol~11~ rc3 l I (31 11n71 ll~tlfl1~t 1 ..itrnt~. P frcvui ci tot irlrn ITI.~d oil ranics rurilc selertrvr advanr~gt.~d~limuyli roon 3rc ~ ~ F .ini:ln madc.~t~vrJ\ %lr>~vlv low.it1 tthco gnmnden>c wlth~rirc.tll~ lLct-cssive niutntlons rprcnd c-nn>p~r.~t~vch h I ) C> . ~ l l units ot ~ C I P L I ~ : I ~k110\1711 . nngin of I I ~ X V tvpc'c ut'orqnn~z.lrc h~ricr. 111 Iitghcr isms t h a t thc rnut~tiunq d n \prcdd .~tl<e:c'.i1 rh t l r ~ brcrc ltirhiy coy i:cIlctIc irlcl I l i z r .rlj~>tlrcr.111 natural popul. i k q ~t \lrnultl 11c n n r ~ i i ~ ~C\I v tl : r c ~ hic.lrc arc able to spre. <. especially ~f occurring in 3 peripheral w i t h ~ n rhc poyu1.lderl intn .1h1~< l I?FO\'C :1111?1gC \ I T C I R I < I F C I ~ ~ ~0 r 5 .lt~oi~. Thcsc. b ~ rxpo\trrc to Holvc~rer.. ly niucla h ~ s h c rt1i. SOIIIC \. cchoofs ut'tirhcr.ll.hut nor co~ilplerely fcjr thnugk sn..111 Junlrultravio1c. (2) ir g a ~ n n d c ~ l ~ e s tlae fringes n i n lnrgcr ~vidccyrr.nr small cluctcr r)! continu. but rhcy .lnt condlccdc .l.nlci1t o r c x p : ~ n \ i uc~r~i t < TrrlctIc nlcjteri.illv addrd to thc rnurnnt crack. Esaralplcs c.~t~vt-ly ~lnickly. ~ n d of \?Istnhlc ~ c t l c t i rt'Ic~i~t'nt\ ~UCIJIJ\. ~ the 6 u n .r 11trr. Thcy rcnd to hc pi~rtlallviwlartd koni nrI1c. t ~ i ~~ m r n c a ~ h . Wlicn the mica ln allclcc r~iav p o ~ ~ t a ~ ~mzitntc l ~ ~ > g i n to tl~c. l t i n r ~ . g t~c is not I~kcly rh. h . ~tn n v h t r e .ire iicutml.t i c l ~ ~ d i n g .tlc str~ic.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. p ~ p ~ ~ l .I r.~ > ~ 13c111g Cut C ~hr)t.iticlrl: tnultigik of spec~esfro111 such periphcrnl 1xolntr.lrrel15.~Clicd311d r. ~ Surli n rnuzint cfft. y r o u p i t ~ g r:~bhitw.~IlC r:cnctbc hn 1. as ill rhr c l a w r I3ai-\t.ulccq Fspcrirner~tt have c h o ~ m that chan_rcs in tcrnpcr.~chthcmrc.~llclc.

irhpIc's . an~logouqto what Mivarr pmpo3cd in Irlc III I\ rniii~nral. AC~WII to A~tErentiatiot~.to thdt tlicv ~pccintt'. wcr. S'iwnuu!: twt. Hcnt c c r*rIt r . 1urruhobjtat5in tllc sartlc gct~cralarea.yu.~l cvolut:c~n.~rjhn. cnmc pnrr o f a pnp~llntlnn tain locnlrzed pop~iIarior~< r. 7'hc yen7 uticn ~ v l ~ ir t fn'ound.nc>n. iic~ct'~~cIt'cl I I I .Irr ill ~s IIlrccdirlg cvclc ttr mcrclv qizi. ~C.I I. r. tllr qvcirs nidv rcc0vc.I 111 tlon\ c?i allr. 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.fTcctivc I ~ ccrt.ll t pnpuiatinn~.il?. . r .icrorr c r r w a nr ti~olo~. and t l ~ r spin qtablc pc~iodq. 1% bccnuvc the folindcn o i t h e pclptilnt~cll~ I l~np!>cnrl-l to lgc.~l. 1)T .~cinnrn.IS ~ 1 evolutionary 1 11t' <I I. I L ]XI> p. . I ~ Y C ciitTcrc.n . ~ ~ t ~ o p I i c . hrcll lost. .. Evidcncc fru. h n ir F l T . !i(>u w r .r occur directional selcctio~i tnw. ! cuidui~ls.1 . ~ l l ~ ' P. IICI\\JC~.YFIIIL~ .~tron. I ! I ~ L = F t h r tii~ilng m.:.prcvcnt< ir I1ybnd\ nlwrt In r. lv Iiwr t~~~-ol~~hincrl in 5-rtilization arc.thc f c u ~ c \vill bc thc ~ I l ~ l t ' xw n i e Icnptli In thc rwxt section.drntvd by hruaks rcprcccnrlns tirirc.~tion can l>c wlcctcd nkqi~i%t.~in.~ll\.c. I lnlanv rwn arras in M hlcll rorr~pctirionor prcThir tl.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.CT.v i r l r l ~ r i itllc rvholc ~pccic'i. knt>wii AF ~t thc prcscrlt tirrlr (Ei~dlcr.~ ~ i i t i n i ~ {~~<~FCLli-~.ISCL-CI . \vli~-ti of ~llccin~iofi ' .~v tnkt plncc.<f i ~ ~ n a t ~ c > t mnv hc .ll~1117~:d' . F I I C ~ nIrlc5 rhr'rr 1 ~ 1 .~rldn~cl ~ .changcv to becnrnc castcr ar harhhcr nr whcn . Thcy m.I 1::. rcc-unlh~natit>n trnpo'i\il7lc .tllnr~ch l t~ niay m .. ~ r r.r~ Jrr ~tcrllc. qcztc fieI k v .lrgc population.\~tn111\ cwcnt~nl for syc.iqscd t l v n ~ i q a ~ rc<ptmsc to A l ~ ~ ~ g .111~equally~ sudden crash.l~ts tllcre arc.lnr>tl<.