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Contents
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Port One General %laeonto!ogical Con...
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mtR-wllst. mmdmd-& the i m w m h l s
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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 ~ofo nh s s d specie5 T ~ s o n o m i r111er;rrzhy ~ Usr of st$1t1stac;11 ~netliod~
~l~cut>i~\l~~.gy P~I'I~o~c~~Io~ Functiorlnl rrlorpholoq, firowtl~and form Strnt~gaphy L~tbostmtiqnphv Hioaratqrrphv Chronostra tlgrnphy H~lrl~cyrapli~ !$(\oh, trcatlsrs a t ~ dsymposia I~idividualpaprrs a n d other refere~lces
2 Evolution and the fossil record 2.1 Introductiori 1.2 Ilanvln. t l ~ ccpecit.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 nthrnugh tlf popiilat~otls Ilutir>i~ Allupatnc rpcci,~nnii klctcrorllrnny T c ~ t ~ m~crorvolut~o~ldr)'i ng pntturnc Ar~aly
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t Two Invertebrate Phyla 4.1 1)31!-lan1Porifcra: spntlgcs
4.2 Cla\\ificatinri -1.3 C~.I\F D~nio~potigra S p i ~ ~ idl ~~ nr ~ o s p ~ ) i i g c ~ S~l~rospotlycs I:l~.~~tctids Strrunaroporoirls Spl~iiicrozt~ai~s 4.4 C:lais C;ilc;~rca ?... Clngc rracfi l ~ l l ~ l i ~ ~ ~ . l p l ~ ~
trcatircs nt~dsytrlposia lrldiridual pnprrs and othcr rcL.rcncc~ Iintlks,
5 Cnidarians .;,I It~trod~~ctiotl .i 2 Mgor cl~arnctcristic-c; ~ i i dC ~ ; I C C C 43f ( I111y113m Cnidilrin 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~ clf vt~ral5 Cnr~~lq ar colorlics: thc lirnitr nf ztlnntharinn evolution Mir~osordcn I!iblingr.~ph?; Uotlk~,trcatiscs a ~ i dryrriporia Ir1d1rid1tnlpapcrs nrlA clthcr rcfcrcnuc~ Bryozoans h l lntrod~ict~nn h 2 T n o tuamplrc of l i v ~ n ghrl;c>zt,nn~ Rrl~r>rrl)nr~ km ,57l~!ttl??,?
83
ix
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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 ~ofy t l ~ r c cgcrlcra I - " r ~ \ ~ ~ v d t i\ot i~l , , d3tlJ ~ c l ~ ~ ~ i f i ~ a t of 1 0 ;lrt1~1113tcd 11 ~~KIC~IU~C~JS M.Gor L~nttirc.?of br,lc I~it,purI~ r ~ o i p l ~ o l o ~ Etidopu~~ct,itinil .u1~1p w ~ ~ d r ~ p u n c t n t lin c mthcllr '+,ti t,phvlurn 1 ~np~lifi>rrnca !-it!y~i/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 ~t ~inch\ f lclu.11 rpcclus 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 ~ 1~1cd'~o111111~rnltv' \ c l'crrrunn 1 c c f . I F ~ O C I tions ~I Mc\c>7rricb r . ~ c h ~ o p cnrsuclatiul~s d 7.7 Faun;] prrn irlcc\ 7,s S t r ~ t i ~ , r p l l ~L c dI ~ 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 not nr ly ~ l l i ~ ~ t ~ o t l 8.2 C:P.IWl hc.lrloll 8.3 S o n ~ c.~cpcct\of thcll t r ~ o r ~ h o 31ld l o ~yo\%-th C o ~ l c dlloftm I~thogrnphiclirncctonc. Hnvann
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365 3 60 3 07 307 370 372 372 374 377 3x0 3x2 3x2 3x3
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xiii
xiv
Contents
12.7 I c h n o l n p Clnu~ifiratinno f trace cocrils Morphologic.11 m d prcccnratlonal rlascitication 13chnvto~~r.~l cI.~c~ifirntioi~ Phylngcilcric cla5sific ~tiola Urcq of ~ c l l n o l r , g C;crlinicrit.ly ctlvironlllent Crratigrapl~y Fossil l>chav~our Fiil~lio~~pli~~ Rook(,,I! l b l i F L ' a t O ~ { l r l ~ l 1002, ~i cI.c~ . Hrigps, r :o~:I! St,rrltton :~rld Ii;~chcl\T!t~nd p v t cxctllcnt take11 iiito a ttlureutrl ill my 113tivr N C W C ~ S ~ ~ P - I I ~ O ~ IVI~-;> . l r tl~chrytitlil~gc>t'tlic p r o j ~ c r;IS to IIOW I T y n c , to cscape from the min. 'rhcrc. in a dusv &rr ,111Iil prr~t-chcdn-it11 thc ti-~urtlj cditiotl, and r havu casc wort two g s n t arnniniai teq (probably 7'i!r111irrs nri t l ( \ . ~ r t i~logctht,r u.itl1 IJ;~vidHarper, Pcrur had lrvud iirdtioris of y e d s J ~ O Thcv hl~clLl~\i~. S~~s;aii Iligly, Liz H.arpcr, Dick JctT'i.rics. r ~ a t ~ owhich ti has c n n t ~ n u e d~11inEnow, and I all1 still lLgt-m~: ( ' ( ~ 1 ~ A . l ~ t lt . h ~ * ~ rL h ~~ ~ F ;i I ~~ I Oorhcn i~ I ~ Ilavc glad to cscapc to t l . 1 ~hills on a f r u h sutrlrncr mom]I!I,\~L! 111i- ,9r ,111 ~ ~ likc. l r l .lhovC 311, r o rccord r r y hc;jstfclt lifc, a t ~ do f the rncthods avallablc for ~ t sscirntific ri~.~:~h< r r > Irly trut rid st:ll\~nrrfricnd Ct-cilia Tnylor, ,111 tlw \ u p ~ ~ o.und r t l~clpfl~l ~uq;crtinttr ~ h c 112s study. If ro, 1 will havc achlcvcd \\,hnt 1 srt orit 20 do. ,
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CD-ROM T h c Nnt~ir;llH15tury Mtr~rtirrlLc>nd(>ndnd Dr E.N.K. Clnrkcon havc uollahoratcd oil thc d c v c l o p m r ~ ~crft thl\ ~ n i p o r t a trlcw ~ f initlatlvc In pnla~.ontology ttcnc11111g. :I l r l c r q J ; ~ ~O~IiIl L'IL < K O I I II C .I carrrbincci Itnage I~bmn;m d database, c o r l t ~ i i i ~ nrecnrdr g fur 1000 k r y fowl gciicra. E ~ c hrcroi-(1 iotltnln.; .I hct of i l l l d g ~ c ,;i11~1itlform~riono n stratrgrmphic rangc, hard-part rrunrmlog. pnlacoorolnt;.v, p ~ ~ l ~ c o h ~ o ~ tcrc. ~ o The ~ r ~ ~maycs ~ p l ~ havc ~ , bren captured 1ivng thc Natural Hirtow Muscunl'r l ~ i g rc\ol~itioi~ l~ I'AI.AEC>VlSION diptal lmnpng cvcte~n.atld the data arid It~iagtgrcude 111 tlic Gvrtlt~,Srrntcint~barcrn;lilnpcr. Thi\ qwtctn ~ l l o w rthe u w r gc,at flcxlhillt): in di~playingdat;l. For example, users cnn \imply broww frol~i rccnrd t o rcrorcl; pull lip fnsrllc frortr a taxorlonlic indru; o r sclcct and sort record< h-! gt'ogr3pl11~U I I ~ C . lift' linhit, ~ t r ~ t l p n p h range. ic or hy ,I v.~rictyof other cntcaln. l k ~ i i g ccl~artsand p a l n c o h i a g c o ~ ~ ~ iniaps ~ ~ l ~+Ire ic pvrr \tl~crcappropnntc. and thc uwr ran print records. or cxport tlleill illto other ;~pplrcat~o~rs. Wltllin each rrf rhr Ir1,ljor gn>i1p rlicrc ,Ire a 11u111berof labcllcd irnngcc and dldgrarns to allow rhc 11cv1 to bccolnc fnm~li.~r n,ltll ke!. rncvphnlo~icaltcrnls. .'llitrvt;(;)scil< (111 UU-KC_S:\I tn.1). hc urcd as A I I a d ~ u n c tto 111vfbrrrhratcPrlfoeot~rth,yy,o r a? A qtn~~dalorlc prntluct. T a v n 1 1 3 been ~ ~ ~clcctrtlfor tlicir rclcv~t~ct. to Earth qcicncc tcnching \vorldwidc.
