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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 I
vili
Contents
Microcvc>lutir>i~ Allupatnc rpcci,~nnii klctcrorllrnny T c ~ t ~ m~crorvolut~o~ldr)'i ng pntturnc Ar~aly
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 S n ~ a l ~l h c l l yfxsils. Prr-calnlinnn tr,icc Srls~ilc CJIISL'F nf thc Cninl~rinr~ 't.xplosion of lifc' 1'hycicochciliic;il facton Biclloyical hctors Rinlopcal c ~ i d c t ~ oc nc mrtazoan rcl~tionsliipq 3.4 MAJOTfcanircc o f thc Phancrozoic rrcord Dlrpcrsr fic.~rinno i invcstcbra tc lifc Cliat~gc:t~ in q p r c i u ~divcr\ity a n d Iiahitat Psol,lcr~~ntic car1y PaZacozoic f x s i l r M.~rinccvolutiotiaq ca'aun~s Clirrlatic ,ind ~ c a - l c ~ changes ~-l Eutinctinllq Pns~ihlccauccs of r ~ ~ n ' ;~,stinctinns r E:.rthhound tncch.anisiri~ F.;rmtcrrcstn;ll rnt.ch,~ni~rns L J ~ C<>i-tlc>vic~ati(Ash~jlliar~) cxtltlctlon cvctlt 1.di-e Dcvt mi311 (Frdcni,~tl-Famcr~nian) uxti t~ctionevent 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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I
Contents
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 ?... Cl<~i.; Hcsactincllidn 4.(1 Iri(.crtac qcdi.;: Arc-h;lcvc*yath;) Soft pan$,orf~~~iixatirlti and e c o l c ) ~ ~ * I)ictril,utio~r;~nd \tr;lti&mpllic uhe -1.7 (.;colo~~ual i~nportailceif sponges 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~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
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 ~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/<~
Orhcr I i i ~ ~ ~ l ~ f r r r n ~ ~ , ~ h ~ ~ b y llurn ir C:r.i~~iifi)l.t~~~~ 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 ~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 d
Contents
203 3 10 300
-7 1 0 273 21 9 230
27 1
771 --371
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224 224 226 726 37C)
Ld
2.10 238 23 1 2.55 2.56 2.1rr
237 Erl~inad~rnts
"I
111tl-odll~tl~ll 'h.2 Cl<~rclacq E ~ ~ ~ c l i ~ t iJIICI u l dthe ~ . 1~ 1 1 o r p l i o ~ 0 ~ c~ h~ ;a1r1~ ~ t of c nC ~ I T C ~ ~ ~ Z I O I ~ C tvolu~io~ 111l c c l ~ i t ~ o ~ d c Claf+Holuthuroitlc~ (:lnsr Z d r ~ u ~ ~ s t ~ r u ~ d c ~ 'i.1 Sl~ly>h\ lu jn Astcruzt>a Cribt.l.~rfA\terolJm 3ulll-lnfsSomastcrr>ldcn S t ~ l w l C)plllilr~iilcd ~~f 4t,1rtirl1bcdc '1 h ~ l y i l i v l ~(;ri11(wo~1 ~t~~ Cl,lrb Crinoidc;~ hlaiti qrouyr\ of criiioidc I',~larozo~c cnr.oi~ls h l c ~ o z c r ~at>c rcc cn t i r ~ n c ~ i d.s~: ricul:~tc.\ t Er olo~l-~ ) cl-it'toiti~ i F o n l ~ ~ t t oni ii L r1no1~i~l Iimc~lorlc'c " (I 5t11~1111l'1ll~llI ~ ! ~ l \ t O ~ l ) ~ l I:l,l\(cc I)~plr)pc~r~w .utd 1~hor~thifc.r.1: crctnidc
xi
xii
Contents Stnicntr~lchamctenqticr ~ T Qni~'tl1rCc C Clnw~ticntlon Ecolog C l ~ vRlnitnldca Rivcreir~;and functlon of h y d r o ~ p l r c ~ Clnrsiticat~onand c v o l u t i o ~of~ hlastoids E c r ~ l o g2nd dixtribu tmn of b l n ~ t o i d ~ 0.7 Sr~hplivlurnMnn~nlozon,orhrnx~lwc a l c i c h n r ~ n t c ~ 0.3 E\~olutinn ' E ~ r l i c~~~~t l l i n ~ dand ~ n tihlc~~ rn d i a t i ~ l l ~ Evnlutinn of the tube fcct Why pcnwmcvi Cnnvcrgcnt cvolutinn J I I ~i i i t ~ m i ~ dfornms ~at~ Bibliogaphv IJookt, trc.nrl\r< and <\+\.rnposia J~ldividualpapcn 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? ~ d of graptolite~ Ul t r ~ q t nhirc ~ c ;111d c h e ~ n l ~ t of r y gr~ptol~ tc pcriclern~ 10.2 C:lar~ific;~non 10.3 Rinlngical affiinl1t1t.q 10.4 Evnlunon Shape oi gnptnlitr thabdn~nmes Iymx~iiialcnti in Lm'ptolo~dt Thecnl qmicturc Cladia Stn~cturcof rcriolitid~ l(7.5 M o r ~d ~ graptolitcr d Iivc? Pa~qivcdr~fting Autotnoh~l~~ U s c otlmodcls in Intcrprctlng thc mode of life of gaptoloids 10.6 Faunal province? 10.7 Stratignph~caluse E~blioqaphv Ltooks. treatises and svmposia Ind~vidualpapers and other references 11 Arthropods 1 1. I tntroductinn 2 1.2 Cla~5ificatiolland gcncral morphology D l v c m ~of ~ -~ r t l ~ r o p ohpes d Fcaturc~of arthropcld orp~nlsarion
Contents [
I .7 'frilobit,~ C;cncr;ll r r ~ o r p l ~ u l o ~ ~ ~
(IOIIJI~~I!~~~~~
i+1
Ih=t,l~lcd t ~ ~ o r p h o l oofp trilr>bitu< Cut1cIc Cc['hdZotl Glabclla Ccpltal~vsuturcq Hyposto~rlc Eyt.5
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
Artluopodr Lobopotis CJtIlcr ~nvcrtc\,r.~tcc Slgnific~ncc.of thc Htrrgc\r Shnlc h u n ~ r 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~thogrnphiclirncctonc. Hnvann
12.7 I c h n o l n p Clnu~ifiratinno f trace cocrils Morphologic.11 m d prcccnratlonal rlascitication 13chnvto~~r.~l cI.~c~ifirntioi~ Phylngcilcric cla5sific ~tiola Urcq of ~ c l l n o l r , g C;crlinicrit.ly ctlvironlllent Crratigrapl~y Fossil l>chav~our Fiil~lio~~pli~~ Rook<. truatl'icc arid synlposi;i (e.rcrptiona1 fauna<) Inclividu,~lpapers .itrd othcs rrferenccr (cxccptioiinl Rook<,trc,lncc< ;ind qyrnposiCl(Iclluolo~y) Ind~vldu,~l p ~ p c r r(ichrtology)
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1 hc f i r liqhcd I 11c\3- 01
Systematic index
h c t all1 tcxt for
General index
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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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I.I(.~IIT
n o record of terre5trinl , ~ t u r t ~rnltll ~ l r rnuch I,~rct.I n
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.lc<, 0s ~ ~ l ~ i t f0~111ed ~ ~ t i ~~r~vcrrchrnte ~ ~ ~ ~ lquups ~ 3 wurc c5tabl1rht.d, and thcy thrn Il.nt~aI?II, ntbl~l.ir dirt, .qttcd nr tlle~t.roll~dedthcy drvcrq~fiedand cxpa~ldcd.Somt. of tl~c'iuorgL~nisrnc I ~:c.rt.,l, c\ rrttu;rll\ ionnlng rhc pl~ncrg ncqnircd hard rhells and wcrc cap~hlt.of buns focOt' ,111 rllt. itil~cplnl~ctqIn thc Sal.~rSyktrnl onlv slllzrd, and only becausc of this can t h c r r hc any 1. rth. ,J< lj: ,IF k kll0~77,~ ~ p p O r t. Sd v d n c c d I l k , clinncc of ~itldentnild~tlg thc 11lrto1-y of-invertcbratu .I,l$.i;h ,ir ;hc tmic i ~Lvntlng f ~nlicli lntcirqt ha< Ilk. ':]I ~ t l t * r . ~ hv t n l thc diccav~.n of orgrlr-~c~ n ~ t c r i a l T h c stmtitied ~cdlli~ciltary racks laid drrlril ~lixcc 11 luyirtm'5 rntcllitc Eurrip;~.It ir. ho\vcvcs. a qtr~h- t h e enrly C a ~ n b n a i ~ n11d , h i l t u p througl~clut thc t l i t tli i t ]ifr ~ c l t l I:artli I ~ c g , ~\.cry n c.irly ~ t ~ d c c d ,nbllc>leof P!lanrrozo~c tltrle. nrc dictinpirchccl 1s). .I 1rh111tlic t i n t 30";I O F tlw ~ J , I I T C ~ * F l~i~ltm-y, Tl~ert. r i c h l ~ c r i t a ~of c the fossil rcunmnq of thu invcstcc r ~ * r i ~ , l l{n>\ < \ I T T I ~ ~ OL .~ ~ I I I \ I I I( ~ c t c r, IjI ~ I L ~'h111rE~rntef that rvolvcd thro~~g!rsucccssivc Ii~qtnncnl ,r~.:.tl,rlc,rc', or L-ynt1ol7acrcn.1)i t ? rock? 341 It 1 ,Ma pcnndq; their ~ t l i d vI$ 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:<' ,:!,* to 2 . 1 ~ . tllt*cr ,II-C all I I I V L = ~ C C ~ ~nrdmal? ;I~C T h c ~ t n t co f presrn.,ition of fos~~lfv,lnr\ ~ r r a t l y , IhIL.!~, h,~c.Lhrlnc~. A11 of thctr~arc i t i . i n ~ ~tlicrt' s: IF dcpcnding 011 t h r ~ t n i r t l i r ca11d cornpn
js.