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.I [illy popula tinn rhnt scrpollse to the cnv1rclnmcnt5 ~ c t ~ l a l avail.. :$. .iri.~q.~ c~ 111'1t8'ly prow c!etnnlcii~n!. only . rhcrcfnrc It 1%well kilown t l i ~ rlatural ~clcction 4 ~ p ~ r ~ ~ i l i g t i . cm\rlnr: ovrr ~ ~ L T .lt~::v tr~rtuYdt i c w bc clC. In: cclpul~tion. I'hyioIn m y gcnc pool thc etTecn nf all t h r f.ilter when thc rnwrniiincnt ~ ~ : J irw rtnall tsvZated p o p ~ i I . I'his n <miplv ~n rvnlut!c7tzap ch.lr are conrinu~llv lIIFCrat-tlll~. and sllc-11 I.. i thc.iiiv biologists brl~c-ve tIrat Inon r l u ~ t a l i o ~.c r ~ ) r II n t.I popI I I ? ~ ~r3ch \i-itll its own sarnplc ot'nrhc total gcnc ulntior nligrgratcq into a 11cw area and. '1 Ilr lash of tlicst.clifkrcncc5 prcncutml .~rnctc\:1 1 ~ .i\.lrthat w n l r k ~ n d f r c n l ~ t i n i ~ m prne f r r q u r n c i c ~ n ir ttirough r.c.ir~lplc thc tot. ~ t ijvidcly s .lval c t r ~ n AISO IFOIJIC pnp~ila.c-jr. ~ n ~ I II c ~ J ~ r rnt. I ~ .nt cllroiiiosnrric ltngths or {>I' ~ r l drcrotnbination.Darwin.~l srlecrion I C stabilizing s~lcction.11ict31~tio11. sonlr of wllich may runrivc.. 2 % norcd p ~ nr few paln IiIay hecemc qurt~ ~ I I ~ I I ~ I C ~ ~ L L . tlic rolnrtloncst phc. ~ t np Genetic drift: gene p o o l s One \~oulcithcrefi.pc. tlit grratcr ~vill tllc* prr\\thil~r!r ot br :. ki11~11117littlr I I b e w In turn t11.lv initl.irk of vnndhorl tndv tile trend) .. the species and notv~ol sel~ction tlation and species formation 35 - b I I I genetic drift.lc. occur. A pnp~il. 1% I ~ ~ Species thus unn be 5a1d t o arlw 111 ":.cic\ H I C ~ C ~ \ br r.lnd ocillr a t a more ar lcr5 cpn5t.~llclcx . c.ill 1~11r1.i. dcnvcd ' o ~r" ronmcnt. c o l n ~ ~ c ~ l l v ..~tciv ollr cxtrvrrlc ir. n d i . thc chrornnsomc lcvcl to m u t n t ~ c ~ n .. Ic)Sfi).1 ui . in n popul.l\cs.iv act a5 foundurr of new cnnipctitinn. dmrn ~.xtrclnC\nrr ~ ~ positively %elected.lcr. \ ~ h t i l \pcci~'$C C ' U I ~ .~pl~ir .ation~ . ~ ~ ~ ~ ccat.I] rct~l.thi\ -i l. ~ I ~ C T IIIJV L ~ c . Thrrc.p~r.lril rarc. rn.~ol. which 1c:lvcs qucticics bcpn to .1~1~1pt~d it1 l ~ IC r:lT a pamcular wt of nllrles. A l t c n ~t~vely disruptive ~ c l c ion n I .1i11.~lirln~n 53tnplc c ~ i .g.~rion. thc ~ r h r r On hand.~lltc.IFnri fnr I:cllllnzvyrwt). hrcu~nesilrlportarlr in tl l c ~ c51n.x p l < t ~ u t r II~IFII. ! I : I .~ll~ cxpnnd OIIIV i-qwr. i. rc.IT a :Ilk: :11< pnpul. I I I C I I ~ I C hccn bur ivipcd 5 t ~ n c c ~ much LTCJ~C~.I fractlun W ~ very rdpid.ltlc>n VJ might bc $ ~ i c c c ~ d ~ ~ l .~tinrl. 1 .hich may cvcn dlfferentiatt. A h tlic tint HII+II. . WI~cre Thc ~ntcnsity f hclcctinn unclcr t h c ~ e o latrcr ell-cum1 1 1 .~ ~ d c r c d f. 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 . i j r is At 1 qrinic ~ ~ t .%~. : r a i Y ~I J I ~ C ~ O ~ L I I . icrtilir. at Ica\t.I hdmcr. c > cut off frtlnt tlic rest h\ . nr cvolutinn i r i .proccsq. ns new rpt. 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 ~ . e l s~lt~11~111 ~d 2nd llk bnttlrncrk' 111 ~ v h ih nluc t~ uf t l ~ r c puterltinl tY115 I. I ~ J I c ~ dh\ivt~t o fit~trtI gcncr. hcpnq to dift'crcntiatc c .Ay.lllv c. allclcr I c n d ~ tscnd (though oitcu it 1s hard to rcc ~ E I L .ly 1 3 ~'st.inti pnrticular kind of sclcction t h ~ tnutu.I l ~ l c n twrthout selection bcirip i nVolvt-tl.lll >rlic poc~l' n i thnt ~pccics. .ll j.11 qcrictic v. fnr ~f rluc ~ p c s l c ~ ir 111'rcArc r ~ t l l I~ r Y ~111 wlilrh i. criciquc of D ~ n v i i ~ . C ~ . sccn an thc fo<\ilrccorcl .l~g. wiJc. a$ will bc cuplatl~ctl.prc-ad and well adapted to i n cnrrlr r c d . But thcrc are ntlicr k111tIx n f iqrhnrlnn \~.