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1 I
PART ONE
Concepts
111
Amixeras cf. hartmanni (Oppel], an assembla e oF immature ammonites from the Lower Jurassic of Black Ven, Charmwh, England [ h e r Sinemurion). These specimens were pro&bly ca~strophicall~ buried, since he soft parts must hare been in when +,e, died prwenting sediment penetrofing the chambers. Po~nf~ng by Roisin Moron; original specimen in the James Mitchell Museum, University Callege, Galway, Ireland.
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of nur u~lrlerstandl~lg of t l ~ c11istc)i-y o f lifr-, pcrl1.1p~tlic rno5t significant of all C V C I I ~ F ID(>^ ~ I : I C C about 343 Ma ago nt about the Gcprlnlliq ot' t h c ~ ~ I I I .(l l, ~ l l t )~i11111ot1 vc.lrc . i g ~t h c Earth carrlr inti) C:nmbrlan Pcriod, For nt tlliq sta.yc tlrcrc \\:IF A q ~ c i .II~:,yrrllh~lllt fi>rriijnqf r r ~ i .I~ lionrlcnr~ngdlrc of den pmllfcration oidifl?sent kiilds nf lllarlttc ii~\vcrsl I I L lr.,. ~ l l t r t,iiid g;i\. I\ 111~11 t;lteci rouild t r b m t c ~ . 13un11g rh~scritical period r l ~ cpriiicipnl I, 51r1i 1 , I r y r p~rt~c.lcleof P!lanrrozo~c tltrle. nrc dictinpirchccl 1s). .I 1rh111tlic t i n t 30";I O F tlw ~ J , I I T C ~ * F l~i~ltm-y, Tl~ert. r i c h l ~ c r i t a ~of c the fossil rcunmnq of thu invcstcc r ~ * r i ~ , l l{n>\ < \ I T T I ~ ~ OL .~ ~ I I I \ I I I( ~ c t c r, IjI ~ I L ~'h111rE~rntef that rvolvcd thro~~g!rsucccssivc Ii~qtnncnl ,r~.:.tl,rlc,rc', or L-ynt1ol7acrcn.1)i t ? rock? 341 It 1 ,Ma pcnndq; their ~ t l i d vI$ the dornail~of in\rcrtchr;~rc .1 ,rl l~ir. prcwnl,~l~lv onpnntrd hciorc cl1t.n. pal.wnntnlo~?;nlid rhe qul?iect of thi\ book. Tt.c\: rrnit~lcform< r,f !iir hcc111 ~o hnvc dom~rl.~ted itcrlth cL,r thc 11c~ut2ll01) Mn. nnd evolut~nll t h . ~ t tllrlt, \v,~c vcn slozv. Nr\~enlieless, tllc rr\cib,ictc.n,~atld phato~y~ltlrcricb , ~ c t e r ~ awerc I [ ,r~u!rlcr~tnl 111 ch.111y111~ thc cnvirniitllcllt. fur they #:I: jltf LK!grn i i ~ t o,irl .Itrnosphcrc t11.1t war previLI~vtjiil ~ t \, L I dint nr~imallife eventually IIIIC y~lv.~hl~. hlly \vll~.~r qnlilc. of thc cnrly Irvitly lbeaii<s o t ' t h i ~ l r t l ? llarl rr,,lrhr.d n high Icl~rlnt' phy51oluqlcal diiri F o ~ v iiivcrtcbratcs l occur it1 m a n y kinds ofredimeny r q l ~ l r t ~ t i i - 1 -u r q n t i ~ ~ . l t ~(,117ci o i ~ Inort part~c~iPnslv c.lry rock depo~itcd 111 the senr driritlg the ( I I L ~ I +r\u,~l rcprociuctioi~nnpr,'~tcd) w.lr the rsjte Ph:rnt.roroic. The?? In,ly be very ablitldnt~t in (1 r'rollrt~r>rl,ln,cl~,ingc,.~ct-clcratrd, and with it all lirncr~e\. FIIJICS., ~ it~rc)ner l n r ~ d~-nucf~tot~cs but o n 11 I I I I ~ ofL 'IIC'W ~ . ~ c ) ~ + I \ > II ~I 'I~~~I Cupened cF up to rlic WIIOIC nrr not cor.larno11 m s a i i d ~ t a ~ ~ e s . ivlv:~~.l~ir. Thi5 \v.ls n n t 1ir1t11coiiip~rativclvl ~ t e Scdimcnt.iry ironcrorlc< iilny t~avc rich fo\sil 11 :~rl~1~~gc,~l hissnn. a t i d tl~crt*,we 110 tbrcil ~nltnals rern;~ins.Occnsionally they nre f i ~ ~ m111dFonw conme IIO>L\III L ~ YI ~ L ~ I I I I P t)ldcr I > ~ \ t11aii s h o ~ ~ 7tbO t MA. mcks F U C ~A F L~cy%vnc-kernr~dcvcn coirglo1nerntr5. A:wn tc>g~-thcr tllc. c!~eiilical h c d ~cullrditung ~l111ci)r ,IEI tlic w$r-l-rnrlicd clr- ~ r ~ r i i t i pornclcs, 111~11tr1t1 t l ~ ct;nun,r tvr-ll. 'l'llr.\c ~ r rrninc.n~cly ? cig- halamcc i r uilrtnblc. T h r \cdirncrrt 1i1111 bc in r t w l n ~ cal cquil~briuni(>lily.iftcr dlngcnctic pl1!.cict7clrciilii ~ i f t u . i r ~itb r pn l,~contc\lr*,q. Il.~vct:tkcn pl.~cc.Tlic\c t r ~ . ~~iivolvr y T h c olclcior.~ ( c i ~ , , ~ i -fti i l r ~ itrncc inqctlc) nsc pre~cn,cd. t l i e ~ cchow rridy b~ V and thC h i v l , ~ c s r r ~ ~ b l .prcrciit yc< in t h t rnck nlny tht* rturly of cucli ~-rlould\arc clcccribcrl in ccctirrn llavc tlccrl ~ T J I I F ~ O I . ~ C CL CO~I I I ~C ! I E ~ J I ) L - C . nbradcd. dnm- 7.2. with rcGrct~cc rt> h r ~ ~ l ~ i c ~ p r >Ifc l \.I. fowl lIy FFBCL-, ,i'r For il~sc.~ncr a ~ r and d rnl1xt.d \I it11 eleinellcc nf n t l ~ c rfizln~r.Even cncloccs ,111 c ~ r i ~ i n . ~ 11ollou~ hchvccn the p,Iir nf khclls tlf .I bivalve ur br,~tliln(hit k-~hcI1c.d ,111j1n.1litycrc ~ r i ~ n , ~preqeiit ll\ 111 .I f>und, t1lc.y rnay not br p r c ~ r ~ \ ' r 111 d : C . I I I L ~~~ ~ 6 1 -pod. tliir qpnrr may citlicr hc lcfi rrr~pr).or Lwcomc nle11t- I T ] \ V ~ I I L -t~l r ~ - ~ ~ - c ~ i~lv. ~ it ti cnrer~~rtci~li~-, for tjllcd w r h ~tclvnicnt.In thC 1.1twr C R C C .I ~ ~ d i l ~ l c ~ i t core t f prcrmn,t.d. ~ ~ ~ l ict>rricc i c h n ~ i illtdct t W ~ I C tI1~1 ~ I I l F t d t I C C ' . C " I ~ C . ~ ~ C O I511~ll~ IC 1 1 1 ~ 1~~ F W I > \vltkl~ll ~ V ~ .I fc\V I C J ~ S L l c f i ~ rt~h r \e~lliilri~t r . ( v > ~ p , \ ~tilrn t ~ d rnrk. rock i c crnckcd ilpcil. Thiu hcnrh upon ~ t cc l ~ r t ; ~,I:)~ t , hinct. ihc. ica floor is !lot al\vny+a rcgroll r d ct-mt~r~~t-intetnal mould of thc Lwil shcll. whctcas cxcerous scdirnetlt dcpour~on.rllanv a p p a r t n t l ~corznliu- nal moulds art- left i n thc caviry from \1,311cli lr came. In rx-c cirruitistnncr< the r-orc nr t l ~ cslrcll, nr ut15 r e d i r n r r ~ t . ~\rqucucc< ~ c o ~ ~ t a ~rluinrrmir ti both. tn;rv he rcplaccd hv an cntirclv d~ffcrcntmln51n,tU-scalr Ihrsa ks ( d i a s t ~ m s )rcprcrcn ting yerlodq lc In iroiit l spccicq \V,IF Irvir~g.!