S ~ I ~ \ I I
i l ~
oripn;ll 4irll, tllr nature ,irlA grJin ~ i ? co f t h c c i i d i ~ ~ -tlnnr durinp rhcw crncioli periods \vun~lJprol~:~l~lv nr d c ~ t r t y c d- .1not21~rIrilut,~tiur~ or] i n s s u d ~ n ~ rtlir ~ ~clicr~~iial t, co~lrllttorif. ~ttl ~ cti111-r of br tr.in~yort.c~d thc n d c q n n q of thc fnwrl tr.rt>rii. ~ t ' d i ~ l ~ r n t .~~i tt dt11c l ~ l S~~ I ~ W C ~ I I pC rI I~~~ c r \ ot'dia~lf~ O n thc rtth~-rII,~TIO. w ~ r n c . rn.lrltic i n ~ c r t r b r ~ l t c c g ~ n c s i s (clrcii~lial .rilkl plivsir.11 I-hnilpei) t3k11lg foiii~d ttl ccrr.ltn rc~cbtypes 11,ivc. i ~ r c r ipr~-\c.n~t-il pl,icr 111 the rock ~ t t r -dr c p o ~ ~ t r n n . Thr qtuilv o f tlir procrcscs I e b ~ d r ~tc,l shrriliznt~nn nhniidnt~tly.in{[ in cxcluiu~tcdctsil, \u t F ~ ~tt 1% porri1, kno\vri JF tfipllonnrny. I n n l w t s:wc only t l ~ c hlc tn )nF~.r inuch ~ h o u trhcis b i u l o p li-r~~-rit l l c ~ r -CV U I ~ h ~ t dpart\ uf t t w ~ l,~rltrn,~F~ , ~ r cpr~-rcn,c.d,n ~ ~Tnrd rci-n.~inr. M,IIIVof tlir ~ ) ~ ' ~ ~ - ~ T C~ ' ~CC ~U II ccwnc clr k i - ~ ~uilty r ~ rctlirncllt\ 1vit11 a high t l ~ c l ~ c1, 0 b r f0~~1117td. ra1plrll'rlirl,~IIF 13(1n11,111y .I prr- from 11n~r.ctc~nc.c rerlurcltr. rhc cofi-llodrrd r l c r ~ ~ in ~ n rllc t ~ EILIIJJ. r.11cnrcous c-(>ntr.nt.I r l ~ h c w(Fig. I . I) ttlc o n g ~ n , ~ ] I tc J I T ~rhurc irlrni, \ ~ 1 t t i tlnu or$anli slirll<. d ~ d Trot calc,l~-coucrhcll\ nlnv h c sct.iinctl in 1 1 1 ~ . f i w ~ m i i n n n ~ l runJli.e l~ d~.~gc+rlc.\i\ ,111d hcr~cchnvc lcfr littlc with rclntivcly littlc altcrntioei, dcpcnciing Ilprln thc r ~ < die wriirnciit , ~ thc t tlrnc o r nrt cvitllr~lst.r ~ rllc.11f c\lwn tc>g~-thcr tllc. c!~eiilical h c d ~cullrditung ~l111ci)r ,IEI tlic w$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 olclcTI ~0111lifc b c f n r c t l ~ cC:niiil~ri,in.4ii.citIlcr ~ l i c h'xvindow.l.'i \ (authigencsis) ns \vc11 d'i C C I T I C H ~ . I ~ ~ T ,iricI kiio\vn rn Middle Ca~nPin,in i-(ir-k5 holn Rrrticli pactlon of tllc rock ilithification), and rl11nn# . ~ n c C01trinhi.1. \\.11~-rc In nr(ditlot~ tn hic nonnnlly onc nf tlitrr proccwcc tllc fowrl~I I I , I ~ hc. ,~ltvrrilo r cxprctt-d trilc~bitc~ .it,d ~ ' ~ . I c ~ I I o ~ c ~ Ctlicrc ~ F 1s .I Lmeat dcctroycd. Sliullc zhnr nrc nriqin;~ll\ of' c.ilcitc prrFigure I l C.I~CIIIIII railer o f ~l~r.cl and dii~i-chtlFcd ~nlnt;ll\ - scrvc hcst; ar;l~nnitctr n It';$ ~ t d 3 1f n~n ~ ot' she11 was n cc.rt,~inIivirtG orghrnsrnk (c.g. spun yes. \vcrrirlq. jdl\-ti\h, ~mnllshrimp-like crr.1- c n r b o n , ~~~r c r c t c dI ~nallyof i z calc~tr ~ ~ l rllir~iig turer . ~ n ~.ulitri:~l< l of qtiitc ~ i l ~ k n r ) ~nFtii1itic.c vn - coral<) and i~~ ~ f t t ' r l~ ~ n ~ ' r t 3 l lt43 genetic inlr~plctclr.. l ~ l l 1 1 ~11ketviw 1. lll~lll2lTl~rill~. casts) T h r tils\ll rrrr~rtF1.r. ,i\ .i gu~Ic.rn thc c v o l u t i ~ nof nftcr its indurntion. Moulds (oitcn inlcc.~ll~d the ord curt:+cch\c)f thc f i > ~ ~111Ri' il JIICICIII I I ~ L . . ~tlqlit'stit>n~tL-l~ liiiitted, patchy and o f thc eurc.rnnl ~ n intCt-tlal ..-+-- I t11c dctail~ i ~ l c o n ~ p l r tU\u,~llt r. c>nlu rllc hard-uph~ C ~~O C \ S J (. ~ r l cI ~ J C ' ~ ~ ~ fO Co ~~C 111 tlic l>ior.~ ( 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 iroii
a
Occurrence of inyedebrote !ossils in Phanerozoic rocks
5
recrystallrzea c a l c l l ~
"
.
CUP F I possible rocerses OF foss~lrzat~on of a bivalve shell ( a ) original shell, buried In mud [leh) or carbonate (right);(bJthe hl *or m t ~ ta ~ , gun& in o m b a n a s d i m n ? m d r o s preserved ~ntoctnha man or o small cmtollired rtch; (c) shdl *rig~ ~ ~ ~ ~ o gnow o nrerytnILzd ~te, to m i c i t e rhlrh dertmyr the fine ~twcture,(dl original ~ I C I C &dl retainJsurroundangci dim - P C core 01 SIIICU, (el u s~lrcorim growing o n the ovtslde 01 the shell, (f) tectonrc d~stortionof o shell preserved in mudstone; (g) i e I ~ r ~ s e r wInr lmud w ~ t horrginol shell moterral leached away, lew~n an external ond on ~nternolmould, svrround~nga rnwdsl~ne ore I;! o coicoreous concret~ongrowrng round the shell and Inside I~f tie orlglnol cavity was empty), wlh p k h e s of pyrite rn ploces; I fonttane re,>locernentof core ond port of shell
B
q?.icr~ 111 tlw ~ h c I l,lrcm i~nprryintcdtv~i-I1 I ~ I I ~ c ~ I1i-I ~t~, 5.11d to hc perminetalized, I* IIIIL tilt. ITOM t h of ~ C Cuid.tl7' I ~ ~ i i n ~ r ;;I[i l
\tr\
5
rllc ulica 1ncre;lqer ~rlarkctil!.. nnd thc s~hca G O sclc,~c~d ill r r ~ v e throng11 l thc rock nild precipltatc L\~IICTCVC~ r l l t pI I i q lowcr. The zii5idc of 3 qt.,~ urchrn cieu,ryirig undcr dl6icrcnt c a n d ~ t ~ o rwould i\ trap juqt cuch an intern21 ~mcrocnvironrnc~it. witl?in u ~ h t c h thr- 51llcn could prccipitntc nr a gel. Tuch siliccou< corvq rctnin cvccllcrit tc..~tur~*q, prc\crv1ng the ~ntcrnnl ~ ~ l n r p h n l o gof y t11c sIlull. i3n chc ntllcr I l a ~ ~ r~licn d, 111.1~r r p l a c ~C,IICIIC ;IF very t & ~ nrhcll ovcr tllc w r hcc of d fosil ns a rvsult of wrnc crrr~lplcus~irKitc GI
1~~1.1iiJ TIICY1 i \ ~ c 1111 ~ 1 ro r 1 1' ~ ~ d ~ i i oi ft cnlcalc~~it tIc~.~rlcrl cvprefsiot~c)f the ~ u r - ~ ou fctllr: few[ and, ~rlud,tlong \v~th111.111v y v n g c c , w h l c l ~~ c c t c t e d m c c . t h t y ciin hc freed frorn the sock by d t ~ \ o l v ~ t ~ g I!IL~IIL*\ r l f htoqet11~-FIIIZJ :I ~l\t'lc'toi~. I n alk.~liiic tlic I~tr~cqtor~c w t h hydrochloric ;rcld, 111d1vidu;ll \II,
I:
-1rrltcrl
dririiw,
h . ~ ~ t r - l - i a~ICC,I\'. l
rhc. srrlubll~t\.of
thrrc d11iir11~1011$. SL~ITIC of tl>rn ~ o C~S t~ U I S I ~ of C 311
/ '
6
Principles of palaeontology tr~lolxrt.~ .lnd bral.hiopcd< ~ r c LIIO\I . rl Froill ~nntc.ri,rl rciilaln\ u i r n m c part, 1i~unlly n ~ h c l of l r k c l c t n ~ 0~1, .j slit-I1 a\ rill?, ~liicc-l~ving urganlsm: (2) trace fossils, whir-h ,lrr A rclar~vclv ~ I T I L - O ~ I ~ I I but I ~ ) I IC ~ T L ~ I I I F I T C 1 * no~1 I ~ E ~ track<. tr,~ils.llurrcrws or uthvr cvldci~c-cn i t h c .irr1\.p r c ~ ~ ' r ~ ~ ~i~t t 'phosphatizatio~l. oi1 C(rin~~tirnc+ t l ~ c it? ot' n r i a n i r u d (71' FOTIIICI. tliilet - C ~ T T I C ~ I I ~t~hLc-rF; ~ cxtcrrl.11 r~,:,lr~t~--d~clltd arc. thc oiilv q ~ i d cto thc fi>riilcr prr.;cilcc nf'.iofinnilnnE~,Inn) I3c r ~ p l a ~ codr ovcrgrolvn bv .i thin bvdicd anirrlals 111 .I pnrt1rul.1~c i ~ r ~ ~ r n n ~ n(31 ct~t: s h c c ~47f p1lo~pli.tt ~ LOT , thc- I,> trcr t n ~ vrciii tbrcc :it> chemical fossils, r c l l c ~nf b~clgcil~c nrganit coinor1~111.111~ p l i o \ p l ~ ~ ~\licll. t i t - 111 ~ l i cf n r ~ qtttidmt~n ~ ~ ~ r poi~ndswhich IIIJV b r drrccrcd gcnr.f~ciiiir.~llyt i ) thc csrcmnl iO17ii r r f t h C I,~11jy1, ~ ~ C C I S C I Y r c p l ~ c ~ ~ t c dtllc . r(7vks. Altcriintivcly J F E I I > T ~ ~ . I ~ I L hllit~yof tlic i i i ~ ~ r i ot' l ~ r , At t l ~ rIleart OF in\-crtcbntc pnl:~cniitc>Io~~ ct.1nil4 thr \lir.ll r t ~ ; ~ form v .I r-orr, yrc-king o u t r~ltcrn,~ltaxonomy: thc clarrlfic~tionof f I > > ~ i 1.111rI rnodcrn stnicturr5 1t1 rcmnrknldr* ~ % ~ t . S i iL l .I LprCsrw.lti~i1 '~~ F\ A ~ I I ~ I ~~~l iI tCvorrlcrrd and 11.1tuml p(111pi11p. T ~ I C W proL.ahl\. .~rsoc~.rrcdwrth Il,ir.tcn.~l .~~.rivlty dirrctlv gn)~ip111gs,l u ~ o j ~a% ~ taxa, i i i ~ ~ r bc \ t ilnrncd .~nc nftcr t h c dcnrli nt'rlic- ,1111111.11. M . I I ~ c\ ~ l ~ Canil-ri,aii ~ l l cirraugc-din n 11wr~rcll:alcyqtcm 111 xvhirh t h r ~ rr.l.~r fos~115 lin\,c brrn yrt-\c,n~cd b!. p \ ~ c ~ \ p h : r t i ? ; ~ t itoi ~o ~ l ~ l i ~drr p s rn:~declcnr. a n d ;IS t:lr ,I< pclrsihl~,murt (Cl~aptcrs3 ~ n t2). d hut much largrr fu<\~lqillay Ilc hc cccn I l r evnln~ttonnrypcrspcctn0c prt.wr\*cd nlco, 67s r.r,~ll~plcc-ru11cc. cnrh Chnptrr 1'. L C . rt~"ll I qpccicq corltillcd to iliir parr of thc ~ucccc5ionnnlr and rcprcscrltirlg tlir timc ~ v l i c ~th,>t 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 r ~ i r ~ d c r p ~ i h r r l (:1 ) body fossils. ir, ntlwr ~vordkthc actual s w i i ~ l i ntlic ~ g c o l n g ~ record ~l in tcr1in of rr.11 lriirlr