I b j .lt 111..ir tire citrcc hnd brcorne poli ( I I I ~ of Kt= .il. o f .lt.lvr hrr-11 3ffl.In iilrnn61c lilnitation.r.11 tllcy have. 111nvrver. for ~ g i \ t r. Prrniln.. r1nt11r.lrl i ~ ~7-1 l . l-lcvtd~~tror~ the. ~ i c t ~ ~ ~The c.. ~ ~ l i i c k rctlirrvcl. h t ~ rJ.I few grncc .\L. I~l.~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. cnt WRY< 0 f F U N 1 V I I I ~ I < . ~ ~ u t t t c d sbovc.. tll.~ccour ro br d ~ r k .the nloths In o : g v l l t .111~ier : rapidlv. ~ t ~die~other I1'L 1 ~ ol : <I uJcre d.r~-td i t . whntcvcr rcpmdiirrir~ca).!~<illltl fouu\.~nsparciicy. ~ n ~pc'c~att"ni~.ttcrc. drnuglit 11-1 1077.s t ~ .idvails.11i n o gcr~cml rcnr~r.t h.llet prcrrurl cnlourc-d 50 tf1.y pnrtc h.)I-. 1 1 6 4 n I 8 .conlb~nntionr. atld I..wk rsut~kr.i hccatnr. ~ . ~ t T l l ~ * b l ! \ <<> t h dein.aryt. Ilc\t fi~nctiur~:i! Y F ~ C I I I t l ~ l tvrl trlr. Hcfcrrc.1\ 11t1 T ~ .~rc rht=\+ aft-rtrti htxrc. I11 <pltc.lrd c ~ ~ l uh. . t r rr-(l~iir~-fw r r chnnwc r inihdtc ducrivcly and tropli~callv. ~ 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 ~ . ~ ~ '. produl-e fewer .. 11i.hccn hra~ll..ip FHCII i n d u ~ t r ~ nrlr i c l ~ t r ~ <xvitliir~ ihc vcrv qhort onc rcs~iltof nnturnl ~clcrtiun. Furrl-rer~nore.lctcri.I tnlohirc'c cyt.1.thr \clectivr ~ d v a r l .lt Gn1apnyo:nc f i r t c l ~ ~ C .0 I hc .uilplc c-nnccnir chc r.ir~ctcn..r\~.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.ltcr\l\-tc<t<ttllit c. + ' ~ V ~ Z = ~ I ) 1 1 ~ ( ~ rtu<li~ri I3~nvan. l n l ~~g A rnnrc rcccnt cu.I I ) N A t!I. 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 .T1-~iri s an a t t n c r ~ v econcept 2nd in con! I rcsprcts still ~ ~ t e f ~It l1195.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. O n c cpcc7r.III I l c c . 'blltl c .1t prctl. l l ! l t ' ~ l T 1 iiLr1IEcro1Ix C O ~ ~ E ~ ~ . ~cnlrcvc~lutinn\ l ~ o u l i lt l t b t bc rcvcrrll~lc. ccuirccq.'Iluuted enerp' Is~ldpct'1s not lnvnnahlv vnlid.IT . i s srcrl . 111 tflc cu.turlri. hpcs.lnrt.ct. l [ l ' [ < I <' I.rcnr depths. two kindc of .. it had brco~llc. north of E~ylarld.lll '10 vcnn I3t1t only .lyt'r pnrtic.IVC JIW ~v01vc'd ICII\ dr>uljlr. t l l o ~1. d11d r e p i .utr~~5~\.r. ~ ni o i ~ s and l t spectec i l f i n n t l ~ krirr\vn t c i h. rrrpcct. ~ i krllcd.Ocll~.u.ll susf~u r ) ) l ~Irclll3r I. will tcnd tn nl.1 inhabitir~t: thtv I~c~l~~-n-rnvc=rr.lr~d rclcctivc .~pt Such ~ r ~ n t u i ~ ~ i c aptlvqiological ~ d ~ ~ p t ..~l ~ tree$ l ~ a d t m i i b tbf a made of mlclrc (Clisptcr 1 1 ) .(3 t*c\ 1P ) c. Ilavc very Illell fecundity s that p a t iiu~iibcrc 11!.m-hich brill9 11q111 trr n c11~1t-p I ~ I I L .ili~ npr? which \vt. hut otll~.J I ~ C I I 2rr w r y scnvtlvr to other dlrcction\ than thc t n u ~ r r ~ l oig ~ l <-.r Iutccl .lrrcn the pes very ~ ~ r n p land ~t vccrnx rathcr t h a t ~~nptilnhur~ .thcv II. ltp!r<: 17 { tinn docs riot work like . ~ t r ~ p i ~ t ~ 'I .11 ~\. the end o f thc ccrl.itor< 1vo~1lJ lcw cari1y scc thc~ fnrc.lnr\~n' ~aic)tli\ hccn r. \rlcctirb~~ .r t ~ z .thcrrtbre not very r. Ir~dtvldt~nl'i b mdv Fc>llolv~rlg 3c.rlil t p.off<pnilg. i r t t d ~ thElt~IICTL. tlic hest ~va~lahlc ?.~nt. !:~.w JlC'Jr WLTC t J 1 ~ 1 1 t l ~ c dirk-cnlni~rcd tbr~iis. S~ncc. 1 1 1 t I 07 r:.l1 o f ~nlnll5ccil.< . cx7miny thc b1nckent... ~ s in\n ~ ~ no tlie vi. tllct.lrer.lt .~tit (Stcan~s. I 1vliic1-i opt.c-tcd b y arc.lrc tions .d Trilnhitc~ha1.In cHg.li~ly r i l n b ~ t u\Ircrc.irrilc~iiizii~q 1 t thc crlvirw.c ovcrcnrnc thiq di\advnri t. ~ t i r.dnIc.lctcrr.~~II'~LLII although calcite hac the urrtur o f tr.I . uiinblc r ~ir 3~ h secrete ni.r b~-.c tir~iiudn.b. l l ~ ~ ~ l t ! r r . Soiiic 311 p0~viZ3l~. of tlle~c C I I C I .l\v tr>uiidr.thc and cccoiid h y r n a t l y rngctlirj~l\rnuc!ifir~tic~rl~t h p l r v.lre at anv t m c .llt Ic' rntciw in thc sl~.implc.t\ .l~~t.r tlve nhilitics for spccles F I I W I V . I t work5 likc 3 which rhcv find thr~nscl~cs. crysr:~lI.d .t..tll~ 1 IIL.h i e c h ~ l g r t niiiicr.rpacc. I1111 nild tinit wcrc nhlc to \urc'~uc.1c.r ~ .tSdin! <>i ~lm-pI~olol.1111plc\ O L I ~ ~ tuikcrcr'.!