-l-lurcm lirr the oldcct 2nd 111mi gcncsal .1ppI1carioil of n~vcrtrl-rr.~te pnlacon mlrrh? : biostratigraphy. Uslrly the qcqucllrc o f fr~\nl K ~ l i i l . t ~tllc . geolo9cal cnhuil~llh a i been c I I ~ ~ F ClipJ C1i7tn ~ .I ht'nrq I~~vcn-tchr~~tc p ~ l . ~ c n r ~ tI \r ~non~i.i!/v l r ~ ~ ~ 5~ t i l ~ ~ JS l ~ d of m.lior gcc3ulapcal t i ~ n eurlttr {prnndc). chit-li ot subdivi~inno f qr'olop, . I < i t iq t ~ i r l ~Enrtl~ n \cit.ncc \vhicl, i~furthcr d~vidcdnltc3 ,I Iiirr,ll-thv nf ~ n ~ n l l t h ~ it< t w n t c c t , ~ ~ ~ l i ~ . ~ lti ir .i >Itt I.l s111 .~lct)t)c- rci.11 ~c unlts. Tlrr \\ 11rrlc b ~ < fur a tl115 li~rtoncnlcSlroiinlop .I bt'ologt'c~l ~ u h l c r t I, x ~ tlilc t rh,lt h,i\ thc uiirqur pcr- i~ tflc dnvumentcd qcqucncc r ~h5cll.; f 111 rhr rtjck~. G p c C t i v ~o f p . v j l i y i ~ . ~ t111l~. l W1tl1111 tllr ~ l t ~ r l ~ ; lof i n Uuc difTcrriit h n d s of (05~11q 11.ivc diftcrrnt iostmtipa- ivhicll thcy occur ~ i a t l ~ r d l ytllc : lnttcr i c ~oinctirilcc 1"" I \ ~ ~ ~ l ~ ~ t qhl .r ~I 'j ~+Rc~ dC ~ C > Z C ~C I C ~ J I I ~ I C I Iknon.r~ ~ ~ . a5 synecology. ;ilrl~ou:l+ h distnl>~~rinn ~l nf indccd thc kcv. to tilidin? out itirtlicr ir~fonii.ittr~t: mnri 12c 11l~~'m'b~.: trs 111 r ~ l l ~ y fKlI I I + C cnii hc cluct:ahnut i t . Stwi~clcI,~s~ltic,i ti011 ,tnd ~ I D M C ~ ~ C ~ . I C II iIr T C dntcrl CIICII c r u r i ~ c q t > K palaeobiogeography (nr nt r31c rnot nf ,111 b ~ n l o p c n l.it~d p:~l:~co~itnlnpii palaeozoogleograpby in thc c.1~~of nnirnnlr) rnrl li~nrk; w ~ r h n u t rhcm iin cnhcrcl~t nnd orrl~.rc~ Zit* r ~ r c t l 111 r.nnlilnrt1on nrrth ~ c + c ~ y l ~ \ - ~ rint.1 i c a lIn y t c r n c)< dot.1 ctnrJgc and rctr~cv~il1.c pn~.;ihl, ~rnrIcnt,~n(Iiny tll~. ti>rni~-r~ . ~ l a t ~ FChlRUIIh \rc ntd T.~x.ionnmy, or systrmatics .I.; ~t winctltlr~, iiiovciiiciitc c~~rnntiiicnt.il m.~\\c\. k ~ ~ o w ii si ,tlic \rrc.ncc o f ~ l , l r q i h ~ . ~ ror ~ onrgnlllrim ii All nf rhtrr aqiccr$ ~ r t ' p : ~ l : i c o ~ ~ t carc > l nii~rclrc~ ' It n the oldesr of dl1 thc h o l o q c a l diwipliiicr. 1.1rcd. rid :III ,~rlv.uiccit1 nnc iii.1~h : l \ ~.I hc.>nng thc pnlKl~!t'> i~titlinrdLlv Cni-1 t;ti\tn\' I ~ ~ ? n , i r r ~ . iipnn nnv ~ > t I l ~ T = rh. ~ w.I p : ~ r t ~ r u lctutly . ~ r i r ~fiii~c- ( 1 7117-1 77s)in htq pioncrnny S)~,ft'~ri~t rrrrrrr?.rtc, ~ r . tlnt~:~I n ~ c > q ? h n l n ~ t11.1y y ytvr irit;,im.itinr nil chll in uqc tr,dny. though ~ ~ ? ; i t lilr.r~diticrl v In8' p~lacc>~=cologv ,1111- ~ ( I F E I ~ IYC~ITIC* Y ~ C ' C C I F M C ~ to tnw311- curcndcrl. o n ~ p:a\ \~1.11.L i k ~ivi\c. rccchllr r t - t i n r t ~ l c r i~l lt ~t.iuonotnlc pr.1~-clccli,ivc cr1.1blcd ~ h c dcvr.lnprncnr . nf .i The species concept rrluc: 11 nirlrc, prrbilsn~piv~~itlc rlf IvI~lnq,?c,~l ongill can Anim.11 ~ p c c ~ c(sc g. Sy\vcctcr-Uradlry, I'l.5f1) Jrl 1101~ hrb rccovcrrd frclm ,rnc~c-n t lrkr ,lnli f ~ m nrl~c q r o u p a f ~r~dividualr rhar gcncr~llylook ltkc cat1 h,lrir of biornolecular palaeontology. Such fnisil ntlicr .lnd can intcrbrccd to product ~tTcd spccics, o r rhow only rnlnnr \.ari.~tinnFa that would hc c ~ p c c t c di r l a lclcnl v,~ri,~nr n.lcu5, \\,l~crcbvt-nc11 cpcclrs is drfined hy n ~ ~ klntl n w t h i i l r h fame ~ ~ ~ C CO ~ tCh cCr .\pccic.\ i r ~th? CIUIIJ ncc: tllc generic nnA specific (or trivial) ndint's. rt.l.~tcd. m.iy hc ~ C W snd . if rcl a fill1 rcchnical drqunption csnnlplc, all cars. 1.11-scand < i i i a l l . h rnlict hc. prcp;lrcd for cnch ncni I onc pnrnci11,ir ~ ~ t w i11.1~ p heen ~ I . I C C L ~ ln t1w ~ v ~ rillu5trationc ~pccicc,~rliirl~ should frc pzihli\l~cdin a pdncontoaoa pt.!i.r. C)t the. IfLlnnuc.;pcClc.i<prcirc >~I 111 pnl.lentrmln=, a< In rnosr Ilnnr: spec1ilreilTIL>ltlK Whcrc thc f r w i l be ~ d r n t ~ f t ca.cd bclotrgnl: ro a Crerius E ~ ~ r i r o n cILccd r ~ ~ 29 17 ~prlIly L J k i ~ o w nTL-IIII~.hut L-.I~~I-ILXI7r .~sirbl~rd to .L spru~cs(as S p c c ~ r si ~ d v r t ~Rccd a 1 9 17 l l l l \ rc.llf~ ma\* l ~ ctlir c n w nhi.rc prr-cqt t pnrncuI.