I ? The wrlovs sukdiriplines of palaeontology
iI~urn
I,tron r i pcnnds o f ~ n i l l i o lo~t ~y c m ) r;~thcrth;ln I pur.-I\frr.l,~tn.c binlc. 1his Ir o i ~ l ypnnihlc a t ,;*FIIII ilor~zoi~~. I i o \ ~ ~ v rand r , for p r ~ c t ~ c apurl
l Y 5 h . is concerned 1 ~ 1 t hthr rclatioi~ship~ of fosd aniln~lsto therr r~ivironmt.~-!t.both as ~ ~ i d ~ v i d n a l s {autccology) nrld i11 the faunal co~nmuuitics In r,~.:, tl~: c~~:c.nlayc,~l timcrcnlr hn~cdon l>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+l[llrrlt' peochronolwp;y. J s r ~ ~ a lhtwtl n f ~ ~ i a lprc%crvc.d, lv but only thelr hard FIICUS, therc arc 7111~ ~-TI;~IT-~LIcJI ~ t ~ l hd i3 3~ hmnng ou wlint liiay he ~~1.1tivcIy ~ C Wm-~ys i n wllich thcir biology and Ilft ~rrr.~olgcochronotnetry'. L1v counting daily Railits car] br ~ i ~ ~ d e r s t o oStudicc d. ill functional :tslirfE~ nrlp 111 cxtil-rct corals and hivalvcr. infornumorphology, howcvcr, which dcal with the Inter: r ~ ! n l1,1\ tlct.:~ n l v ~ i ~ ~lr,c~l : ~ r ~ ~;ipon i ~ g t l l ~nurnher of pretation of the b ~ o l o wof fo<~ilizcdskclctons or i n > 111 III~IITII~IT 111n11th,111d yc:~r111 : ~ n i - i r ~t ~ i ti l ~ c s .h m l c ti1rc.s in tenns of thcir original f i ~ ~ l c t i o nI~avc , 1111~1h.1~ Il~.lpcdto (*oltfin~i ~topliyritalcstrnlntcs rill hccn ruccc
I ,
15
livcrl whcn the. sr-rl~mr*nr \\,a\ hr-lny rlr.pnr1rc.d and 111 Tlicrc word? cliould ~n.ikcunnrcly clc.lr tlic ti111Inteq~rctlrly thr. c-ontr.lilpor 11lc*o11ccn\ironIncnt. darncnt.11 trltllrc o f mstlnomv. For, n k ha7 rrt-rcn hcct Finnlly. it I \ only h t t l ~ r~nrcp-nt~on ni tflsoiinmlc wid, to idei~tifva incs~lc o ~ c i t irl ~thc' t i n t rtcp. ,ilv Ci~t.1rln loi.~l Cllinn~thnr rlir ~ l t > 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, prrbils<.ctmttg;~ph\. Thc fi~nd~~rncnr,ll unit n f t.lxolrtln1v r\ t11c qpcclr# Chcr~iic,ll r.r>n~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 ~tT,~\s:d ripmi pra %tchvn s ~ - q l r ~ n r i I1-lln\. lg clnrrlv rel.lrcd w hov~K I ~t \ v ~ )o r r~101-r ~ ~ 1 , 1 tOc~~~ lJ I I I < I I 1I $1 ~ v r ktiowii. I3t' ,111 thc ricscnhcd 'cpcclcs' ot' liv~iig,JW rlivcrqrd horn ,L ~-l-rmrni~~i ailcr-cttlr, nud to ~ornc ~nnl<,l ~ n ~ v ~ vonly c r , ~ 1 7 i l l 1 r n rivtli .Ire 'good', I;. cstcnr thc .~v.~~l.~l\ltrcChni~luc\c.111 ix-. ~ p p l ~ ctod rhc propcrly drfiiicd, c y c c l c ~ l ~ i f i ~ r ~ i i . ~ilpnu t i o i ~ tlit r r ~ t1 r I T ~ I I ~ ~ L T I ~ C ) I ~ ~ _ ~ ~ C - ~ I - ~ rcpmducnv~. ~ C I C ~ T ~ ~ ~ ~ ~ pl-cfrrci~cc< ;II~~ n f t l ~ cotlicr tivc-q~url~< I. trclln?clur\ can hc ~iw.tlrt) IIV~CL t prot~.il>c ~ n polvd .11I n.ltlii-.~llyocrurnng .Inirnal pop~tlanniiric p r r n r ~ n ~ ~ * h , i 111 r i dh ~~r i ~hcllc l htrr. thl- rhc tnntncnt. c.t,ly docui~lcntcd. ~ I i ~ l l younycllr t l ~ , 2~ Ma r ~ laavc prnvc.rl :um.cnal.Ic t43 Tlic rlifl;.rci~r~ntionnf i n o q limn2 3nc-I .rll fn\\l a n , ~ l y \ iTli~.rc ~. .lrc .11~opm~tll\tny C E C V C I C I ~ ~ ~ C I I,115~) ~S ywcicq thcrcforc Ilns to hc based lipr>ll ot11r.r- .In, 111 palact71-~ocIicrn~\tn. ~ r l r f ory,11111. ~ ~ ~ ) i - t ~ ~ tcchnirally ~ i i i ~ t y Icw v,~lidrntcria. which arc .~pplat-.~hlr> to thr. f i ~ i 1rr.r.~td. t h a t l q h COf tlle5c hy fir tth- inmt ~mpnrtnnr.r.spccinll\ 11 t11l.s~.>rcbcvc~ntlrh~,ut-npr nt-tl~rrl,nok. p a l , ~ c o n t n l o p . I < morphology. rhc kncncc r i h n , slnc-c Innst nnrur.11 spccir-F tcnd to be cntr porr-d r.rf ~ n d i \ n d ~ ~ of , ~ l csitn~lsr cnouvh rstrni Taxonomy ;zypc;lmncc to bt- ~ d r n r i t l ~ b lnrc o!"rhc vnmc klnr K h + h n p l ~ r l i ~SnPgC C ~ C S of l i w i i ~niiimnls ~ by moq-llo lnpcnl c r i t c r i ~nlnile I< ilot \vlthottt h.17nrdr. c q i r c~alEy\vl~crct11c ~peclcsin qucalon .Ire ri~ni1.1~ all1 cl{xt.ly I-el,ited. S~~pplclneil t a n ' ininnnn tjcln. a11:11I rhc ni1.1lyur of spcc1c.;-~pccific protcitlr, may he hclp 111 some carer whcse thctc 15 good rc,lqnn h t t so he
L m
Divisions of invertebrate poloeonto!ogy
9
4omenclature and identification of fossil species conlparing the cpccicc point by point. S n t r ~ cn i thc lit. fnrn~ali ~ o r n e i i ~ I . ~ of ~ uany r c ~pccic<, l i n n g o r sprrlrs m3y prnvt to bc ~dcntical u~it11 nlrc.tdv 11, taxonomst< Knllu\v the biolcrpc-nl fi'ftrrli of dcscri\>cd spccics, o r rhow only rnlnnr \.ari.~tinn<>Fa 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.il .4iid leopard, rcq-cc- h.ncd upnn type specimens, which arc n1wav.i rherc,~kcrkept 111 n InuGcmm o r rc~catch~nctitutc.. *. 111 fulF t;lronotnic t ~ o m c ~ ~ c l a ~the i i rn~lthnr'l, c~ ' .ltld the dLltco f pllL7l1~,ltl0tl 31.C glVrl1 R ~ T t Cl l~ ~ U ~ u a l l yniic of thew. tl1c holotype, is. ~ c l c r t c dnr thc refereilcc spccimcn and fully 11luc;tratcd;coin<:, e,p. I:t,lrc irlrtrx ( L i t ~ n a c u177s) ~ (rcc hclow p,lmn\lc dctnil 111.1y bc ,~ddcdfrmn o t l ~ c - rs p c c i ~ ~ ~ c n r ~rthcl-d i c c u ~ ~ i n n ) . pnlncnt~tolo? I[ c n n nc-vcr hc kiia\;ll for c,~llcdtElc paratypes. Thew arc v a n o w othcr kind\ L L I I J ~ \vIlcthcr ~ n poplil.lt~orl\r.trh n pnrhriilar mor- o f n p c spccimt-ns; fix c x . ~ ~ n p l,cE,neotype mnt. bc ~ h n t wnq n ~ rcprndtictivr-Iy tsolntrd o r not. Hr.:~cc r,rcctcd ~vhcrrJ 11nl01ypc h n ~hccn Inst.. o r ivhci1 ;I s17c~1rs h n j : rcdt-ccnhed In fullcr and 1ttorc 11p'hr ~ C ~ I I I ~ I C c>i<prcirc >~I 111 pnl.lentrmln=, a< In rnosr Ilnnr: spec1ilreil<. inuqt b-. b,~cednIml-rvrr.nnly rlie hard t-ccrl dcngnntcd. A i~tw gtnur wtll ht' d c f ~ p ~ t as c dgcn, new. 17): it\ ~pinrof' t11c t'mcll ,1i)tn13l Art= p r e w ~ c d .3114 much nritlior. t l i ~ q follnwing thc gcncnu rlntnc, a nvw 11rr.tl1l cl:~tn1 1~q ~;1111511t(1A c.~rt'fi\I~ Y J ~ T I I I I . ~ ~ I C and ITI I!lx~rment,~tic~n nf .]I1 tllc an.~trrniiral~ C A ~ L I of ~ C tlit' C sptcic\ ns sp. nov, niid n ncw suhgccics .I< FCP. 1101.. Thr ncur qpccies must hc narncd and allocatccl t o 1 ~ 1has 1 to be the maill giidc 111 c.;tnhlichir~gthnt I~IILI ~ P C C I C FI \ rlifl>rtwt irnrn ,I rclnted qpccic~.In rnrc nn cs~'itinggenuc, or if rhcrc I$ 1 1 0 dcscsil~cdKcnus t~ ---: t h i q ran bu ~lipplci~~r.~ztcrl b\- .b cornp,rwon nf to \vhtch i t pcrtnin~thcn a ncrv gcnuq muqt ~ 1 s hc l:herr~i~tn. of thc \hell. .I< Ilac provcd eir in,ih 1.r l onsidrr.ihlv lew, incrcn5ilig urldcr a dlffcrcnt gcnrric tlnmc, thc o n p i ~ n nutl~or's naiiic i q quoted In p.~rc*nthcwr. In 1917 F.R. nrk tit‘ gt,~ trr E I ~ ~ > J ? c ' ~ ~T v IlT~\ ~ t '. fp-nups ~ nrc I V I ~ I L ~ I ~ J I~I ~~ r i c l c l r r c ~S~~ C! C I P S .\ \ ' l l i < I ~ 1111lqt th~11 Cowpcr Rccd, t k c i ~w o r k i ~ l gnn Ordn\+Fc~an:~nd ~dclmtirrl.This I F doll r bv r-oiw~ilr~ I I C p.rl~ennto- Silurian hrach~apodrof thc Girvan clrcrricr. Srotl:find. recogived r n m y ncw spccics. C)nc of ~ ~ C F had L ' : ~ i r~il~ o r i o g r o ~r~ ~. ~; ~ I ~~ cCPO; I ~ ~ ~ I tkrnilrd T U I ~ ~ lu~r(.~l d c ~ c s ~ p t ~a ~oi~d~~lcl u < t r ~ t ~ of r r nprcv~ously s simrlantie~in r ~ ~ o r p h a f o to w Cllorrr~r~~. hut it \ I ~ J ~ :trlhc~l I~r.~clunpctdI a ~ a n a ~of slrlulnr ayc, nnrl s~itllcienrlyrlifl2refit to 7nc rcgardcd a 6 A