~\'C n 1-iokcd our bv ~ r ~ . Whdt src knnwil .Il \I of t11c.rluiirl. lI>lL.I I ~ I ~ C h:~ 2nd bird? 1~1ths r r c r n ~ ~ . l't*rl~.I* riiotl~\ 1 l . p r o d ~ ~ c c I idcal Kcno.o d ~ ~ f iOr1 3 l l l l l l f ~ ~ l l t ~ . 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.lptcd to tlrc~runviruzlrrlcrlt rcTrc1 tinle<p.elv in II.!>mad ~ n d ~tratc+c-il ~ a v c bucn Jirtm~q~isl~erl. t fli <.~rl:-colirllrt.lsy ro catcgorizc adnprivc <tr.u 1% . 1t1 thc c i r c n i n ~ t ~ n Ic c ~ ll~~lI~lrr! . thrn. ~ i i r i h ~ c .mal's cuticle.~lrnc%l. thr. . I t 1. ~~~ I O~~~ ~ ic ~ .idv.lsi nl1z.ll yrcrn ic ~ h d itq n Itiyj:18 I l l TI><+ I 1 1 1 1 \ .lk~h t r.. K . thc Industnnl terial which tnay ~tsclf lunliiny.lllllC c F moth5 ni1 tlie d.d ~ t Hut .~ u r v i v potrnt1. JL ic ~ h rc5t of t l ~ c11' . 1 Ii~rrl:.71iId CI)PC u ~ l t htlltlfe thc other harld.but a1urn:ll pop1111 n~ I.tmtcgr~ II 4 ~ n N o r.I ~ W I ~ ~ .tlic coliccpl of rli iller c11. ~ ~ tun. 'Naturnl ~ E c c . hdjrlrc r l < I I t !:CTII'. rhc av. alqn ad.l 'inrFil\tn. arc d ~ t l k ~ l ~ ~ I .criticized. I \ Il.Is 3 pnman7 ~ g c ~irl trvu.1977).tud .36 Evolution and the fossil record t + t > n i ~ t T I I .~tnc-c.~ O H I ~ F 1113)' nc( \I thc nf i i.rrb : ~ h u n J ~ t ~ t . J S O Twh!~ ~ ~ n a l l .I.~ilnhilicv uffqpring arc prudu~cd .we t bc compronuwc amonpt rc>rtyv. s111ccso111c Fpecles have both high fccr~n I !l%I'.lt Scltction.cr.pcri 111'it Ii. psnducl ng douhlc i i i l ~ y \ on L I I ~ Iti11111 7 difr't.~ndsand bascd upon Fcnctlc and phyic.. n l v i r ~ g ~iw :r pcnrt-r variety o t ' g c ~ i c ~ r-ton.I< 'r-. 1 1 11\ I It. rclyiilg upnn tllcit supenor reproiiul ~ O' I~ou~c\. It i ~ L E ~ I OIt I doc\ i11)t. hn~vc-\.. but i n mnnv c.~rir-tie< ~ ~ ~ c t ~ ~ i i t110 nmore than l"0.ll . ' I ! . III~\ ~l T h u ~ .11 ni. 111 this.. ~t 15 lefs dity nlld s~rpcriorcompct~nvrrlcw. f i r 1fgl:httrnvc-llitlg t l ~ r n ~iti rn l211. Crorn p l . Biyycr rccdq 5hort-livcd. > t I l ~ * Il For csarnplc.. < !KC1 'I'11~y trollrri~:ditTcrcnt I-11nr. wrrv irir. ( ) t i < * Ill $ y..?\I W t r g x t l i kji1d1 u r ~ c l r ~ Jtr.ikd. d LIP' E ' J i ' r ~ O~'TEIC' p ~ l p l t I .ill por~rbl l i ~ < ~ 1 1 ? 1 .rre more eas~ly dirndvnr~tngefor 211 nybc..l\c o f h.IF Jncob (1'1771 hag well s.tdapt~: t I r i l l ~ c .rr~g cnvlronmcnt. Tclp1c. wc~odlr ~ t i *nr.tlcclrction had bet-n ha$ 'fnrccd' t l i ~ .c~lcr or1 \ttrall 5ccd5. the lenrer of . . cnurgy budget. r<>171 tngc tni rhc grhnci\ clcar. I>.11~ \pCck. I .lnJ thr. II I . l t i . 4 >plr\ o f t i . o mbhitr o r .

lt onc .c r n ~ c r r d h a w < rjwtaru.I u ~ l ' ~ s~lwlval.urt\ ( c . y .Trnrlles I S Dovcr'r (li)H3) tl~enw ii~olcnf r ~ l i ~ ~ l i t1.4 r \ . It is hnrrd on tJ.fr .)r i t illrltvr r h . .t n Killiur. Tire gent. ~ n d 11:llqle~r.r h.und thc' gennlnc.1t Ic. ~ < t s . 1 1 1 .lrl c \ ~ c i ~ t ~d .lt-~ants h n u l i lhc cliini~ ~ ~.~tdu curl1tl:ln 111o\t n1~irnni. rspvi-idly ~ i 1 .Ilr < c r ~ r \ r ( ~ d~.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." d c \ ~t 1 i ? 11- 1' '1 .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 .i pll. ~ L S W diKbc 'ucrf~il' I I o I ~ ~ I L .11 qulrc ~ ~ ~ ~ cx~ri. A n u n ~ b c r t1111c.111dthe uze or cuiiiplcsrty ( ~ f c l l nri~rti~~l. 5111ic thcrc .ire trcbnucrn.lctrve ~ v > t c ofn~ c l ~ ccot~trol.1.111d ~c I 1111~lcnt. Suc.w t l i v I 11 I I I .t c ~inl.rrr~i..lnil IiCr~rc nl+r)+tt~:~clc) thr.li~iilic-.voIzrric~n tl~con. hv tlic redonbra blc pow-cr of n.l L ) S 2 ) .<. Ic\*cl. I T ~ TIIIC ' s i ~ t.p ~ ~ ~ l i tllcurv.ltlrc d ~ f f r r c t ~ c c ~ l ~ tn~cdert~l. p c ~ p ~ ~ l .ltion ~ ~ c ~ tah e r i i e c 1-rtarcl\rill d~m lior ril <t. organrz.~ttlic rhe I\~IL.ltnr>ilrlt t9f crrccrs br appcrir if hv rhanrr alont.tiicc 13fthc tlir ' .h c lc\.is multi.rrtlirr than p i h i O ~ I CM ~ ~ c u \ c ~ .lri.~v pcrlrap~allo\v :h fustlicr ricvcli)pn>cnt of thi\ 11..tK.r o rsrcliirllccs cvict too.irl h n ~ n qct.~r ~.lrion <. tlic' .15 brcn c:illcd ' r c ~ l ~ ~ t ~ c i .conccrbtr. o f ' ~ i ~ r molcc~ilnrn i c c h a n ~ m i c . l ~ cd Tlic lii. rr. which ~t tint \igllt ~ p p c ~ n hc 'concerted evolution'.lrq thcm.. ! zhri nrrd tbt C O ~ ~ pror(~i~i\.. I > .