~rcct of n~les,whrch h a v t bccn woskcd out s11r ulnrity rcrults fror~luonvcrgcnt cvdution . Hut I n I l ? > V flcrcl t t by a SCIYCC of 111~ P ~ I S ~ CC,IIIIT~~\SIOIIF I O ~ I ~ ~ and arc a5sccsiilp 'rin~ilantics' hot\* doc\ nnc d c c ~ d c~~pcln 111111-c i ~ l i p d o c ~ r n ~ c u t cin d hll 111 the oyrnllly paKcs of cac11 which cliaracccn arc in~portantiHow h o u l d tllci ~~l)tC1l~1 11lL-2 ~-olun,~c of rhc -/'t.,-nrr ,111 I t : r,t'rtc*/rr~~rc P 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~ t contlrlulng wnec ni vol~uiirs p ~ ~ b h s h c hv d the natural order h ~ o u p c ? T h c ~ic n o ~inivcrf.~ll\ l'he tllirr Gcolng~c~ Socicw l iri A~iirnrn). accepted rlrttllod of E ~ c i l ~ t n t i n gthccc cndc ,111ri tnnc. empl t.lxonolni~tshavc uwd diffcrcnt m c t l i o d ~ .In rtccnt ~ r i . ~ n i ! > q t , 11, Taxonomic hierarchy ycnn th ruc f l ~ ~ r p l ycnntr,xsting ~ ~ l i o o I 11avc < :Ill. distribut Althntr~h313 tnsonanuc catcgorlrq ahove tl~cspccj~*s c n ~ c r ~ thrcc d : arc rht. schnoh of ( I ) c.vnlutio~ian !>Ills i t i\ lcvcl .II r to samr c s t r n t art~ficlal 2nd ~~~hjecltivc.,taxunonly, ( 2 ) t ~ u n ~ c r i ctaxonomy al and (3) cladlqn~ FTS tcmatic ~ d e , ~ l they l y shanlcl .IS f ~ ns r ~ O F F I ~reflect ~ C evoluUllril the 197115 rnoft palncorirolopsts, crprcl,~ll\ l . l t ~ ~ 7 l ~ l n o n a y relation\hrp.c. tl~oscwurkins with fowil liiatcrinl bric h tlicy h ~ v c , i p p I ~ ~vtgr ii similar speclrq ,Ire ~;.roupcd III genera (rinp~bdr rallcctcd in thc firld, wcrc cvolutinnnry tasonn- rr~dCracrnf genus). ycncr"in families. t;lrlul~cs 111 orders, miqts, 1n crccting ;I hicr,~rthicnlclnwificntiou, circh t1i~v.c' 1vct.c o r d c r ~m rlasscs artd clnrsrc I I F tilt Iargrst C11~1siut1 ~i ilasqical tasonornlsts lircd a tr,~d~tinnnl and very flex- ,hftlllt, € 1 1 p~ thc an~mnl klngdn:~~:phyla ( a ~ y ~ l aphylum). r ihlc ~ i m i h i ~ ? a t i uonf trlrcnn. Fint thcrc i~ morpho- i.i olvcd. c l , ~ r7dt~1rilItct'd
6
[
t a t
02
8
Divisions of invertebrate paloeontology
11
logical (nr phenetic) resenlblance, thc c s t c n t to Kir thr 11105t etTcctivr tri~th~>cl for ~ C C I O I I F ~ T Itin? IC phylogeny. S l ~ ~ i t l r( 1994) '~ c ~ ~ l ~ n of r nrladist~c n ~ ~ r r 1 ~ i ( . I l ~IIC ,71111~1;115 ~ L Y C ~ I ~ J,111~ ~ C ,~rlotlicr Sccnnd, ~ylageacticrelationship? .Irr ~lr,iigwith phc- nlcthodology i ~ c rn L-c>mprrhci~sivcthat mJv hrlef rrc r ~ ~ r c r n h ~ .corlc~dc.rc~I ii~c~ ~~nport:iiit, By thlq is commc.nts ,trc ~ ~ v hcrc. c u rhc 1v.1~(2s h r .I\ c,~rl bc d~.tcnnincd) in Hcnirip war of rllc aysinlnn that rccciicy of coluI l r 11 ,~nlnl.ilq nrr a c t ~ ~ . ~rclntcd llv to C A C othvr, ~ 1.c. Inon o r ~ g i coulcd i~ b c ~ hc t choliln 11). rlir sl~nrrdpoctrnnv nf rcccncv o f c c m i t n o t ~~3rigi11.\vIii~hCIF w ~ snil r of cvolu tiom? novc-lnc~ clr 'drrlvcil urw yr.rrlcr intn ~ ~ ~ l l ~ i t i i~~. IrS~O. Ii I~O\ ~~Thr I I ~ . ~11.1rnttcr;~. Thus in clorcll; rel,brrd g r o ~ i pwc ~ \vould srr~i~r OV ' i l l ~ ~ c ~ ~ 111 i 0 1C7 ~ Crot k r ~ c o r d, ~ n dgcn' ~ l ~ a r cderived d c'l~ar,~ctcrr' (synapomorphics) ~ ~ p l l i c ndincupt war that in a11j7 g o n p lTrlnnmv,whi~-htook . ~ l l h c t n n into ronq~dcr.~tinn. char~cters,lrc ci thcr 'prim1 tivc' (symplesiomor1.11 for n Fni19 tlnlc I ~ P T the~ backhcli~cnf p a l ~ ~ o r ~ r ophic) or 'dcrivcd'. TIILIY 211 vcrtchsatcr hnvc lurk! lac,rf cP:i l t ~i tt ~~~~d ~t ~~q ~. ~ I . An assemblage refers to a ~ ~ t ~snnlple g l c €rotrl a ~ ~ l v ca n ti mlcrocoinputrrs nnd thc propatticular horizori. pcci~lt\t. wftw,lr~. ppnckngr.i dcqigned 7. A n association refers to a group of assenlblagm, u rncct thc nerck uf pnl,~tontoli)pstq all ~ h o w n gsindar recurrent parterns of spccrm ''1LC; I A I-; Rn~rrtli .mri tl.lrprr. I qr)C): Ry;~nrr composition. lLj()i), tlie me o i n u r n c n c ~ lnnnlvsir 1s becnn~lng 3. A palaeacomrnunity (or fossil community) ~ ~ ~ ~ Sr~ti~tic.d d i r ~ l . rnctliod~: I ~ P likewise enrntial refers to an awernblnge, assoc~ationor group of d;tinrny p,~l,~ctlcornrnr~nitil.c. i n 'undcform~ng' nssocianons inferred to represent a oncc-distinc;rrr+ of dcfttmicd fclc.;ilv xr~dcornpanme them nvc biological entity. Normally tills only repre:,I k9n~ltt'rt.rl mntennl. sents the prcsenrable parts o f an ongitltlal biological cotnnlunlty, ~ i n c ethe soft-hodicd animal$ are not preserved, This definitton cerresponds more or less exactly to that of Kauffman and Scoa (1 976). 11
I
(
1'
C r011
ctc [I.*(
11.11 111-
es. 111-
:nilur cnreynrlcr, msv hc iricludcd In palacohiulla. palatclr~ulclg(hew diwus~edwith pnlnro\~icr::qnpl~v). fu~lctln~lnI m o r p h o l u ~~ n ~rlinology d , :is 11 ot wluch scquircs wrne d~scussion.
\
lull 1c:11
13
Palaeoecalogy
rl y.