~ I
ncnr ~l~llgcnnis; r l i ~ \1s nrnrtr-n U/rt~~icrr~ (Eorl1trtrrrr.r) Thcrc nlay hc vnriuuc ~ t ~ b d ~ ~ rofi ftil~l ~n ~ r l t > ~ i r : t lf c IL)17. WIIC'II111 I q I X thc tasnnornic g ~ r n r sc.g. ~ s u p c d ~ t ~ ~ i lw i th. ~o r. d c r ~ ctc.. . ,mrl I i\ Il1c.h tllc, p r n h l e ~ ~ of ~ s C/rtvt~,tt'.r211d un11l;lr f;\rrnh wrrr tnln grnnpc thcrc if pvun n cnrc for rrccrinc pl~ylnycn addressd b\ tJ.T. I n t l c ~ , the nrlv S~rpcrtirnily 'rupcrphyl~', Tlicrc drr only . ~ b n l .3n ~ t phyla 111 tlic 114.I1~ 1 C'\Cl I ' l t . c . t , ~ ~ ~ ~ b o n r t ~\!.ns ~ c t - ~crcctcti to . ~ ~ r o l n m t ? . d a t arlitn~l ~. kingdom. and nnlv ahont n do7cn of tlirrc. Illc*,lnt rllc r d r ~ ~ i !,iiic( , ~ Iilnny nthcr rc1:lrcd hmcliiopodr. At a c.9. Mollurca and Rmrliinpod,~.Icivr- nny fn.~~11 Ill( I1 . I l l i I Inter st.~qeI;tlOr~~rt,!vi\vns c l c ~ . ~ t ctod t l ~ cmnk ut' a runlains. En !I'TITI\ I Full gcrius. I n tllc moxt rtcrrrt trc.itlncnt, l>.A,T. En tnxoriunlv h i ~ h c LIXJ r Arc t i r ~ ~ , ~dlil~v h n p ~ i ~ h r r rt!llr\C* l LTc H a r p e ~dcwnbcd n Earsc inuna frmn thr Glrvan hy thcir suffix (i.c. -ca, -3. ~ t r . )AF . ,171 c-xamplc, thc oi(lcr o t d~qtnc-tof Scnrl,mJ. 2nd ~ ~ r ~ t cl hi t~~hc n recop-rlizcd folluwitlg docurncritq rlit= clns~~fic~tion of' tlic ?I l ~ ~ l ~ c hvo w b < p c c ~ cof r I:'. i l r l r ~ i ~ r r Ear t. ' t o n ~ c \ v l ~ ditfcrcnr at CSrdnvics,~n hracl~iopnrl Fnriroli~ta ndru.lrlr pi?rf/lc , Llct !ding r n s r q nnd d ~ s t ~ n p l ~ r h chv c l rllliinr rlltFc;.rc~~cc+i In rcfcrrcd tn carlicr, accnrrling to 3 tnsnnnrnlc qchcn~r fi l \ ~ ~ t ~ ~ > n r o r p h n l o ~ Thccc. . 111 I-larl?cr's ( 1 08") ~ ~ l l o ~ ~ u p a 111 p hwhich , thc aurhor of thc taxnn : ~ n drlic ycnr nt 11~1,: lilt :I 11-1 ,ire \i7ntrc11 ~ i o d ~ ~ ~ t~~ it d - r~c ~poctlI3urrlcril 1806 1'1 I L L - rrtucl ~11tllnn1rlT1 refrr to rlic I.~rter.111 full. as Et~clivx~r~tcsSubphylum Khvr~clioncllifonncaWilliam~ct , I / . 1 !t-~:k c ~ l ~ f , ~ ~lr/~*r ,rmiilt ~ ~ l r r. Hatprr I ')ti". or 111 d l ~ h r ~ v iforrr~ ~ t ~ac~ i 19ofi I'~'t0). 1:. d t , i I ~17 , p ~ i ~ ii ; ! : Class Stropho~ncnat,~ Willinm~cat #,I. I Ylh !.or cc.rni Where. d u e tc9 ~ndrtYrrt-ntprcscn..~tiorl.o r lack of CJrdcr Strop11r)nicnid:i a p i k 19.31 , t ~ ~ clrlgcc rl nn up-tir-darc 111onopap1i~~b.lcc, J ~pccieccanriot S t ~ b o r ~ iStropi~onicnidir~.~ rr apikl 'U-1 1 111~1ot. CITIL>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-snnnrn~c~vtrrk .ruth as tlriq Irlilht foUow a clgy). Tlik is always so trnlun, for othcr rcnwnr. t 1t>\v l>cqt t pnrncuI.~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
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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<~it;catinn,anti ir 5t111 C C I I I F I C ~ ~ *tn ~ Chc ' ~ rhc bone<; rlls puslogi~tr. whn chnmctrr [or all vertt.bmtus ~ ~ i rl not, d o f COLITCT, :.'Ir.c much ci~~p1in~i.i o n t t ! t l r 111 st.t.kiiip .lnuu.;tor- indic~tive of any c l o ~ t -rcl,ition\hip bctivcrrl a n y i: cndniir rclarinn~liiy~ (Hcnn;, I QS4; Gir~gcrir-h. gmup o f vcrtcbrate.;. What is a pninitivc chnractrr l l'h1) fnr all vertebrates i r ~ 3 Cf O I I ~ Co dcnved c h a r . ~ c t ~~i r Itlr toillr t n x ~ ~ t n n i i \I~owcvt'r, t~, tJrc ~ i r r ~ c n . i i n h ecot~ipdrrd ~ to i~~vcrtrhratcc- u~l13potnorph-y nnd I! %t1h,jr~tlvit7: ~ v l ~ l arc c l ~nlnicxt m c r c . ~ p ~ t ~inl cany c v n ~ p l e s i n m o ~ hthilr v dclinc.btc thc rclntlvr xt.~t~is IJ rliil~rrc irnpartnncc (ag.1111 s ~ ~ L ~ J L ' c ~ L Yin~ I rlrclcr ~) to assurncq that d;vl~otomou~ splitting had occurred t,lln rneanlnghl rccultr. Hcncc the olljrctivity nf In cnch lincngc 2nd c o ~ r ~ p i l cnn r (tinwc~gh:hccd) ~ncricaltaunnorny 1% less th.m i t r n ~ g l ,~i tp p r ~ r . TIIL,tllrrd ~ c h n o rl ? l l e ~rlporl phyloqciict~ccriten;~ I I I I I ~ . cniph,~s~zing that icaturcr rliarcci hy c\rganisms miftct, in narure, a h l c r a r c h ~ ~pattc-rr~, l cvidcnt 111 d~~trihtitinii of U I I R ~ I I C C C ~~ h ~ l r tnrnonpt .3 orpnIs I F ~ ~ C ) \ V I IA< dadism or phylagenetic rntematics - n SCIFQO~ t'o~~ndrri hy tllc C'T C M ~ R I ~ sfl>rnolopstWilli Hcilrug ( I W C I ) ,i~ldWJ+ \ O C ~ ,rlllrd v r ~ m r o u ~ lto v pnlncoutc~lnp(c.g. Eldrrdgc 111 Cmcmft. l Qt;1)). 111 t l ~ ct.drlv d a y of cl.~diitlcc srr. wcrc iiinnv d n u h t e r ~(ir~clitding,aq I havc tn m m thc ~ ~ tpresrtlt . ni~thor).Lj~lt,
12
principles of palaeontology character datn tnktns. T h e r31urc c l ~ n r ~ c and t ~ r ~ opporcd by k d l r y , 'E .'r%_S). tlic succcssjvc ay-pear;lrrrt character stat<-T tl~cre JTC availnl.lc the larger the o f nxa in ~tratigrapliycannot bc denied ar an eqrendatabnsc, and large dLirabaqcc arc. ~ifrcn routtncly tnl qource of d,7t41, hnwcver itnprrfect tlic I ~ W I ~ ~ ~ O C C F S JC ~I the ~ conctnlctitm nf a uladognm record nctlially 15. T h u ~a5 C;lnpmsh ( I 'ju0) conisprcdrd up by u u n g oollc uf rcvcral cornpurer pro- mcn ts, 'Time i q a fiindarttc~ltalJ ~ r n e ~ ~ in r ~ cvuluon gtanis. T h e PAU?' (Phvlogcnctic Analvsiq W i n g bonar)' ~ t ~ i d j ~ e ~t, 3~ major d god of p . ~ l a e o ~ i t c h o ~ P a r s ~ n ~ n n vprogram. ) fur C S Z I I I I ~ ~ C 'is, .1 technique vhould ccmtiilue to be t h r sti~dvof rhe diurrtiiii~-at~o~~ tvl~ich1nakes thc i c u w t a~sirmptro115 LII or~iering:I nf rl~njorg o ~ i p sin rclarionshrp to gcolo+cal tinw So. hnuing can~tructednn JpprOFnate ~ - 1 ~ d c r ~ 2 1 n qec of nbsen~nt~ons How can we d i ~ n n g i ~ ~ rvrnple~iornorphil: 11 thc ncst ~ t a g c111 rsplonng rc!~tionrhlpsrr to raml ~ r hblc).;tratlgraplllc r i d r ~ .Fo: (shared prinntrve) from ~vi~aponlorpliit(shard hine this i n h ~ m a t i o l w thic, Smith ( 1 944) d i s r ~ ~ ~rsnec\t h n d o l o p 111 J c t ~ l l dcnved) cl7ar;lctrr s t a t c ~ ?T h e most ~ l s c f i ~w3y l Ir 'out,gmup cornparnun'. Here ;In '~ngnup'(ofwhich The result is n phyloge~~etic trrs, wlvclr ' i l ~ o ~ tllr~ ~ s the r c l ~ t t n n ~ l ~ are i p r ur~dcri n ~ c c t i ~ d h o nI <)
Divisions of invertebrate palceonblw