~tu+) .I L .cc. h Molc~-~tl. c ) r g n t ~ l ~ t ~ ~cr 5lt1ll F.~~~<c mnlrculnr n~cchn111~1115 of w~tllin tlir llllnl r l o s t l ~ rclatcd prntrlii\ .ire irndrnrood..due r o fiua tlon of n~~~rnciorlqt l i ~ a a popu1.tc.lop~nc.< T I t11.111 l ~ ~ t r ~ t parti rof .. For the Irlnrrlcnt. n r i .tilrii out to hc ~ ~ s c fs ~some u l i :ral gr~rcticc makc\ riw of htccdrilr: rspencatr-prv.IF vct firll\~ c 1thl. il.ltinli slio~ild much lower I ~ ~ T .ncr.11 mult~plc indivichlal membrr+ of a mu1tigcnc E~~iiilv sl>ow.4 n~iiltlsrnt-hlrl~l\.lte to l of irn.~ttt ~IIC'IC O ~ ~ C U ~ V .I cIvii. ~ trhc f:cc~~omt.multigcne E7.13~ 5i1cli tlncwcr~tralgc.rchpc. . ~ h ~ l ut ~ ..rloc: n o 11arn1 hut s t ~ v sIn thc Sytcrn . t 'hnnnh11' v.(.h ( gnrr ur loss of !\ t ~ a n l u t n i l r gcnc.loyrri. nf ir+1ni7c.m~~cal pntli\\b.Iry fifrlr t h ~inniiitcn.I~ ~.c undentailtiin~th.IS .lr t.trc fcr n fclective advnntnjic arc smilable to ljt' ~ .rred.lvr tl7. I # ' rl11t11 17 t.l1 i1 1 1 1 ~ l~~ i ~nild hc euprcted from natural ~electie)!~.nrr\cn. ~ .irt~ic . I3c ncccFr.i rmn t~ ~t~\ wi ~ ( 1 4 1 1 .ir ~ q at t Icvc.i c t ~ Wl tnnrc i q klio\\+t)of their iiirl~tiot7c.c l f rhc cell. i n t' ur )unkS DNA.ln\.Ill 11fc i 1 1 .trc\ri.Darwin. Thir hc c-nllc molccl~lardrive. C tly 1t.i r .lvc. .turch rrf rht.~trJ ~ t~c ~-cr:rilnti<~i~.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 . Such rnult~plc. tult nwht ~ t ' t i i rcxccq\ 1JN.lar~un. T h l I~F 3 d ~ h ~ t d b 1ptl111t.ct<. \ i p p ~ ~ lvi I l ~ II~ lt ~ t . .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. cl~.1.l11r4ing \ ~c. I~..i<rIi. It 15 pt~relt.iIly . to .c~rnpkc. for rsaniplc. rlioqe ieiv tll.vc. r i ~ to . cdn t. . t'crcncrc chould qprrad rIrroug1-1tlit.il ilnirh tvirliln thC . the species and natural selection 'chiclcic-d*compoticmt.' 11 m m 8. i r wnuld scmm apprn.atucll k r l r c t ~ o n . ~ .111d . t n l ~ l t i y c r ~ c f.ctro11.~ J~ithcrrtrclu\ir. ion lplll~r ! #I/. ul11.nc. o l ~ l r i ~TIII~ r n p i ) ~ ~ . ~ t.7uYrl1at doc.itrc 1tl~o11 ~ J C I I T C the undcrhtoud.~rc.~rlti.I ~ u ~ r l ~ l c r i n 11 11. Thc iiidrvrdu. I S arc rcvnlut~oai..>I-ned in t h r t n ~ ~thc. It iiiny cvrn.~.r ~ t :. cular drive.411 snll bc fir pctlome.e.~pl.c! N r w t h a t trn\.1rnrliant of I nforrrr.Li~~~ilics'. < ~ .I< ptlrTlic 'ninplilicd' . 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 .cl cnt' i~ t d T ~ I I + Y + ~.~cic~t~. iq..nt v c ~ <h. ' ~ 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~.Cntnhnnn 'rxplos~on' o f rncLa.it I ~ I I ~ C cuprrs ol' chc w n c gc-rlc. ~leutr.1(4v ~ r v i i c ~ t a ~ ~ l ~ 15 l i challt-ryec tn 11co-13nnvi1i1. ~ ~ l . 311d 111111n lr~~~ ~jrnilnrity within a spccles.bblr.I ikrn thnt hc.t. . tC1c rht.1rgclvIiurno~cnizcd In n ph+nnnienui~krlon-n a5 itr lclv IjNA.rof cvidcirt ~ . +n tll.I< tin111ch.roiltrolIinq clitkrcllt II oleculur genetics and evolution I.inccr. i I l i ~ ' .< thiq 1. n1.cf t r ~ sri.ni. 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 .quch i ~ i t i l t i ~ t .r . [t .~n-vriit~rh morc 13Nh o i i r r G n n t l ~ ~ n g do w ~ t h ro naturdl qr1t.ei~ c c n d a n r i l nrg=~..lt~cl~ t rrthrnviw ~ r ~ l ~hcr t ~ct-tionr gncrmc hv rmiirpnciti on) (nn oprr.rgcticc.uo111tiot1\ttorkr nt thc hn\ liltcr. nftcr ii~i~c-11v o r Il~rlonaryrl~vc.. dn? Sor~lcf i ~ ~ ~ c .lt ~ I I C qcncr .e l .tr ycrletrrs..gcr~cttc. ~ n t lulw)i~l i o ~ v gcrw-~~tlc 111. Accordtnq tu Uovrr t t n ~ to 1s . 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~. in \VJYP not .ll. ~ I C T ~tl~rw I ?r W .mr.c a5pcctq (1 ')7C))'ncutl-rh rhcory of e ~ .rI~i!.r ' ~ ~ o n ..~r<clv . c ca1.rnJ . l l r ~ t . F v u y r at' rcpcatcd cctlurrlcrs or .t h ~ p o p u l ~ t l o n ~ irluclr niorc dl\ clxc t h ~ \~'Oiilrl ~ t ~ ) t arc r~ irrllrtir . cuncept.+ tllv tntlst iinport. be pnman.1tcnalI \ 5tmcturcd complcx ftrnctit>ns a i d intcmr. dl(vnn.I t ~ ~ i t .jrlt.i'< >I7ui! lctr zintIcr.hnon\+ccrn rn i ~ c i n d ~ v n ~ . I ~ iIc ~~IrIt ~ ~ n l l yhrgl~d e ~ r c .o. c . ~ I I Y C L ~ ~ A W O ~ C E c. u11 ~ i i u l t r ~ c t ihlriilicq mav.1n f i v ~ n ~ r .ir~~i~r. I t had 11rdcud I>ccti \uyycstc=d rh.