\I:,LL rcolog
'"S
111,lr
IIF
the shdy of animal!, i n rc1;lann to the qtudv of
r~nlronrnrrit,palac-occolohy i r
Pataeoecolo~y must alwayi remain a pnrtial and incomplete science, for so much of the infomaoon available for the study of modern ecosystems is runply not preserved in ancient ones. The animals themselves are a11 dead and their soft parts havc gone; thc o r i ~ n a physics l and uhenmtry of thc cnvironlnent is not directly observable and call only he inferred from such srconchy evidcncc as Ir
Modern 111ar-int. csivirc~n~nt.rit< arc qradcil =?rcortlavailahlt-; 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 d d l drld .I[ ~ h cdgt c tt7r f i l ~ ~ ,IS~C~III-I.I~C~ ~l that .ITC f01it7C1 m,)v well Fr qhorr pmdc into tliu ' r ~ ~ b t ~ shelf. 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. 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 VofT ' I~j'l~ic-h ~ ) , ,117 ,~dtlrti~~r~;ll rlitt~en\intr with tlic sl~orr. I n addit1011 thcre is n ~ e n c r , ~ l ~nvnlrcr h u r i ~ l prvcc<sc< (biostratinomy). Tliix dccrca~c111 abundance (number of individudl5) I7ut 1;ot r~rcrrsarily diversity ( n u n ~ b r r01- specie ~ c 3 l 'l \ v . ~ t r n;Ire r - ~ p ~ myd (
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
1.2I12rc1 31-
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 ~ lI\ r > hich ~ ~ c:111 c*rcinnl i k "lighlv cnn~rrriiii~-rl by g c ~ n ~ c t nrules. t' Il* L!h*iii~c.cri trcvl~fu\r~lrzcclrcrnalllc ,~lotir-.IYC p-t~wrh ~ o \ v t l i~ ~ O C C S F C F2nd , t h r proprrtlcq n f !l~:itcrinls ~ 1 1 ~hn11 1 Fn!In\viii~ tlic C I ~ I ~ \ I ~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 sw e n h w n ~ cnrt. n t t r J c t n ~ ,ordcrfy and ctablc contipurarlotls uF nl.ltter that nllirt I I C C C F S ~ ~CIncrTc ~V ~n the 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 ,~vnilable l 11~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, i rathcr than a< JII c n ~ t 2 r e (lacob. r lEJ77:scc ~ l \ Chapter n 3) - a point to be bonlr In m i i d in ccm~~dcrirlg =llIlivin? 311d tbssll org.lt-tIFrnr.
stratigraphy nnd chronostratipaphy. aU o f wl~icli are w n y of ordering rock stsdta Into m c n n i n ~ r l utlitr (Hedbcrs. 197h; Hnllatld rr n l . , I W X ) .
I< t
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I
~
7-1
i ni ~ % ~
rhq-
4:3tly
!'
rill- tlir~ct (-8
Lithostmtigraphy ~l~llllil~l rIIIIIIr~llIr[' Lithostratipayhv 15 i-ot~ccrnrdwith rhc c-rec-tlnn 171 IlT t 4 \ . tfll? becn deposited . ~ tthe o ~ i etittle; slilce it cut\ acrnr tlorl \ \ ~ 1 t I i z ~ ~ nplnnes e i t i q snid to be diachronous. Sucf, i l f * ~ ~fl~l ) l l l ~~i.irlr M es~creda t the qamc time XVithiti a qnl;lll qyacc - ,I has ch.ltigcd itz ch.br,irtrr r,lpidlv. The 11ldivid~131 ~snrI%ronc l.l\tl.(l \(Wll Inycr, for lnlta~lue,passlnp ~ntna n some diaance nw,rv. Litliustratigmphv IS t h u ~only r)l bedding p1a11c.s.T l ~ c hcciJ~ng ~c pl,~ncsare time honi t 1% $111,w of bottom- notably bv thc Gcrlilan palaeontolopist A. Orjpcl 111 du dliny invcrtcl~rawr,arc hcicc roiirrnlled. Thev the 1R.ills. lqvcd In particular cnvlrnnmcnrq nnly. c.g. I ~ m e It was Oppel too w h o fimt recoplizrd t l ~ there r ~ n u d\chi Ruor. sccf. l ~ )trcut) p x n c dijtiiulticu in t ~ ~ n l i i nprcii5r y zo~r;~l Ivriri!lcvr. NC1I~~tiotr. C l l ~ p n l > nk ~ 1 l a ~ t / . 1 2 ~LIOII, 3~~ :#,I, rrf cq11iv.1lcnt 4 1 ~ r 'ill \ v i ~ l ~ ~srl-r>ratc~l ly arc:)$, (Nc\.v. V ~ ~ ~ ~ .IIILI U I I rc;~d;thlc. F tcst) i II! fn'ni~ilxrcIur.nrrn\, tl~cl u ?I, \vull d o u ~ n r l c n t c i\r'itli i i$ricgs+D.E.C;. ,ant1 Crc~\vtIlrr~ i3. (I t3tJO) f'L71L~r~~/jr+ll(!tf)~: ir ,111 c l ~ ~.Irc.i, c rn,iy collt.iin vcr! fcrv r . l c ~ r ~ c ~in~ t c .Sptrhcsi~. Ill.~c,liwrll. CJxlrrrLl, 583 py. (-Tlir. ~ L * < I ,111~1 . ~ l ~ ~ ~ r~f t i ~n i~~r,d ~ ,.1r1d1 ~, ~ t;ill, E1nt.r. d i c y hcltrr~gtn Inmt 11seh11p 4 r l ~ ~ r u r ~ t o lCo ~~ ~c IC~~I I~ R - T 11f:dI) I ~ ~ I J ~ ~ I . ( 1 {)74) ilf,in'1r~Z ~ ~ ~ x ~ i - e i ~ tblcllnlitntion of G~ln.11proviilcc\) !.!? nc5, IIC)LVC\~~,T. tilt- I l n ~ ~ i ~ c lof ~ r siir'l~ i~% H n ~ t o ~ i [>.I-. . 2nd Harprbr, 13.A.T. (1 9')4!) Micm-$:,I\ y I I J I : ~ n~cillatctlto . ~ n dfro. '['F~crcnr:~! L ~ O I ~ I I J I I ~ C T \ 111 p ~ l : ~ c o ~ ~ t o l of .p' ~. ~ r ~ ~ r i i ~ rqtt/' i ~I//(~n>~ .!~lri.~r)rc ,Iilllcal- d c r l ~ r - r ~o~f c~ d l n c ~ hullal nt prtlvP ~ i ! ~ ~ ~ ~ +3 I~I tI X ~ t~ tJ i~I.Ill~I ~/ o~370. ,t i ~\ -1~ ~fE.( # pqwrs) IU.A111 vrrtic,~lSLICL~ w i ( w ,T \ I L I $ C I C I ~ ~ T : Isrrilti~I~I~ Cuwcrr, Irt., r l r . ~ s r,ldinnwtrii cl,itin: lvill Fnlrhndpr.. R.W. ,1nd J~hlnrlski, I). (ctls) (11)7')1 '1111. .!~!+l*ncilc p ~ l . u r o n r n l o ~ ~ccvrcl.ltion; ~-,~l t31t. two .arc I ~ ~ ~ c ~ * r / q~j~Efirt/~ p t * ~.Sr;ct~r-tx, l i ~ i Veil, V l l + TJtt*L't~yc!ryi,dr(r 1il'r:I~ ( - t ~ l t ~ ~ 3 l e l 1 .