b; 111~lcx,!nil fcrr~nlcrrd the onc spccics m ~ br v 50 ancierl t ~r~rnnisrnsin their cnvironrnent~lcontext. t l ~ r t ~ n ~111 ~ l aappcnr.mcc r t h ~ tlicl: t havc wmctimer All animals are adaptrd t o their environment in all 111'1 11 dcrcr~\~cd :IF d i f F ~ & r\ ~ ~ Cl C ~ItC F ..liarl the true si ru- of-it%physical. chcmjcal and biolocjcal asp-cts. Each . I ~ I I + I ! I I ~ . I \ IT h,~rrI tn d ~ r ~ n t a n ~(a? l c with rpecies I S prrclsclv adapted to a particular ecologic~l .11ttlllcvlltC"~: Clllptt'r S). n d r e UI w l ~ i c it l ~feed.; and breeds. I t 1s the t ~ of ~ k Il'llci~ ~t C O H I C ~ t o t h ~i l l ~ t i ~ ~ c rni ~f iCiI C I T F I ~ palneoecology to fuid out about the rlarure r,f ~ I I P F C r,,!irrIl F ~ L ' ~I C F ,I I ~ W ~ V C T .tlicrr arc a number of qtaadaptanon$ 113 fossil orpnitms and ahnut thr rclnt~r::\.,~l I n t i t E ~ ~t T I A V11clr to p ~ t.I 'hiyhrr drgrec of tionqhips of the nrun~nlswith e ~ c hotlier ,ind tllc~r environments; ~t involver the e~ploraaonrlf both IT, tnr c.u.~tnplr, c:ln he iircd whcn .I wries o f present , ~ n dpast ecosystems (ScI~Sfer, lW2) T'IIIIII ~ ~ I Cr l ~r ~ t;llli~d F tngctlrcr. If n rnllectlon of Al;houph palneoecolo~is obvrously related to l r ~ L l l ~ q ~ r rIri < rntdt from lot.~lity A, the l e n g h / ecosystem ecology, it is not and cannot bc thc s.nnr. \\ dtlr r l t l l l + nr trmlr nrhcr appropriatr parameters rn modern commui~itvecology much ernphacis i~ 1ri.t Iir. plllrtcd on a p p h .I< a fcattcr dlngmta. A plnced on energy flow throusi~die svstcni, but t h ~ g IIIIL, nt'llev iit ( e . ~;I. redr~c-cdtmjor m i x ) nldy thcn k ~ n dof determ~natio~l 15just not possible when dealI\c tlr~~vt~ tl~rougZithe 5c:ittcr. Thi5 givcr a simple Ing 1 ~ 1 t hdead commu~lit~es. Instertd ~ a k ~ ~ n t o l o g i ~ t 5 !. = ;r + q~p11.u'hcrt. it 11, thc padicnt a i d I, thr perfarce murt concentrate oti ertnbhshmg thc cornwtcrtcpt on tlic y ax!?.A sttnilar qCdtfeT fronr n poppnsitioi~, qtrwcture and orgamzation af palaeocom11'11lonrollc~tcrlfronr locality R may hc plnrtcd on munities, in attrinptii~g h r n here to work out. r ph, anti t11c rrdncrd major nsns drawn linkage patterns In food w e b and in investigting 1. Thc rclatlvc s l o p c ~and c conrparcd ct.lh~tlcnlly.If Many attempts have her11 made to sunimarizc I h ~ n ccrtaln threshold rhe p o p u l ~ h o n s categories of fossil restdue. to providc the backI ~ c p r t l u d.I\ bring of tlir same SPPCICF; ~f ground for iiiterprenng anginal comn~unityscruu.n the, yxcicq different. ture. The scheme proporcd bv Pickerill and ,. , ~ , , ,nnt. v nf ,I ivholr r r n c ~o f pix~ihletern. Hrenchlev (1975) and amended by Luuklcy ( 1 qX3) i? I I I , ~ rnorc clnhoratc t t ' c l i n i q ~ r ~ o f niilltiv3n~te used lirre: :#),I!\$ 1 .jrc. ~ h~sanlrng~ncrcavngl\Important In LIXlliLt~'llIt. v ~ \ > 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<) 171: 11rlpq to rli~rrlt,lnglt.thc \Ianoor f ~ ~ t rc\pon~iblc o r ~ for d e p o ~ ~ t i n nf l i .I ~.r.lrttcul.lrfcj';\ll .irleat p e a t depths. TIwsc regini~c. ~ r r Ilow, 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\. A n l i n ~ krit~dplmit~that livc on rhc sea hour aic ~ncllidrtr~n<port.ltloii.pl7y\1c,iI,ct1cnl1c.11n r hiolt~yt l ci ~ drict t paw~vclyo r slvlm keblv rll ic,ll dot~tngcrn rhc shell, and tlic ,lr~.~chrncnt nf cpi- benthic; t l ~ o ~ water crduini~ arc planktic (= planktonic) fauna. Po~t-hunnl prr,rc\5c.\ m;ly l t ~ v t > l ~ ctlrc . dirturhn~lcchy t ~ ~ ~ r r n s r;~~rn. ndsrdlrrlcrlt c.ltcr\ (bio- qmuc tlic): are the plankton. Nekton (nektic o r turbation), currciit rcwr>rkilly. colur~or~ .~ntlor11c.r nektsnic 'CNIII;~) 011 the other hnnii coialpnsrc actlvc cwlinnlcrs. Nerltic anitnnl~belong to the ~li,rllou di.~pcnrticp r ~ ~ ~ n ~ n t~h.111gt'~. i011 watcrs nenr land a ~ ii~cludc ~ d demersal rleiaicntr ~ fie+ Modern environments and vertical distribufion of wlilch livc above the coi~tineiltnl~ h c l v cand on bcnthic arlir~lnlst l ~ r r r ~ Pelagic r~. or oceanic animals ivater? or i ~ ~ i d ddepths lc 131 Figwrr 1 .4 shnirvr [ h r matt] rnvlrmtllcrit+ w i ~ h i ntilt' 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~*
( )lilt' tl tIl
r11~.ocL-;l
O ~ L I C FFCC! ~
.I?C.
.1211c\
# 1 1 l ~ ,l1rr 5 0
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 11s0 ~ 1 nr \vhrrc
Horizor Thr. 171~1iil ticln of rec tlnrurc idIcncc. TI14 rtr.cnn\ if I t l ~ cijthcr 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 iT ~. ~ l > ~ 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
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 rnf ~ t;iur~,~\ tn $1 hi#- and low-etier~y e~l\~i-rorinlentq Arc oftcii vcrv disp:rmtr. Robust. thick-slirllcd and rouiiltcd .;pr.c~c~ arc normnllv .~ihptrd fclr h l d i - e ~ i ~ c~)rid~rior~'r, 'r~~ ' I l ) , l ~ r x L ~ ~ I \ L I I I I .C~ ~ tt l ~ rS U ~ ~ I I C ~71311CS I O ~ ~I V I I I ~ \viicrcas rhin-vhcllc~i nnrl f m ~ l cforrrlr poitit to a l\\*l (I\\-rlw r ~c J V Ca qtnhlr predator-prev rela tiotlsh~p. IT I %rrI F .I prono~itlceclbreak ~n lope. Cornn~llr~iry qmlcturc is norninllv wcll detii~rdin cold-tcmycraturt arrnq, l-rlit 111 wnrnirr scar wllrrr Horironbl distribution of marine animals divcniry ic higher it IF ~ e n e r , ~ lless l v clcar. T11r m;un rontrol5 atkcrinp the hnnsontal dtstnbuPctctrcil (1')lX). working o n the t i l ~ r l ao~ i thr rlpn nf rcr.rnt . ~ n dfnv11~ i ~ ~ rare ~ ~teniperaturc. alg thc K,-lrsCg:lt.tint ctudled and dcfiilcd mrrie of tlirkc rt.it11 m r r tit r l ~~ I I~J [ ~ s ~ s T~, 1 I3 ~n ~ l.~and j: w;ltcr t~rrbu- ur,~llyoccnirring cnmmunitlrs. Hc 2 1 ~ 0 rrcnipizccf 2:nru T1ic Inrgc-~c,ll~* diqtnbz~tin~l ~ l,111lin~l~ i 111 tllc two c ~ t c g o r i cnf ~ bottom-dwcllinp nillm.rls; infautrcans is largely n f~riicriniiof rcrnperature, whcrca5 nal (Isuncd and lnvinK warhln rhc sediment) . ~ n d : I C rvhcr ihtrnrq gr%ricr,~llv nprrarc nil J illore local epifaunal (Irving o n the sea floor o r nil I-orks or ilc T ~ C I F ICT~ .Y ~C ~Trcgnus ~ c a r r y the inost divcrsc w ~ w c c d ) I. t \\la% ~ o o i fi n ~ t n dthat parallel ct)llimuni;%tncrr, .~nctIn thew thc E ~ C C F P F.ire vrn. uil1slzrau.i tics ucc-\IT,with thc wmr senera bur nor tlie h a n l t . lllrt t l i r nurnlwr of ind~vlduaho i JIW c\nc spccics if i-r~munities, rhe hnbits of rhc animals currlposing rnrltinctl M w ~ t r nof near-normal salin~ry.A thcrn, communitv w o l u t ~ o nt h r o u ~ htililc nlld. .I$ .,re ~uryhaline,r c. vcry t o l e r ~ ~ of~ reduced t Tar 3q pw~ible.the controk acting lipon tElc1n (erg. I1rr1n2.Thc hr~ckivlirvarer ravirnnment ic phyrir,- Thnnon, 1'357, 1971). This is pcrh,qw rhc nlwt I~I
1
I(
Divisions of invertebrate palaeontology
c
pure 1.5 Disbibution of modern marine
shelf-living onirnals in founol
o
l
d currenfs
provinces, using Winkel's Tripe!' projixtion. (Bused on a
s w r g In Br~ggs,1974.)
r ~ a m p l r .tlir t ~ po f Florida c~rrir.:;a tropical shclf snl,>redto r l l t r ~ o l - t l i e , ~a~~t l d11orthwc9t hy wlcr ivat?r arm\, biit tI1ouq11 this is part of the &'orAtlannc troptcal shrPtprovincc it ha? hcen iwtrll for t, 1";' Mciierrow and Scotese. 1 qclO). Whc-n uced in ).i!uncfion ~ ~ i tpahcocun li tiilenc;tl rn,tp.: rt mav hc )~,ibli ta s r c IIOIV the dictributirlll pattcrrl n f ~ c ~ ch~tnas nt relared to thc pnqitior~o f a n c ~ c .con~~t ~ t n t a lma<nleti~ilcspalacnI O Q ~ ~ ~ ~ L ~ard , I ~ grnlog~c,~l ~ I C ~ I ~ evidence may havc b;.annc upon palaentmperntllres 3 r d cvcri allow me snt'crciice to be ~ n d d ercprrlltlg anclrnt ore;m llr
11111.
~trrtltsyctenlr.
Modem and ancienf reek Thrcwtghou t gcalogical n m e anirnalq have 1101 only bccornc adaptcd to particular ctwironnients but alqo thcrn~clvescreated ncw hatmats and cnvirclnrnentr. W i t h ~ nthese there has heen $cope for a h n o ~ unlimt itcd ccolclgicnl differentlatiot~. Pc-rhap the most striht~f:examples of such biogenic cnvironrnenrs are tllc reefs of t11c p a ~ tand prc~cnt.Rce& ( h g . 1.6) arc marcive accurnulatlons o f llrncrtonc built up by bmc-~ecl-rringa l g c ar~dby variou~kinds of ~nvcrtcbrates. Through the activihes of these harnc-builders grCnt mound$ may bc built up to sea lcvel, wit11 cave< artd channclq witlwl them psowdmg a residellcc for RA5IN
SLOPE
LbGOON OR
AIM
reel
SHFLF
reef fac~es
/
Figure 1.6 Generalized =tion
through a reef {olgol or coral).