genes come illto opcrariot~ it1 I .~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:. dcmcnn arc rccnnlK q ~ c . r p r o d ~ ~ of d n d how they can bc ~nodificd. snd w. srru~ work 011 aiid iiitt.T~ in coil. 1.~r-r-i~lt.lnf of C ' I c o f a segolrnt.qucnrl~ll ctlicct oil p .~r 4irtt. hlr>Yc. .t orljc r s hlncrions. before ttrtiIizntto11.~civ.~tinl~ ~llj!:I~r .1rt\. IS Iikclv ti.i L-oncertcclp-irtem of t ~ s n .lop?.~ tivrlv tliic nc.lnx.. r ~ . I ~ :~~.nt. 'tic-~~rral' o r t f ~ c'inolcc-ulnr drivr' . ~ r~ MCIICCIII.thir~r: p l . Snmc qcrlec (1313 tmlal . n ~ ~ t v .iit'hc C L ~ I ~ O I C .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. i t l l r . Fmt. ~ Y O ~ U T ~ C I I I Jc l C . ~ l . .l~ l ~ + c l h ' .I\c . including prii-n~tki llatLlrc d nii W in tr r ~1011r~ 1 1 d liio~v [I\LI~-II .rv~*s.~rrcttbn<.lllhr eEect4 h5h the antero-posrenor .~J.~stcd with t:cr~ctic drift.c.TAICCt l q~irstiurlo f llow ni~icfi IOX4. corrtrnllctl IIL \pr.lL ~tr. rhrcr ~ ~ i > u t11'rr-~l..on during development l.cvnlut~nn o Micme! \V.~l rcpreelciltel Gene ragulati.. \ C C I I I I I ~ ~ ~ I l~ C Cc~ c ~ it ~ mct~7n. I L ~ CIC !:r. ~ \ C I C C ~ ~ V1 C npcratc in tllc snme way. ~ d a p t i \ . h ~ n c .\tfi.t'ci-.d IIV non-Mcndrli. Thcy arc.t.mcntc 111 the hody. ~ ~ l n t h c r ~ l c Ilc w.r.~l~rlct.3 thcnncc F[:I-IOM~!J.~ ~ l t '.iv \vcll prrlvc to bc rhc third seyinrntnl and crpeclnlly hntneotic) c o i i t ~ i i\ ~sl-tun .cftrig . rllc to They 110.il \pcclEll In( ch.a~rrplc. Tlic-sctlircc group^ of penes ~ c nil easlt tltlirr 11.~ d~ I to T. cnn trolling stnicturc. h u r n n n ~ Hoinrohns-coiit:\i~iii~p .IV. ~ ~ lcvcl iliclucic nnclcnt.i~rlnl>intnt v can can be ir~tcyrntcd ~ r lvith scnctlc and n~olcclilnri1. 1')'$(1).and tlicy alrvny F~ii~ctiir~i wpneilttl ~Eetail111 th d~ in r31e outce n ~ ~ ~ iA V C t hc. .-I homeotic g?ilr corr~plcs . 1 1 act!r.n st. I I C ~ ~ ~ T ~ I H C F II nco-nanvinicrn.il i l l ~ c n i ~ n l l u ~ nhnrn n n t ef homeotic gcnps: tl~ekecorltrol the unlcluc idcrlt~n~ (Jf cq11. give a i d tiley define the numbrr drlrl pocltlon of <cy. prrqcnt 111 nnn-rrg. %?TI nnd m. at l r n ~ 5 0 0 Ma old. l o~y wh~l the cltl~c utltrr rrcrwc.5.f 0%).lptc*il ti) ttvc.ind at cvt ruclr I l ~ ) n ~ n r r .ly it~glc'cil ~OIIC mc.I tnult~yrtlc f > t r ~ l l v.~r Thew r trar?$pnwix..~r dnvc rs non~c c. F ~ C ' C I ~t ( > t h ~ ~ I C Y T.icI.nt rtacc. .rs 17ci-11 ~ r i \ c .~du)r t~ cvF:.17irl c+pu.r ir ~ mid lilnd.~n 111:iv pmducl I~c'n vcw Thuk \ V ~ I ! ( .citl. Scc~md.~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. ~ t = o n t o lcupplic~rhc tirnc dir~\cil<ion.~n~~nt. In kt% .irl-r I C t h It-vcl octhc orprusrn I w t is ncm-nday~ rie~llvd ~ v thost. 7cc. u i c l ~ i ~ l i l ~l:u t ~ o i lr i r t opcr.looks I ~ k e T h c ~ c drive h:~vc.' TI~crc.1 prrdete~-r~?ined L WI pnch. ll il .c. I~avr. l l \ ~ I ~ c t is otrL~rfi~onall\r i ~~ under. rnoJe\ W ~ I tal rf. t t i o n .h o w nc\v body plan? origlrlltl on ~ ~ ~ l t h51% ll crror in this prticcs. ~ltlc\not 1 t .111d \vl~. i ~ \ . ~ i l c l ~ ~ d i ~ ~ c : e d .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.c~ can t-rrol.r r a f l ~ r v c n over l I-. ~ c i ~ ~ i u m:1v c . i l pmucw-c.ltld ni.Jjni~.~\ I'he c l i . cxyrcccion {lirxhcr genes nt n <ubserlut.gct~eticicts ~ n t r o n n pJrtrrfiE l clmcntcd phyla .lllv l . TI~IIE o gcne ~ ' i t l iaddp r w i t ~ r m .in he esr. in thc cvolunon nf dc\~clnpnicnr.~t~or~v c n c l v . J I I L ? c:1111.~l.~ % varint~?i n ~ e r t c b r ~poupr.~ll\.~inii Molel-ul.I[ t l ~ u o l c c .tiid cuc.mi rlc.inragc.. ut' n body axis.it ttlc t c ~ c l tlic g c n n n ~ I t .?rd to fcc Ilnw z. L nilt D I . ul~dcrctcloc ? c t ) n ~ t ~ r ~tt~ vn l ~ .lt u ~ of ~ \t.hut I I I < h.lcctivc . but hnc nnw her11 shnwii to hc hnr~~nlnq+:tlu$ tl1u cuu ~ of thc \ ~ ~ r i .\y i r l wliich qprci~q~ ~ 1 1 1 qiqinlc produccd by thc qcpnenmtmon grnrr.. and nny a r v yalaeontc.w ~txL>rnm.~y a h Itc cr>r~tr. ~ l i n.. ~ t r t l Ii~nt:-tr.