~lFTLi ~ ~ ~ tI1c t , ~ lp-vGlr ~ ~ ~ sLlc.c-r-5$ l>f (I{ Pftlt~i~trrolaqy.I ) n t v d r l ~ . H u t c h ~ r ~ r r >a~~i r d I G l c s , :l ~ ~ ~ ~ ~ ~ ~ ~ it]~,p~ttm ~ tr d~ 11% g tl -~ ; i ~~ ~i ~ lt l~mvcs ~t .i v ~ ~ ~Strnudslm-g, . i'cnn. (lnv.11u.tblr rctrrcl~cc) II,;!YII to 17vtl1. Tlic 11s~c~f'.rutnln.~tic d c ~ t a - p r i ~ c c w i t i g G n i c I r i ~ ~Ii~~tralrr~r). ritr~ ~ m f t c m r r ,Wl~ckw~ll, C3?rl'nrLi (Lucid rxplanatiun o f rladiitlc mcthodulop) Iluckwnrth. Lnr~clnr~(E:nit~rnr~n~ty r r r n i ? ~ r n i t ~ t ~ n n r ) 5ylve~tcr-Hradley,1j.C. (cd.) ( 1 Y 5 h ) 77ic Sperie'cr Colrit.ppr rrl! P I r b t ~ t ~ t ~ r l ~S/y~~~tqcyn,~ r t i c,4~~ocintio11, < London. (13avc M ~ - K c t m \ v S, . ~ r l dSCC?~CCC, C . R . (cJcI (IC)C)O) Palaeo701c. work nf r c v c n l papcrq on pdaculltological tas@narriv~ Tcicl~crt,C. (1075) 'Trticltrsr o ~ rIrrr~cn~bro!uPaleattr(~lq~y, I1nrt W (Suppl. I ) , 7i.iart. F ~ r r r l r ar~ti Pro/~ir.t~ini~m. p + ~ r s ~ I I C B U ~ I I I ~I'~~.I'C'ZCIIC \1orI11 n1.11lc) Crolugc-af S o r t r w oKhrlerica, Lawrmcc, Kan. M~ddlclii~rs,F. A,, I i i ~ ~111r rI ' W B I ,111dI I U I ~~111~1islict-I by (.'3111l~i~!i~- u t rcirrlrrulllir> r c o l o q .~ardp : ~ l , ~ c n e r o l (ill Ii1;d C~~t.~~~!ic~ttorr L! / ) ~ T ~ ~ ~ : ~ ~ O I ~ I I (~ cI Ih~ II tI .IW. ~ J P Sc~ntt T Univcruitv 3 'rc.;.;) 'rh0r.t311. (;. ( 1 L):l 1 ILtfi?i t t t l ~SI>J,UJor!d U ~ i v ~ ~ k i t y ,111tl 1 t . K LVrstl. Ilou.rlrir, I ! U I C ~ > I I ) F ~ I I J I I ~ J ~ ~ t39, o h ~ 1y-IC]-3. 511:d~b{Lo\vcr Carbonlicrnr~lution ailcl rlrvcrqzty in by the rondridc +~nrl~zatchingtthc car7 go p x t but the fowiI rcuord. 111 t 1 1 ~rrc.oild part of. tlliq chuptcr thc G.11.1p.l ~ t ~ t l ~niiy a n ~de,i t oil>c>\vtheir cngincq ~ r ~ o r kto' . UFL' t h c r r is a inore cutendcd auccrunt of ~ v h a ttlie fosril r ~ic(~ii~ Umziwtr's ( 1 973) p ~ p h ~wct ~ i l e . For tlic fos~il rccord can tcll uq about thc n;\tnrc of rvoIutio!~nr)' EX.73. Icd rrcord cannot wvc n~lirli~nform,inr>rl nu the xncchchangc. d~vcrrrtyin .11liqrn of rvnlutintlnr). rhangc; this has to hc progroup "f bl iriiled by gci~ctlcq,c v r o l o ~ , niolcculnr , b l n l n q and origii,~l pn population dyn=irn~cllttivn.in t h o u ~ h t .,tnd clrlwl!, cnrrcct, hut to this pniiit wc \l1,111 rctrirt> I.ltrr. ~~i:~.mv~nctl \xitIi I T I < cornlp.lrJrlvc aii,lLrlrllv. Thu C3f tllr prc-1Janviui;lii rvnlut~otii~t:,tl~c. ttiort i~l.-finp-rrr\,n r prilt.ui,~c~ l I i. n ~ l ~ higlicr vrrtr-- p r c > ~ n i rt~ r ~ i the Frcllc h i i ~ t ~ ~ r , l JC~~~1-l%iptir\I.i17sc.,~1111rrstncnnn nf n ~ v l i ~ cmade h nni1nL~l< bccnrnc iilorrnC ~ I T I P I CHC E . frlt 1 1
28
Evolution and the fossil record clrpnrlq ttruct iri\tv horn lrciv I I C ~ C L \ and t h ~ a thu nnturc of' v ~ r i a t ~ r ~ and r l hCrrdity wal I~scc there '~url~ritc~d c.h,~r,)c.tcrund. fi>rt l l ~ ,n105r S C * ~ ~ I I I I~C ' c ~ k ~ivwof tlic r h c t ~ r y , Ir pr~,ily, ~\ n~,rw. F n l l i ~ w i n~~h l c p-occ5s of sw~tcliingolr of stnicniral gcricy rs FIVCII I V ~ I C M .I rr,murription', ,I fu~rthrr ~ r ~ ~ l ~ c(mcwcngcr ulc wnsnr gciie receive? a n JpFropnatc 7 tllnnl~rc. 14 t~jnnerl.ivhir 1 mcwcr oiit of' rhc t ~ ~ ~ c l e i i c , Slriiplr orcnrusrlrs such A S bactcr~aand f ~ m p have I tlir~utghprrc.5 ln thc nuclear u~crnlir~ir~e.cets of;l$acent grn.ut-sk r ~ o ~asv operons, ~l cndrrrg h r Ill ; ~t%clt' rn thc nlwconir. (TI116 I C 1101 a a p a r t ~ c u l ~ tl~ctaholic r path\v,ly. a'l~chc can b ~ . a1 nlI 1'1 ''CiF- b l 1 3 ~ . c ~for ~ ~.IT r . tli~sst,i?r the Tencs r~vitclicdr r f i a n d or1 tngetlrcr. In Iugllci- ~ ~ ~ J ~ F T I I S tls*~~~\r.l~cr arc innil~frc~l. I t IlaX 13ern chorr~iirrccn tly Ilnwover, g e l w ~vhicllror~tsold~tlircritp.lrts of .I 111:
Ijllr
'
1
,
30
Evolution and the fossil record
---.-..-,
%conrdin,itr.rl Ilrogr:imnic, m.iz hc ~itt' tc) t h r broprrrph,~w.t.rlhcrc two ccnrnolcc moyc to nppnn~-r~trt>t~ of a pnlcq nnd a ' ~ p i ~ l ~ f l(>T c ' r r n t ~ i n t J i ~ c . ~.ippc> d~ : n A t niet.~ph.lwthc ntirlc.i~-tiicn~hr1 9 /--*\ pnrtic~ilal- c l ~ . i r ~ c t ~ A r. . s o ~ i ~ cgericas .ITC h ~ h % , ~ {thcm. pleiotropir; thar ~ u , tl~r'y i k h t .;vl-cr,ll cli.lnctcrc vnnlsJlcs . ~ n dthc chror;lmolnc< 11nc up .1Jn11g t l y -\;. . uncc tht. v n l c gciir- prod~rrtqmav uwJ 111 d~ffcr- m c d ~ nplanc, ~ ~ ~ 3 ~ 1cpllthng 1 111 h 1'0 FO that c.v c n t biocl~cmic~il p.1rl1w.a~~. A p l ~ c r ~ c ~ r n c\n~oli ~ i ;c ~ h t chrcrt~r~ttd bccortlcc 3 Jnughtc.r cl~rcrlrlotlrcr.TIir\. r n ~ c ~ ~ c r nor r n t l l e ~111ob11e ~ or yztnf;l- thc h u n ~ ~dcmcq,hut nor co~ilplerelysn, fcjr thnugk ~ r n ~ yl cn r t i c c v c i ~ t- ,I p h r ~ ~ o n ~ r n1%-Enrh on has c.~chp ~ n o d c r n cc ~ r r i c s~ t so ~ v ngcnc pool tl~crcir clc.~r irripl~ratiorl\Lor r.lp~drvnltrtlon n l ~ dfor t l ~ c come pmcihility of r s c l ~ a ~ i g31id e IICIIC'C q c i i ~flnv; nngin of I I ~ X Vtvpc'c ut'orqnn~z.lt~oi~. hcrwccn the pools. A II~IY .