17
~ i i 1 1 tllickct~ r l r l t l l ~ rl-cf w r t t ~ r ii1nn111cr~iL31~ k11I & c d : I I I ~ II,II<. I .LII ~ L L ~ I ~ ~ L ~11fi:r- J I ~ Y in!: in ~-1~il11pc \Yihc.n thccc rnrlic\. ~ c \vc.rc tlcl 111 ttw h r r ~ r ~r 1 5 p ~ ~ 1 t ~ -rrrL li of tlic t r o ~ ~ 1 WIS ~ ~ 1 1 rurtnct in late Midrllc C.nnhri.1~1t l ~ thcrc 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~ 1vstrr5 it nllo\v r.rplrl cnlriun~ rrc.f\ \vcrc stlninatolrt~c.In Middlc t lrd~)t,ici:rtr1tt11c tlrc~c ~ l g , ~ivcr~. c joinetl c,\~l-,ll< .ai~tlrrrotn.lrll~ 1 1 ~ t ~ l ~ ~ 1,il!tj ~ ~ 1!1~71cr ~ 1 1 1 ~-~3~ltl1lll~7ll~ s~(>t\'tll,-rll' ~ Clli.~ptrr4) . i \ \I rli pritlc.ip.11 I'mliic-builclcr< ,Ire .11,~.:.1c. ,111d c-orX>l\.l y u r pnroid\ ( I I T I I L > - ~ ~ L~ L T I I I\pi>ngv<; . trrgctll~.rhlrii~cdJ rCc.ic.nvlthcrc. .lrc 111:iny nr.1cr kincls t i ~nv~rtc.lir.itcr in thc .w by rcd , I ~ ~ , I L - tvhicll . ~ t t l ; ~ c t l.I r ~I I~O C ~ 01- otllrr I ~ I V C ~ ~ C I M - A ~ C ~ reef ct,nHrlLltrjb: \pony.r. b17,070,111< ,111il t t ~ ~ l l ~ ~ rotlrncnt cr'~ i r ~ ~ l u r l i nblacliiol,rlcts g ,111cl trilllhitcr. Fnr M I ~l;nc)x~.t nr~longiid l ~ t cI l c \ , ~ > n l ~tirnc. cuntirru~ll~' beep? p,>cC1 ~ 1 t hr~iZ~~rdriicc, bur ~t 15 nlso tilc ~:arhnnifcmuf cvcrc m.i!~ll!' .tl~;11. rnLeii i r n m thc rcrf $mint. ~ n l : ~ tw.rc dccpcr bnqms in ~ { r h i c l l t h c 1~1tc1. ~-crir>rf~r:~Il\ Uchlt~dit is a lagoa.711\ ~ l t :Il ~~ - n r .~rrttd ~ l ~ c d ~ m c nz~d nr ~ C ~ I \IC ~.lr,C 1il.r~tidal fl.~t<~ l o n qthr \IIOIL= r-olr~~~irrdl l n rv:uinl~acrcri.l hccnnlc mnrr xaE~iiv IT c\r.~pn~-.it~C1; t Ilchind t h ~rccK , 1' ('blur-qrrns'). C;rcrrl ,ily,~c.lrc c.oinlr,oncclr 111 tl~ra n-~cvcv.lpor,lnr>n in p n i i t ~ a l p.lnr b x k - r c c t ' t i ~rr\. r d ,11q.1t'.Ire thc I?n.itn Ilmc ~ccscrt-r~ wntt-r clr.~wii thrn11!:11 t t w rccf rlric,tl o u t In tll, Ingoon$ hclllnd. 111 thy Pcrmi.111 rcrh uf TL-Y,I\ .IIICI of tllr rrrt' rrcrlf Larqr rt-ris r t ~ ~ .,I\l i t l ~ t ,* I I ~ Y ~nrc ? kno\\n nq bio- nurtlicrn Eiigl,~iiri,w'licll arc vet?; I.irgc, t l ~ crrc. (runt s o q ~.I\ .I V ~ ~ C I C ~ 1li.111 , I I <~.fl.~tt~iii.ltcc~ n l q l \llcct\ hcnns; thry krill dtsc-rrtc ~ n a u t ~ dr q r < i t q tironi the g ar rhc rcr-f rrr-rt ~ v l 1 t . ri ~f ~ C . . I C ~ Ic(~ IL ~ sr,l floor. Biostronres. oil tht. utllrr I ~ n t ~ d ,Ire . t h t t ~ ~ r n i ncrrcr I . ~ ~ I I I I C'UTIIIIILFIII~ICI I.I~ of rt'~,f-h-pt',11111t1.11~ 2nd h,~rt-ly lcvcl (Fig. F .i). T h c uppcr \ust:~c-c \ \ . I \ fiit~iirc.ly colorii7ctl nw nl-iw~.thc ica tionr. -, < r r o ~ r ~ n t c ~ l\ivt cl ~ ~. crllrl r r l cwt tow.irds thc I . ~ y o r ~ ~ a l 1 J ~ r o z ~ q l ~ nqcnEoyi~ ut ;1l h r \ r t m t l ~ t a ~ - t .h:lvc hcrll rct.c.lfiui gci~cr,llIrlorI ' J ~ ~ C ~ L I F~ I ~ I L nf ~ F rrct' ~ ~ r ~ i ~ l l i iwhi~-li i t t . I I ~ V C ii.lck-r~-t.ft:ac~c\.Tlii\ kintl { l\ tl11 .1TlCf?ll, H c ~ L ~ I - I s ~ I .I!IC~ ~ = L ~~ I C C ~ ~ rIk YI tIi ?~T C t 111 .111 ~ i t l l pl~olngy ~ ~ thr-rcic)rc cine\ n o t ~ l o ~ : c,apyroslrrl.ltc. hllcll the frarnc-bu~ldr~rili,ivc l ~ ~ c l ~ i t i cnlyac. tl hut thc stnnd.1ri1p.lttc.n, 13iFip. 1 .h. Whcn tlir Pcrrr~i.~ri ~~~\.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~rpresent 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 ~oi fnthr p cclml-,~lg.~l rrx., 3 srncs o t r r e i - l ~ u ~ l ~ lalil~n.ll< ~ n r : (Nc1vr.11. 1 C171) Thr t>lclt.rt k11~7wt1~ C C ~~ Cr cO V ~ I -30(111 Ma 01~1. qystcrn rlrnt nrrbsc in thrm lntc T n ~ n l c .Cor.11 r c r m t i~ladeup tntlrrly nC;71ir1 I REEF-CORE e frtrrri .I dccp-tvnlcr rurhid~te-filled I ~ . I F ~ I I , r l ~ Arc ~ r ~ n o prcqcnwl ~ J I L - [ ~ I ~ ~111 > J tI l~i\e ~ crt't'if. IIL~WCVCT d ~ i d tlrclr ccolog~cnlsrnictilrr nia\. 11.1vc.hccn VLTV rlniplr. Wlth tllr ricc t7r'fr~mc-bu1ldir1ymct.17oat)r u~ thc Luwcr C,llll't~ri.~r~ 3 iiciv k ~ t l d13f rrcf ~-ornmuliin. madc i r i . ~ p p c . ~ ~ l ~ Strotrlatnlitic icc rerh were Figure I . 7 Crest of u Permian olgo reef ( R d r w n from Sm~th invarlcd 1.v tllc ~pr>iig~-ltlic. nsch.~cocv.irIi~d~, qrnv1981.)
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 that tllc ~1+trc\r111~p c r ~ o ~111 d r11c c,]rlv C ~ P ~ ~ I L - L -cu;lL>i-ti~,lt~tl~) OIIG of ~ I I C p.11-[Sof .Jn!- ~ ~ ~ J I ~ IE$ Stllc I ~ I 1111 I~c'rt.,lrc- nn r c d ~ktlo{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 ~\zc.rc j 101nct1 i t 1 p o \ ~ t l ~ Thc . ct*r~tral~hqut,licrc (1l~oinn.;. i r d Invcrgcnr 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 t r ~ ~ l u l i c a t l ~ lII,ITTL~\VF y LI(w~,iit l ~ cCpcctnitll of p0wi-11 pncubilitie\ li; . t:.!1 Ft tlic r ~ l r - l ~ t ~ cc ~ l ,n~cc.l , ~ r ~Tllc c ~ t ~liltl~rc. . ot' ctrll further. Sonic rit'rhcsc y p c ~of ~kelrtonsn r r J\ ,% l~rll! r n i i i ~ n u ~ ~ ~ ttlir i r . \ .mort complck (IT ,111 & l l ! O \ ~ $ , I I I 11i1ir c t mylInr u Lv qcrii. Eutcrnnl flicll\ STn\Ylng a t thr edgc5 oidv bv Functional morphology, growth and form ~ c ~ r c t ~of o inin z nlntcrinl nlni~g:I p.lrtlr~ilarnrar111: i~~nc.trr~nk rif p a r r t t - ~ ~ orga11.i l . ~ ~ - 111 1;3.;<11< c;llinvt 51na\ 2 0 1 1 ~o f s ~ ~ ~ wVt chr.) r s f i c ~F~I I T ~ g~mvtll I?.v~r.11~1i~hi~i hy III,IIIV .?c ?T~LTIIOL& A V J I ~ . JL L~~ ~ ~ rcc~1ltqill n lcly:~richiii~c rp~rnl41tlll fit 111 br,lchio-lil~l~~!y~t+. l?llt 1L I < 5tlU ~ l O ~ ~ l l . 7t'3 l r 20 3th,rt btwr thcln \vnt .~ln.v. Such Eutcrn.ll nmnllv ii~ wcrrted alcltlq J i l u c F ~ I I I ~ I ,~ I \ ~ I I , I Ii l i t ~ - r p r ~ t ,15. ~ tIIC>IVCVCI, ~ ~ n ~ Il~ilitrtl,111d III ztnilc wllirh 111,iy13~. hut I < not .lln.,lv\ i i ~ n r ~ r ~A~ a l . I I I U I \ nn\,<~t i i h.lrrl tr> 1>1-\'01icf.I ~ ~ r r , p0111t. ~in ( r rLn tvllci~ tlic i ~ ~ n ~ - noi u n,i p n ~ r i c ~ r l .o~rrp n 1s 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 I 1 I 1) I ' d ~~onto!ngirt~ .IrC S ~ T ~ prth~cntrd lockcrl into plncr, though thcv in,l\r th~*rcaftcr I I c t t h t ' ~\tfitIl &T~I\L in~livirluallyh ? ~ .~cr-rcrionof n ~ a t c n a l corlI ~ ~ : I E I \ w,hnntitcrp,lrt~ In Irr rnq . ~ r ) i r t ~ a l s01-. w ~ t hh r ~ i l c cciihjc zoncr. t717,lrrc ~ppc.1r~lw-c xx~lirrh.11-cyo ulud1fic.d horn Estcn~nlskclctnnq ,111 f~~n11r.da t the m e tinrc.. 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.II of .~rhdcpicoctic .und I < ol~l!r pocsihlr rlunll? moultifi~p. Tllc dr~,irlv;untngcoFtlilr ryctertl 1% tlit v t ~ l t ~ c r ~ b ~ l I;*l[!natk 4 lLJOl), 1wc.n ~ n ~ i cdiccur\crl, h but i t i l r ; Itryrly O I I ~ E I ~ tlw C SI.C)PCot' 1111s \t.r~rk ,inJ ir\. ro c t ~ m , ~ .g~cr r dprcdatinn I . ~ L I ~ I I ~< r ~ o u l t i n g . ,lnl\ cnmr cx.1111p1c': ~i ~ t \ . I F P ~ I ~ ' . I ~ I L I.trc II , I ~iin~!iitd. ( E\g~b~ltt;tll~, ,III\I kind o f ckeleton. ~ntrrnal o r T~rv)rrl~tod,Irpcrt\ nf p . ~ l ; l r o r t > ~ 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