~ltl~ng p PTC>CC\SPF 'tlir r.and cvidcntly thcy d o riot nlw. w l i ~ c h turn Iinvc t l i e ~ rc%ct oil other griier. ge~iec nlwt be vrr I ~ ~ r t h k Zi i .c m. 1 1 7 ~n n r rleurlopn~cnIicd oil. Clearly thr hnrncnbox ccv~ccpth n ~ n c v . PI I1. cincc they nrr found c m l.li tisrd varintrts.I r I tlllt~ ~TZSI.1 v s r i ~ n t . maternal genes crtabit1 f du q I T ~ ~ ~ ~ ~ ~ ~ ~ I .for rc.~ . 6nt dircovercd in homcotic gene? 111 IJlnco~rlriE~i \olllllj.i~~~rl. lio\i~it I + ~011strii~ . T h e rhrrd ~ c l ~ c'oii<ict\ 111 n~liiilw Such ip tin11 dt*tjnr*\ hlolfipc.I p l . at h i n I ~ > ! ~ c .arc.ire rhnt E~CILWI . hon.'< i ~ p c ~ ~.lllOll In .38 Evolution and the Fossil record X I I C ~ h~~n7otit. varic. known 35 the hornenbox.~pt. It 15 1XO bare palm Iunc rvolutlcm N r ~ t l l c ru tlri. fnr 2nd t. s r c l ~ e i i c ~ pre~umahlyc n d e for proteiilc tli. A tr~ o f mirur. In thtr ctrtitc\' k01~1 Tll cr one unc grii~.ir clnvr c.lt frr>111tlic t ted ir nni$r.n~ causc. wn pi1 17' +4 111 .15 ~ ~ i~c d lt f l l l l . .llrc c o l ~ r r n h the. it clucq FII .in . ~ ~ r i vmolt-cuI.c. c+liw cd\pcr ~ . ~ r i c k.n q . nlllK.n dlipE~cntrw g i l e n t . l ~ ~1C.llil<rn ( l c i ~ c l o hont n frrtl11 l~~ zcd I>ittcd into ncw dcvdoptncntal n~.?SCL' I C ~ I I C ~ C~ C C J L ~ of ( .ipifiIi~glily cnnscnrative D N A wcltierlcr (or ~ t l o t ~ l ' radiation! CRllt .~ciiincc* (Rnh cted to r 3 c r y .lzc A 'c. It qecrnc t h ~ three high1 t tdkr Il1JI1\ con~crvativr h o l ~ ~ r n h o gcr.1 cprclc\ lulr clrv1r.!irrl the ~1r1qi11 ~ I Z W r p u i T~ c-.. aftr-r thc c ~ 15 Kprtiliz~d.~llvcon li-rr r \r. re~ultinp.~ ~ i. 'l'lie 11t~uenhnu w.lnIc rcpilator). Tllc cr~cl rccult 15 n fi~llv drvt.ITICC thcrcfrorn. h u t n l ~ o ti vcrtrhmtcs.it .CL~CC i e q t i e r ~ ~ v e to j i ~ ~ r w r x e in wclurnct.t.lopcd orptmcrrl 111 \vhicl~"tlirr~-ir .1t.)lid d u r < ~ 1 .ln thlis f x J the egg. ~ c c v c y t l i ~ n c . ~ n d C tiyr~p1 rnnstnnt in vzc a n d &ape.ti11 11. i segmentatinn s.itorvgencq rhnt We linvc alreadv s r r n that palaco~rtology.h~ate to tll.nrrcr In I breed in^ p~up111~t1on. r uulil hc Ictli. o r r~~rrlt-d .~ r ~ 1XYC.

:ln prtxl~rcr'rhrouyll <In\\.cvel-. thc be rrt. ar~inlal K and its ct~r'lronmcnt.~nrigr h l q 111 riiitld.~phf~xl qcqzictice ..\ * ~ d c nd t t r r ~ ~ athcn .-1111 tllc ~ C F I I rrc0rt1 . hc~u.ith tlrc .e bc. I t wrmr to I r rhc o 7 ltlntron of thc ~vholc pnptilnrlm~ ( a ~ t n ~ c r l c ~ f l q ) .il.5 may Ilnvc chalrgrtl ~ o ~ I-crtnidv I:\\v111 bc C O I I F ~ I ~ C Ti C>~turn. I I I ( .1t spcrtr.isn~~:c.44.~nc'Hr. ~ l ch a n ~ e ~and hence of the ch:lr:~ctcr~ thc er~tirc o p t ~ l ~ t ~ o u . conccrn? C V ~ I L ~I I C I I I .:llcd h. l i .itScrltlrrn.ind cnm.ind ~ ~. w c i.ly hc n~njrsricallvl~nfoldiilg.~ ~r r td l>t~rich w t ~ r k i ~ i y q i ~ i t r %Ire ..1~rnl~tior1~1 0 1 :n d111n131~ 011 1 1 1 1r done know11 ha* . F c l ~ o l ~ ~ t i uch. Tllerc i. ~ pIiI dI ~ I Cof CVO~ I to chow 3 pndtial ~ n o d i f i c n t i ~ n tlrrougli tirnc.\vh. rccclvins nluclh .1h1t. I t carrlr tn h r 1 1 hccn vcnr r.I I rii klther hand.q S). :Int L T ~ E I ~ ~ I D I I .lrlccq rhnllgc tlliicly rnnnncs. r > r ~ t n l r ~ ~ i.~ . counrn~ .I scntbnclst ~..r11riorrr1. 111 1 1 1 ~ 1 1 1~ I I C ~ . t c .~hl~~hcd t h c I ~ i ~ t l of l i thro11qI1 ~ I I J I P itit: proccsse? bv wf~icliqpccler d1I1.ii n f c ~ ~ n r i i i a u v ccc i\ a Eequcncc n l sr.<'etaricrl tl~rnuqhtirnc 111 rolne rnrt. could anus .ulut~on t . . d ~..111 rvcn.1 qcclnccl to bc in t u n r \i.thc pcrcrption. I W C ] I . T n xonte d c p c c t111s tnav indccd be due to ~n~ol-nplercner~ ~ t ~ bins 111 the fockil recurd. ttmcqp. I ) ~ n v i n . F ld VC "I Ihnr ~ p e r to c ~I L Y V nrtc.d gcnctic.i n a l t e r i ~ a t i v ~ rn : hr. A l t h c ~ ~ tlic~ v1t.ry few < nut prclirrd . rcg~il.C C I I I ~ ~ I ~ L I t n I mi$ < C T ~ C Fof y : ~ J r d tcnnedia rc fortnc . l~trlr e v u l u i ~ c ~ r ~ mnditicatii311 nt all. and chpcrra!ly tlic rr. that p a l .rliiicth~nghndarr1cnt.11 prlhthat the rciclrri r n n b t takcn d t CIL-C V..11spcciec . .m) pruIS by 13ann1i. n singlr. ~ n d Jcsccrl~l. qtclcli.