~llclc,f'ornltd llv ~ r ~ i ~ t . l t i n r ~hcntab'c , I t i s within such nn.111 natural popul.ltiol~r t11,1t and will prnducc chatlyc5 In our or mnre chnr.lctcr~ f;~vourahlctnutatiorls. eeprc~allvthnw nllorvinq llc'r111 thc ilry3mstn. Thcsc. rh,~tl<e:c'; .lrc lrfunlly of cmnll tcr ~C(;lptatic)~~~. arc able to spre.id. A iiiut,lnt condlccdc .]tad uf lin~itcdc t t k t : 1f they .ire I . ~ r ~ tl~cy c r arc tion c-qt,~hli~hed cvcn In o n e btnmly Ins!, l i 11 vcn7 oticn lut11.1l. hlclrt tnutat~ons.ire iicutml. Gomc pnrsccqu\ sdcctive ~ d v ~ r l r ~ l p roon e . sprc,~d~limuyli 3rc ~ ~ F , I ~ V . I ~ ~ ~ I ~ Ch' tI ~) UCt C> ~. ~ I C IITI I J ~I C A ~ tn ncw or thc. p ~ p ~ ~ l ,it1 ~ tthc i o gnmnden>c ~~ wlth~rirc,l,~t~vch m r n c a ~ h . i t ciiff rcrit clinr~crsm ~\pZl~cl~ msv c a F fr\v grncration~. It 111;rv then ~ p r r i r lt ( ~ nttlc't qomc .~r.lcctive; i i f v . i ~ l r a ~A~n. ~ c l v a tagcuuc n muta- p x i ~ o d r n l ein~ thc v i c ~ n ~ t )iucrcaqiny ., tlic c f k c t i \ c tion. wl-tich IlIav ctnlv II.IVC J n i ~ t p n n l l ybriirtjcld rnngr of the u s e f ~ m ~ l l t n n t coiid~non.Surli n rnuzint cfft.ct,will nrllv be ~1-3cto sprmd ~f (1) it ic rccurrcnt condition, especially ~f occurring in 3 peripheral w i t h ~ n rhc poyu1,itroil 40 rhnt net%, recruit5 arc isolate (i.c. a s i n ~ l eg~rnwdeitle.nr small cluctcr r)! continu.illv addrd to thc rnurnnt crack. (2) ir g a ~ n n d c ~ l ~oil e s tlae fringes n i n lnrgcr ~vidccyrr.~d ranics rurilc selertrvr advanr~gt. w that fhc populatio~~), will he the finr step toward7 thc frmn.4Hardy-IWcl~tberg eql~ilibrrutn19 ~ h l t i ~~n d I ~ Ffavnur tlun o f a new and disr~nctspecrcx. The nccurnulaticlr ,111d (3) thc p o p ~ i l a t ~ o11rn qtnnll c t i o ~ ~ gfist h ~t t o bc o l a fe\v more inutattnns rnny thcn iin rnut,iticlrl: tnultigik of spec~esfro111 such periphcrnl 1xolntr.s i q kuowti ;llldt.v h ~ v cx-I\eil .it d~tf'crrnttiincs. 5 o m r inutatlt allopatric speciation ( r ~ ~ n c 2.3). l n Wlicn the mica allclcc r~iavs p o ~ ~ t a ~ ~mzitntc e o u ~ lback ~ ~ > g i n to tl~c. h - 1 7 ~is sterile wittl the pfirnt srnrk, Iuvinp rliverpcd nriyinal qcnc csprrlclorl from uhic-h they arose. c l ~ o u q h grr~etrcnllv, thc 'parei~r' 2nd ' J ~ i i g htcr' T h c ratcq of 1n~tt.iti~111 \';In wztll ti]? sr-1lC. SOIIICarc spccics n r t then c r ~ t ~ r e rli4titlct. ly niucla h ~ s h c rt1i.111 ntllcts. but rhcy ,lrc h~ricr;tll~ lLct-cssive niutntlons rprcnd c-nn>p~r.~t~vrJ\ %lr>~vlv low, onc n ~ ~ l ~ ; ~ lt t~~cE 0l t0l1 10 ti)r .I p.irti~l~lar ~ e n t ~ r l doidy in urlall pnrr~lnnnnc~vithclosc i 11hrt.c.dirie bcing c a n ~ i d c r r d,25 I~igh.The t7tc of rrultat~onc ~ u I F thcrr any I~kclihaod of ulorc r ~ p i d+yrmd~n: Z,t. 1~1crca.icdb\ rt~dl.itrnt~. ~ t i c l ~ ~ d irxpo\trrc ng to Holvc~rer,recewve n ~ t ~ t ~ t t ~c.111 o l i \OCCC>~~ICJunlrultravio1c.t light, u r 1v . cl~etrucnlnlr.itnr. I I X H ~ i~ridrr certnln circun~qt,ulccq Fspcrirner~tt have c h o ~ m that chan_rcs in tcrnpcr.irurc .ilon. mag bc ctio~rgli ro rnndlfv tlir dornrnnncc of .. geuc. I r l n n h i r ~ ipoplilat~on ~ 13c111g Cut C ~hr)t,c.Hut 111.1y I\c cq1 It
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!4?ylc-:llor l7 t l O 1 1 ~% t11;i1 rh t l r ~brcrc
ltirhiy coy i:cIlctIc irlcl I l i z r , bccaua L
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tllc Ilvhridv T l l r t c nr >clr~c.vcJ.P frcvui ci tot irlrn ITI,+IJV ~ ! , I t l ( > l l 15 11
I1 1,IY L-ollle 41111\'
,3 ~CIV
tCl.cd R E I e~ ~mc't. The* I' C ciitTcrc,nt cllroiiiosnrric ltngths or ~ r l drcrotnbination. Thcy m;ly 1 3 ~'st;ll~1117~:d'111 ,I tlon\ c?i allr.lcr, rcc-unlh~natit>nir trnpo'i\il7lc .inti pnrticular kind of sclcction t h ~ tnutu;~ll\.prcvcnt< I1ybnd\ nlwrt In r.~rjhn.r~ nr Jrr ~tcrllc. cvolutinn i r i ,I I,lrgc population, fnr ~f rluc ~ p c s l cir~ 111'rcArc r ~ t l l I~V rJ Y ~111 wlilrh i.~ol.ltlc>n might bc $ ~ i c c c ~ dwiJc.;prc-ad ~~l, and well adapted to i n cnrrlr r c d . A pnp~il,ation' ~ - x p l < t ~ uot r~ II~IFII, r" dcnvcd ronmcnt, t h t o i i d r ~ ~ m h l are t s \vccdcd unt -trid tllc I I J f ~ i l l ~ d cpopillation. r rn;lv initl.~ll~cxpnnd OIIIV i-qwr.~tinrl,~l srlecrion I C stabilizing s~lcction. I lnlanv rwn arras in M hlcll rorr~pctirionor prcThir tl; an~logouqto what Mivarr pmpo3cd in Irlc III I\ rniii~nral. A h tlic tint HII+II. IICI\\JC~,CT, criciquc of D ~ n v i i ~ On . thc ~ r h r rhand, qcztc fieI k v ,111~equally ~ ~ sudden ~ crash. which 1c:lvcs qucticics bcpn to ;ilter when thc rnwrniiincnt : J irw rtnall tsvZated p o p ~ i I , ~ t ijvidcly ~ ~ ~ ~ sccat. changcv to becnrnc castcr ar harhhcr nr whcn .I popII I I ? ~ ~r3ch \i-itll its own sarnplc ot'nrhc total gcnc ulntior nligrgratcq into a 11cw area and. ki11~11117littlr I b e w In turn t11,iv act a5 foundurr of new cnnipctitinn. hcpnq to dift'crcntiatc c . c o l n ~ ~ c ~ l l v . :$, at Ica\t. sonlr of wllich may runrivc. WI~cre Thc ~ntcnsity o f hclcctinn unclcr t h c ~ elatrcr ell-cum1 1 1 , : r a i Y ~I J I ~ C ~ O ~ L I I . I I I C I I ~ I C hccn bur ivipcd 5 t ~ n c cis~ much LTCJ~C~.. At F I I C ~ nIrlc5 rhr'rr 1 ~ 1 , i j r 1 qrinic ~ ~ t . ~ ~ t ~ otllr p I iqvcirs c, nidv rcc0vc.r occur directional selcctio~itnw;~tcivollr cxtrvrrlc ir, 1 ,YFIIIL ,~q.