~:!IL-
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<,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 T datiy~~ ~.~.t~r~ that cvcn in J small arcn rock utdts ctrt acrns.; rrrnl ~ Y I I I V , I ir.1 pl.ii~e~ For . instance, if a rhorclinc has becn advnni\ 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. al will be lcft in 11 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 . ~ 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.irl~7i~ layer of ~ c d i m c n r \, ~ l ~ i cqnrzlrtir11t.s li Plas been much ~~rlpenistcnt: diffcrcnt ~ t d ~ r r ~ t ' r r thcier n y nlav hn\;~? ~ ~ \ . t l ~ q l I.,< the sanlc for I n t ~ ppcnodq of ~ F I I FGut ~ a t other tltncr t i l a , ti>r 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,<11' real valuc within a rclativclv srrlalE rcglon. zons, and the ~ I F I O T \ ' (3i 3 rock rcclliciice rs rcflecred pt*l3~3d\WI The rlivi5inn~ ercctcd in litho~trat~graph an in its Inyenng. E ~ r l lavcr i 111 tilc rock sequence i1uirt armn!ged 111 3 hicrnrchial cyturn: graup, formai n ~ ~ l l ~! n havc been Itlid dotvn a n a psc-csl~tit~g l.~vcr,so t h ~ t tion, member nnrJ bed. A bed ls a distinct I~ycsrt T l ~ r * pr thc a l d r ~ trockc arc ;kt thc b o t t c ~ ~oft ~at1 rxpcr~ed a rock qequcncc. A rt+crlibcr is a p o u r elf hcd 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 ucliaractcr~. n A fonnntirri I t y i r ,II ~ U C sion I ~ n sI-recii ~ ~ c t n n ~ cinvcrr-rcd. a l l ~ Thlq IS the prin- is a group of nicmbcrs. a p i r l uuited hy ch,~rnctrr (11 I I ~ P ~ . TI1 ciple of superposition,rccogtlirrd a5 long aso sac w ~ t hkaturcr in romnlon. It 1s thc p n i m r y u t ~ i 41't thh') bv t l ~ r l - l . ~ ~ ~ iccicntifi logcal C V C I I ~ Sb u t J ~ C Oiip~11tllr hi~toryo f rallks abovc forr~lntion;it 1s corl~poqcdnf hvc-~c-r' 11fe 2nd is p c r h a p thc rrlrlrt bn.;~upart of all geolop more for~natioiisand 1% oft~'11 U S C ~for s ~ r i ~ p l ~ + i i i ~ (Harlni~dand Annqrroi~g,I I; Uc~tton,1W 5 ) . srr;lti*qcraphvor1 a ~rn~ll-talc rrrap. Mucll nf ~ t r n t ~ ~ a pi q hmr~ccrrlcd y ~ v l r hclxonol09: thr. ge:t'olopc,llrccord h.1~to I'c CIIVIC~CCF ttp into Biostrotigraphy tulle penodf w h ~ c harc qt.~nd~rdizrd, A S h r nq ~ O F ~ I -in b i ~ s t m t l p p hthc ~ fo~silcontent5 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~~rnpllvhas 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 aclaisil 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 fC ~ ~ s~ iClS C F I I ~ ~ S : (-'
F ~ ?
Divisions of invertebrate palaeontology
OF
the
tllc bnlc that thc 4. Since fowl speclrs could n l i ~ a t ct'ollow~iigtheir dcpo~ited.Thus d u n n g own cnvironmeilt through tmlc, thcrc Ir alw.1~5a tht r',~rlyl'~I,~ct)znic rsilnhitc~and brachiopodq were possibhty of diachronouc f i ~ i ~ ~ The a c . zolir as rhr niurr rrvnnion ~OFSIIF. Ry thc M ~ S O Z Ct11c I I Cnlost 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~~jvnlcrlt to that In al~crtllerregioll. ~mrnonitc\:tlicv too bccnn~ccstincr, 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~crimc r,rniy< ilf i n d i \ ~ ~ d af, k ~ ~l i qpccic~ l art. biozone of a species 1s not likcly to be prc5cwc.d In uamliicd ~t ili rvidrnt that wrnc nf t h r ~ cInstcd 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~ousboundaries or time line<. #.q.prcriqrly a p:~rticul:~rhricf hicrnrical penod. Ideally, zone fossils s11~>~tld IIRVP;I p ~ r t i c ~ ~COIIIl;tr In In\ lr~calxcn. nncc tlic xcqueiice of-fbsr~lfau- 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 vcscahlishcd thrclu~I1awduous for bio\tntigrnphy. T h e w would be: ollrction ,ind d o c ~ i r n c ntlon t ~ horn cspcrccd qrctlti\, th~rLr~nwn~ ~ r c c r ~ rail ~ i ohcn uccd for cnrtclaa wide h o r i z o n t ~ ldi.itrilv~rinn.prefer,~blyin~ri-oli with clrhcr 3 ~ ~ Ccrtain ~ 5 . fowi1 qpcrics havc cnnrrnmtal; b?ctl rn~ind tn bc p.lrtir~~l,~rlv good q r r a t ~ k ~ ~ p h ~ c a ln ~ 1 1 ~ verfic~l t-t r d r q e so that thry could be uscd to 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 ~~ nq zones. To rake -111 Ifnlll; t,m~plc,~mrnnnitcq,Ire pjrmrulnrlvs{,rlorlzonc foqc n o u g l ~iiinrph~tuqical~ h a r ~ ~ tto e rcnablc s rhcni 1% Mcqrvnic crroticctphv. Thr J I ~ W W pcnod IC to hc rdent~fiedniid d i s t i t ~ g ~ l r l ~easllv: cd 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 r t e hy which 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~ Y C S ~ Fc,ic 11. 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~sarc scldou~f~~Ihllcd i l l tornri rvcr. vrn: cninplcx, .uld FornL* part$ o f tlic gcoFils used for 7nnat~nn:perhapr thc t~crlrlcnnlrrloi~ircq )8fm31 ruct+cvCion,Irc t ~ ~ RlOrC c h c'li7\~1y~ C I I I C tllnn L~ come cltlse~tto ~ t and , ~t 1s not ruq3rlslnF r h ~ t111~ t11~'~. Tlir i ~ ~ ~p ir ni ih ~l c ~ arc t ~ ~J F fnllo\vf. p n n c i p l e ~nf reallv precise stmrigraphlcnl c n r r c l ~ t i o i ~ werc tint worked o u t hilly with ~ S F C F C f o ~ ~ i l ~ , I Minv k~ndsnf f i ~ r r ~c~pccially l~. rFtc>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. titsidc rhls cltvironrnrnt. arc four l r n a i ~ ~~ I I I ~of F ~ I O Z O ~ Igenerallv ~ C uxed Thn rncnnc that thcv can o i ~ l yht* uccJ for come- (Hcdhcrp, 107h). Assemblage zones arcL beds or. Innng p ~ r r i c u l ~cnMronmrnts r and ~ I I U nrc ~ not proups of beds with a nnmr,~l~~5urnblagc 1 d fovcilr. un~vrrmllvapplicahlt T h e y Inay bc based on dl thc lo9qils prc~civcd 5nmc k~ndrnt'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 c vcn. FIOIV. 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 ~ dnrc of v c y arc. perl~npq nf nlorc prlcral applicatiutl. A rnnpc littlc IIW for ~ v ~ ~ h l i s l iclmr l n g r~lbdlvi
p.11~rcflcctlnn
21
tacloting \cdirncnt~ wrrc
itjr
Ilft n t