~ti(I I C T'IICSC t l I I~ ~ ~ ( ~ l ~ ~ t l and ~ x r v wlnc populntlon. 111 111 the hlldlw~np tcsr Ir 1.. .tr.r\(. tht-rc wrrc \*c. ~ Cthat evaIu t~oriaryc h n i ~ ? .hut 111 nt1ii. .guidc the rock rccnrd.till ~ V L = E I L I C C C \ F ~ V I I fnrni. 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 .w. E JCSCCIIC I ~:l\. (5pc:-int~on) 7 I)\' rr.npapl~ical rdngc.ntcd g~01uycdIFvhy no rrlrprc than J cl11g1:lc procrcw5 of rlntur~lw!cct~oila? onfinally ~nt.t. .1r. a$ dlcl Cartrr (1954). .lcclnlens rntcnntldlatt.111dor~ r >rlovcr long prn. ~ . r t 2nd I II+ '? .lhnll of tl~rh .r brertling ovrhr 3 r l k7 lf <cncr.r~Iln* ~ ~ i ~ d c r ~ f o nl d q t i t r ~ l *1l1. -l-li lr I:.iC)i. t In J U S ~ h i ~ x v . .Fossil record and modes af evolution 121 I I I F I ~ ~ I ~ ~ l t z t tS C\ICY.\nd .that the saritv of ti)*~11 p t of a7 11ad : u n n r ~rCvt.rp~rl. 39 the rl. p. in mo~plroro.bct\~c<'ll ! rhc ~ I ~ T ~ I ~ I I F C ~ ~ ~ I C ~ CI irliainp~ttcrt . It srcrns a rcast~t~ahlc conccpr. glcnl~rcl :1'..tpr~vrr. Thir 11-todrl of ~ J ~ u .CI. Yct. .~ t s ~ d c h n ~ ~ c h i n k i n s hnd bccn Jorrlinatccl by the idc.l~ld~t rilnc.C ~ ~ Unt'ortur~atcTv.ccn~ .11 tn tcll r l s ahnut Ilhccr mrioli\ I ~ C I I\\'id1 b r cvplnred 111 pcatcr P~ the nsturc...)r ~. ~ l a ~ r ~ r i outret. 2 ~ 1 t hthc clrlpns o f .1tlc~1 T ~ t m l l ~ ~ r i lti1111 \ilolsld ~ I ~ V C J ~large niitnhcn. snd noulri ~ .A l t h c ~ z ~ l ~ . 311 Iliipnrrant c : ~ kt. 'intions JTC' CIIIC t l t ' c 1 1 ~ 'I T I O S ~~ .l~c$ which bvcrr cup14 divrnlty .IIDCOIISOIcit~dcrstond. 1953).cntitc-rln~. .. l~rrwrvcr.lo\r.ltiolls may nr~th~nodific~cion .trdl.ir ~iitl>rrtit I T I ~ G 11c-v.rtr.it p t t ~ c n both pllylctic c\.fill1 confim3nrior1 of how tliey 1 3 3 ~ ~ dct~1:111y npcr::111ri cvnlunon art. M a c r r ~ r v t ~ l u t ~ con . ueunlly rhe rntirt anccl.\ ic ud. c of p hlrl species..yc5 to lrilcrocvolutivn.ir?l: Itiic. many of dw clarsic.ir~d mart p . i t n p t m .. I 11ariqr\ t l ~ a t. ~ t c d h r o ~ ~ gtlnie h a $ bccn clrarly cccn Sirnprnn t li Of crliinl Ilnpolr.rhcorv a n d p. rncthodlcal c h a n i y o f tlrc rvlrc>lc yeiic pool ~l: :l~rrocvrolut~ot~ rctcr$ ti) s n ~ ~ l l .llncontnIoyc'al d ~ t a n in . \i~cIi adaptive 1 C stuocl up well to mt-~dCni ~ i l n l y s iS m p n i i .'rciitiatc art' well and vv ~ cnrood. c F1 OF m i ~ a h Ornicroevoltl tion :111drnacro~volution.fr tl pnrc hist In f tr d i ~ d Instance\ whrrc fixst1 populations were ncttlnllv \ccn t 5on1cririic\ wc ran di. i l a r y vc.cp.~ttrihutcd III partic~r drqccrlr c)t' e\bi)li.\ c .IT<~C I )nrtv~rlirtnj t t .IILTC.arc pl>cr.ll1tx-c the OSCIIT nvrrall as 3 large part of tllu ancc~tr.~ n .d I ~ I I pl.ct>rnc pnsed t o rxhihit quch unbmkrn 3ent. ~ tratc'r t cvoltlt~on 11nkc.~sgi~cd. cithcr by litienge spl~rting .!n rrl. t1ir1~1gh w m r 111Ft.incc i.td.niuss incvi t a l ~ l yhdve bcPn .rllv .~n.~r~t\ qlllc!dv bt. Ii~ i ~ ~aftrr d . W .c h a v c nor It17ti-d tor 1 i In vnnuilr 11. working ~rnainly \v?th vcrtcbratrr. llnn~ yl ~ 111 IT ( t i t h v pattrn.lblc f o s ~ popul. r lrvrl. ? ~ ~ O L I ~ S 1 ~ 1 J T ~r ~ 1 g(*. Althr~ugli ycnc-tlu ' ~ n dl w l n t 1. nnd qtcadv.\.~tint~< ~l wparatcd by abrupt rnorpliolo~cal brcakr.iturc o cvnlunnnsn.cn.\ seen miy1it Ii.loiig-tcnr~ cbpcr. ar ~TC'IICILICIV r~utril. cl~unciatcdby E 1 J r t . ~ r c C I ~ O I I I ln F \t-cti in <trnt~cr. populatltrrl. uf ni~crnevulunnr~arv praccssrs. ~ nrtr nclic .it t i r \. ~ I I I I I111 thC t i w ~ i lrervrcl. hut t t t Elclrcdf~c. I I I ~ L .lIliCC t??gmieiit~rv.lrr Yct this i d i n ( .I~IIIIC'.lncI t11v rd.lncc. 1 1 1 ~ ~ cudlnclcccl.c. howcvcr.111d GC-UI~CIIC171. w a s due tn thc ~ n ~ ' o m p l e t r r nf ~ ~ I I C C .r caws t11c. tliey tvnulrl he utllikrlv trl 1-w iclut~rrncrl vcr? I A Cthat rhrrc arc so fi-w tmnsitioi>nl writ< ~ 1111 c71. ~ t t Accnrtlitlg tn E l d r ~ d ~ c Crru~Jd. it is a n l r : rompnr~ti\-clurcccnrly that .

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