~in. ~ !i(>ue w r , I L ]XI> p.iqscd t l v n ~ i q al ~ rc111tlic t nni$r, at h i n I ~ > ! ~ c~.l~r l~ ~ + cMCIICCIII.T~ lh'. drive in coil- looks I ~ k e T . h c ~ csrru~to work 011 aiid iiitt.h~atetll,, h:~vc.varic.~ ~ c i ~ ~ i u m:1v ~ l c cIlc . ~ ~ l n t h c w.\y r i r l wliich qprci~q~ ~ 1 1 1 qiqinlc produccd by thc qcpnenmtmon grnrr. PI I1;rs 17ci-11 ~ r i \ c . .~r-r-i~lt.nt.~ll\., . ~ n d IIV non-Mcndrli;n~ causc.5. nlllK,lllOll In .-I homeotic g?ilr corr~plcs111:iv pmducl I~c'n vcw Thuk \ V ~ I ! ( . r ~ , i t l l r ; l l \ ~ I ~ c t isi otrL~rfi~onall\r ~~ under- n dlipE~cntrw g i l e n t , re~ultinp.for rc,a~rrplc.l r r a f l ~ r v c n over I-.ire rhnt E~CILWI .'< i ~ p c ~ ~.it; \r lt ~i Ic'vc'! r~ ~ ~of thr n r ~ G ~ n i smid r n 1% 1wt11 four pcrfcctlv Conned w i r l q ~irlsteaduC hvo. lilnd,~n~~nt;lllv ~ c l . ~ ~ r i vmolt-cuI,~r c, dnvc rs non~ltlc\not 1 Tlic-sctlircc group^ of penes ~ c nil t easlt tltlirr 11, . ~ d a p t i \ ,17irl ~ c+pu.\tfi.it ttlc t c ~ c lof tlic g c n n n ~ I~t . s r c l ~ e i i cnnd ~ pre~umahlyc n d e for proteiilc tli;lt u \t,1rt\. %?TI m.~y a h Itc cr>r~tr,~stcd with t:cr~ctic drift, w l i ~ c h turn Iinvc t l i e ~ rc%ct oil other griier, u i c l ~ i ~ l i l ~l:u t ~ o i lr i r t opcr.irl-r I C t h It-vcl ~ octhc orprusrn I w t is ncm-ndayrie~llvd ~ v thost. cnn trolling stnicturc. Snmc qcrlec (1313 tmlal ,~J.lptc*ilti) ttvc. hlr>Yc.citl,~r 4irtt.c m.iv \vcll prrlvc to bc rhc third seyinrntnl and crpeclnlly hntneotic) c o i i t ~ i i,\ ~sl-tun c+liw cd\pcr ~ . t t i o n ;hut I I I < h.?rd to fcc Ilnw z.ipifiIi~glily cnnscnrative D N A wcltierlcr (or ~ t l o t ~ l ' radiation! CRllt .icI;~pt,~tinl~ ~llj!:I~r .ITICC thcrcfrorn. known 35 the hornenbox. It 15 1XO bare palm Iunc rvolutlcm N r ~ t l l c ruC tlri. 'tic-~~rral' o r t f ~ c'inolcc-ulnr drivr' , ~ n drnnstnnt in vzc a n d &ape. 'l'lie 11t~uenhnuw.lL ~tr.3tiyr~p1 thcnncc F[:I-IOM~!J. IIIIC~~~T~IH nco-nanvinicrn. CF 6nt dircovercd in homcotic gene? 111 IJlnco~rlriE~i \olllllj. rllc to tl1u cuu They 110,Jjni~,c.t'ci-.TAICCt l q~irstiurl ~ o f llow ni~icfi IOX4, but hnc nnw her11 shnwii to hc hnr~~nlnq+:tlu$ of thc \ ~ ~ r i . ~ l i n.? n i i SCL' I C ~ I I C ~ C~ C C J L ~ of W (.h~nc-~% llatLlrc d in varint~?i n ~ e r t c b r ~poupr. tr including prii-n~tkir ~1011r~ 1 1 dliio~v [I\LI~-II 11.15 , \ C C I I I I I ~ ~ ~ I l~ C ~ Cc~ c ~ it~ ~ mct~7n,lnx. i~c d ~ cl :t e d ,ti11 h u t n l ~ o ti vcrtrhmtcs, ~ i l c l ~ ~ d i ~ f l l l l ,17cc.1 act!r,~llvcon li-rr r \r.lcctivc .~civ.inragc. h u r n n n ~ Hoinrohns-coiit:\i~iii~p . ge~iecnlwt be vrr I ~ ~ r t h kcZi i . ; i ~ \ . ~ r i c k.I[ t l ~ um o l c c . ~ ~ lcvcl l . ~ r iliclucic nnclcnt. at l r n ~ 5r 0 0 Ma old. cincc they nrr found Thew trar?$pnwix, J I I L ? c:1111.~ tivrlv tliic nc.w ~txL>rnm,~t~or~ ~Eetail111 th d ~ v c n c l v .and tlicy alrvny F~ii~ctiir~i in wpneilttl r31e outce n ~ ~I ~ iA V~C t3 hc.~ri~ir:lipon m p d but coordin~tcd anim:lE< in conrrrrlling t h r r p o t ~ ~cxpiyrccxinn l nf tl:, ~ Y O ~ U T ~ C I I I JcTl C ~ .~ ~ l t '.!irrl c - , the ~1r1qi11ut' ~ I Z Wn r p u i body axis. Thcy arc. hon,rv~*s.prrqcnt 111 nnn-rrg- gct~eticicts ~ n t r o n n pJrtrrfiE l clmcntcd phyla ;and cvidcntly thcy d o riot nlw.~\ I'he c l i , ~ \ C I C C ~ ~ V1C npcratc in tllc snme way. It qecrnc t h ~ three t high1 tdkr Il1JI1\ con~crvativr h o l ~ ~ r n h o gcr,c~ s can t-rrol,t orljc r hlncrions, snd w. in thc cvolunon nf dc\~clnpnicnr,~l rcpreelciltel Gene ragulati;on during development l.c.~ltl~ngp PTC>CC\SPF 'tlir r;.lnIc rcpilator). dcmcnn arc rccnnlK q ~ c ,in t ~ ,~du)rcvF:,llilittcd into ncw dcvdoptncntal n~;~ciiincc* (Rnh cted to r 3 c r y . it clucq FII , I ~ :~~.cftrig L WI .1 prrdete~-r~?ined pnch- f 0%). Clearly thr hnrncnbox ccv~ccpth n ~o n c v - cvnlut~nn Micme! \V,IV, corrtrnllctl IIL \pr.r.~ficpcrlcs Thrrtl yerres nsc dcnt hearing on unc o f t h t m q u r nrcnr of cvclluriru>r\x*~tch~cl o n : ~ t l r l uffLl r i r e q ~ t a rrcqlrenrc. and nny a r v yalaeontc,lop?.;h o w nc\v body plan? origlrlltl ~ ~ ~ l t h51% ll crror in this prticcs, r uulil hc Ictli.~l.Tll r p r o d ~ ~ of c r d n d how they can bc ~nodificd. In thtr ctrtitc\' k01~1one unc grii~, IS Iikclv ti, I~avr.n st.qucnrl~ll ctlicct oil p , ~ l ; ~ t = o n t o lcupplic~ o~y rhc tirnc dir~\cilintnt~ v can r can be ir~tcyrntcdlvith scnctlc and n~olcclilnri1.1t.1 cprclc\ lulr clrv1r;lzc A 'c..I\c , I L ~ CIC !:r.iit'hc C L ~ I ~ O I C . TI~IIE o gcne ~ ' i t l iaddp r w i t ~Iicd r m . o r r~~rrlt-d oil; ,it , 1 1 7 ~n n r rleurlopn~cnrnoJe\ W ~ I tal rf.llrc c o l ~ r r n hthe. cxyrcccion {lirxhcr genes nt n