:.tr>~~lphvA stage, utl thc t ~ r l l c rhniid, I* :I p-oup I I I I F ~ ~ , I ~ ~ < ~ J thc ~ I I ~ .n;lrrowest t h . ~ tc;ln ~uall!' I>rzbsccl cjn ,I rcLjotl.il sr,tlr. A . . 1 I i I M C< ; ~ . I M . IIy nr t11r ~ t . ~ gIt'v~*l r r h ~ TO&< t 1 3 cu ' I C ~ C I ~ Ilrll. Ncw Yr~rk.(U\rLll bntlc tcs~;t) r i c s c;>n bc c-orrc-lntcrl. A i :In t.sL~rriplrs Ll~rringc>n, F .J.W. t 1 CEt,71 hr~t.rrrhr,~rrStnrrrrrrr ntrd ~. z r j c > l ~ )trcut) p x n c dijtiiulticu in t ~ ~ n l i i nprcii5r y zo~r;~l Ivriri!lcvr. NC1~lyll prccicc 21111~~-r0-201lr C C ) S T C ~ , I ~ ~ O I I S EI~cvicr,AmrdChjopt>ddiutrillur~nn) ,r,I+?~-tci:l~i ~ a p ~ ot cl i znijCs ~ - i o m ttcl ~ I v s t ~ v i ~ l wilt11 ~ IErjuc-ut, A.J. (1 9% I ) I'ritrriplrs q{ Rr,rrfhir- ,I h~rirrr :!:rm< dciinrtf all ihcllv t&il';. P n l ~ r . o r ~ r ~Acntlcll~ic b ~ ~ ~ . Prccs. N r w Ynrk. (l'n111.1 blc if Io~91lrp v r a rcl,~rivt-~ l i r t ~ n l h o\ vp l ~ i c lr.tn ~ bc idlr-lu?wcr.it~i-) ,::J , I ? tlic prl111'1q- i~,wi\ ( % < ~-l1ri3tjt1strat1g~~pl1y, L I ~ - ~ I \ C 1'.I I ~. I~I ~I ~~I l.irprr+ f ) , A . T . ( 1 '1'5') k ~ ~ - r ~ . i p t < ~ , , ~ ~ , ~~vt~nliclcrc, it is r r t i t * ~h:~td ~ to inrrclntc p r c c ~ r ~ l y f!t-ol(y]> ,.it/(/ /~~'i>I~~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 ~ tblrr I l i ~ r dt ~ 1C - ~ r r i - - I I r ~ g p J,C, N c w Yirtk. (I>cllnlitntion 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, I<, 1 L)LjJ. Hi$loly '!I' L+, 7r1d call. l H : t ~ k ~ ~ v I l , ,.*!,,1 I I I- ,I~I L.O T T ~ 'ti011. ~ ; ~ Ar nloir m , ~ r ~ ~ ~ . t p hori~ans l~icaI ( )xfi>nl. ( E l r l i r ~ ~ ~rc.nd.thlc) ~tlj: 1'1t.r;.,Ire ucu.~llv.;clrnc uhl q i ~ t t c m\rorld\vidc ~ trwils. I)odrl. J.R. and Smntnrl. 1 t J . (13c)ll) l'tiEnrar.rt~l?q),, 5 . 1 r i 1 . 1 ~l n t e r c r ~ ~ r r i n c ~ rr)rrt.l.~tii,n ~t:~l tr 11ot ilnpcrc.iiblc. Gvrrt'pts ,rrrd .-.ljy~licrtrii)~,-, 2ntI cdrr. W ~ l c y .N c ~ vYork. Ill ~ . l l r n ~ - r o ~ m . l t itgl ir ~ ~rc\.~tivc ,~ ~ ~ \ i ~ r r ' ~ 1 i l c n ~gcv c n .5( 12 ~ 1 1 .(111trrc..;ti~~,c approacl~) !,: :hc t;:t~r~I, is cuppI~.n1~11"'1 a n d i - n l ~ ~ ~ l c hy c c i E~TII.III. S. (1053) Z ~ ( ! ~ V I . ,,!/' T tht~ ~ , )5 iJ~ 1Sidpvlck , .111ri ~ll\rllutcdatci \\,hicli c.111 hc .rfi;ucd ; ~ rertair~ t points J.rcliwl~. Cr~ndon.(All o l r l c ~but still uzcftll text 0 1 1 tl-iannr lifc 7 0 r i e ~.111dfa~l11.11~ ~ L I V ~ I I C C F ) ,! '):*rc.vcr approprialc r o c k rxcnr. 'Tlic~c.nrc usunlly F ~ I c I ~ c L ~ ~N. c , L a t ~C ~ ir ~ c r n t t1 . . ( I W I j) PIiyltyt,r~,,ti~ i'~fttr~tq '.~V.I\ b(=~i-krtcd hrt\wc.n ft7crilikmur seditncnts, :tiid NNJ rllr Evr~l~rrioritiryPro[-css.C u l u n ~ b i Unuvcrc~ty ;~ I'resc. . ' I : ' IibTCLCrCCtlcc ~ I\ 1 1 1 ~; k ~ 1 COIIIIII~III. tt i\ m o s t Nc\v Ynrk. (Exp1,11ns rl;~iliat~i. n~ethnils) #:.~lil;rl\..thcrcti>rt., r l r . ~ s r,ldinnwtrii cl,itin: lvill Fnlrhndpr.. R.W. ,1nd J~hlnrlski, I). (ctls) (11)7')1 '1111. .!~!+l*ncilc p ~ l . u r o n r n l o ~ ~ccvrcl.ltion; ~-,~l t31t. two .arc I ~ ~ ~ c ~ * r / q~j~Efirt/~ p t * ~.Sr;ct~r-tx, l i ~ i Veil, V l l + TJtt*L't~yc!ryi,dr(r 1il'r:I~ ( - t ~ l t ~ ~ 3 l e l 1 .~lFTLi ~ ~ ~ tI1c t , ~ lp-vGlr ~ ~ ~ sLlc.c-r-5$ 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 ~ ~I<. ~ , (1Wl) F~uit. . :1
a
8
K.C. Tr~rherr,U., Rohlqnn 1i.A .III? KJCIIC~. 7 'rratrrt. 011 It~t~~ft~I~ t rat~ i Ccolog~c~l Socicty r ~ i Atnrlrican ~ r l dthe LJti~vrn?;ltv Kans.v I'rc<<. Lswrcncc K,I~I.(771~ c t ~ n d ~ rrferrncr rd work u11 irivcrtrhratc tbwF I I ~- ench phylunl truatcd a l l r c p m t c ~ . o l ~ ~ n l e s ) . MLKKIX , J.W. (id.) (1 (IXi5) Arlhi L I \irr~t~rtc/*rm,~fr ~ d\l,~mfi*q\~/( Lonsv~,rn.f l a r l r ~ ~ for~ ~rlie . IJalnrontolr>pcnl As';rjr1.\rir7n (Exccllcnt phocopapllic coverage of ~ n ~ puuprj in bun.i\) 1-1.1lk11~i.4, ( I cJt){~\ .-It1 t1111i- Plt,~tic-r,~zt~ir l?i~~i-tlrrc~p/qj.l'.n~l, C.!<.C. ( I [MO) T/w >\Tnrt~rd Hi\-hrd. rrting) Il.~rlnnd, W H (cd.) (I"7h) '17rr. L l r $ 1 1 Rrt-t~rd - o I ' i v c t c ~ ~ i1. , (cd.) (1 % - I W I ~ ) 'rr~~rti dto j)ti!C[~t!f~llo?rr~ M,rr.~tln, 13.1nr. (The Frerich 'Trcanw', : Pr~lcotrrt~ll?yy, 2nd rdn. Frecinn~i, S.ln Frnnclsrn I'l~trc Ffltt>. C a r r i l m d ~ rPJruvcnin; P r w . C~nlhriiidjic. (Exrcllcnt tcatl~ookcmpha>inng apprudchcc 3 r d rnn353 pp. (l:wenn;ll nnd up-m-dm) icpn. 1ir1t n o t i n o r p h n l n ~ c a lor W I ,Ytr,;fi?rc~pfrii, ~ T I ~ ~ ? I ~ Grti~lr. ~I/ ,~rr,f (:li~.<.
R.C. tci~ccc.~$ivrcditorz frorrl 1453)
Londnn and P . ~ l a e n n r o l u g x ~Assoc~ation. ~l Schsfer, W . (14)77) E ' r o h ~ y rritii I~alorclr.colcyy4 .?.lrln)l[ Irt~r~rrt~rtrncrtts (trant;latcd c d i ) (cd. G.Y. C:m!g). Cll~vrr 3 r d Uoyd, Edinbursli (St,indard work on Kercr~t North Sc;l cnvirormlct~t';with applicatio~nfor p.t!aeoecnloyv)
S~=hoyf'.T.1.M (cd.) ( 1972) Rhlrit.ls it1 Pr~l~ol7iolrrq Frccrrlat~. Coopcr, San Fcqilci~cn.(10 ont$nal pdpen. l w r ~ n nO i L I U ~ ~ ; ~I IC~ :. I I I \ ~~11.1nrd0 1 1 gloh.11 n~ap\) t IIIS\IY\. N F ( l t l S t ~ )l, n t t ~ r,\rw ~ ~ r ~Rtlct~rd1n2 nlanv q g ~ ~ f i c a n z ) Scott, K.14 and West, 1t.K ( e d ~ )(197h) Stmrt~irrarrd nwrl Stmml Corrr.b~il~~n L-rlrr~rqtrt.r.Carr~hnclpeUniversity C i n j ~ i f i r ~ ~ tof u r rP ~ ~ ~ ~ o c o t n t ? r ~ tDowdcn. nitrn. hutch ins or^ TErrw,Ccrmbnrlqc, J-tpp. and Row, Stroudshcrg, 13cnn. (1 1 nnginal papers) lowcv, K.A. 2nd F r ~ d ~ vA.E. , (IqH?) I'~clhl[,rrtr of I)(t)d~4pcmr~rirt Rt~tt~~r~~rrrrtwrr. Cyqtrni~rrc\ Atrorratinn Speci.11 Skinner. 1E.J. (cd.) ( I O N 1 ) P~fcntrtolngy irnd p~lt.wrrr~~r~~r~rirrn~rt.*: Rmdi~l~gs.fionr .4rw~siron S C I P IrrI ~ Kalr J fnlann. 1-05 Vt~lurt~cNn. I I , A ~ n d ~ n ~ Prra\. i i Nt-w York. ( 1 1 Altos. Calif. (21 cclllcctcd papers) Sttiith. A.H. (lL)94) Syrtrrrratirs arrd r l r ~ -fossil rt~orti D a r ~ ~ ~ t t c . r ~rt,t>i~~tralrr~r). 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<,l\v+nn, I'.F .111d NCU,111. G. (eds) (FC)71) i y i t r ~ ~ r rPr,l~vtr,rr ~l rri .+II.I. t~rrri'lirtll,. Xcc.1 f-.lol!';r 'Thomprot~, d'Arc7: W. (1917) Un Grorrlth nrlri Ftrnll P r m , L I r~q > ~ ~ ( Ii3I .n ~ r ~ g i ~Iprrq i . ~ l C ~ LI I I I ~ I d~ I~ < t n h i ~ - C:nnikrid_ee U n t v e n ~ t y Prec~, Carnbndgr (Ind~vldu a l i ~ t ~ clawir c work i>n phwlcrl lnwr determtniny
K,~nfhanilF . . G . .)nil Scrltt. K W (147(1) t l . 1 ~ 1ctnllicprr ~ qnr\-.'tl?:n t l nbrtdqcd r d r t ~ o nn f this honk nr.isc.rlitcrl I.? y , Sr~rrir~rn, 1.7, l k > 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 ~ Jram,I.OIIL~OII. (Si 11 I ~ I L * . IVC-I1 i l l ~ ~ ~ r r ,text: i t ~ ~~iri~ls ~i Locklcy, M. (I"X.3) A rc\-ictt. of bmclilopnd doiilzn.~rcd y ~ l . ~ c o r r r r ~ i n ~ u n t it ri nr ~ i > t l ~ t - tvpr C)rrlnulcinn. P,~fi~ci>t~ttdoqy 2h. 1 1 1-47. ( T C ~ I ~ U I I O \ Ltix T # p>het>coluililt~nir\)struuturc.) M n t ~ r cJ. . nnd Willtncr. 1'. :1'1'17) C:~mvcrgcnr c\.i>lutint~ j11 i11\~crtel3r~t~<, I ~ I ~ * I K~ (!- I~~~I ~I72. *~I ,~1~#-O( \~ h Ncwcll. N.13. ( 1 1j71)Tlic cvolurlon uf rccfr. Stit*rrttfit .-ltrir~rr~~~rr 226, 5447.5. { l n f i ~ n n a t i vrllort r p3pCr) P ~ l i ~ i r rA.. R . (llJOS) 1310tnrrc. - 3 IWYV k113ciof *trLltlyt-*~pI~i~~ ~ ~-{011nid i i t t l f P~:/~~c[>t~i~>ft~yy 39, I 4'1-53 Individual p a p e r s and other references P ~ l ~ l i c)\.It. r, ( I 984) TIlc hjclnlerc pmlllctl~.c\~c)lutionof I'r.llg. G.Y. {1"?-1) T l ~ p r a l . i ~ = c ~ ~ c n lt)1' r ) ~t \?r ~ *Top Hucic .In idcn. J>r~rrl~il of ~ ' d r l r t > ~ ~ t39, o h ~ 1y-IC]-3. 511:d~b{Lo\vcr Carbonlicrnr~<) a t J. ln~-alln-rlcLlr Kilsytlr. I'ctcncn, C.C,I. ( l V 1 H ) T h e rea bottom and it, prcjduc( ) r i l ~ ~ r ~ r ! y j ~i?f~ thr ~ r t C~;r~liyi[td ,~/ ,at~i\h R1oloyrrll 111.7-1 0 . (Clnrr~cpnl;r.-nc.cnltl.~cal .irit(ly) Statton 25, 1-02 ( T h e tint drtinlrivr 5urnrnni-y niljrnFortcy, R.A. a n d Jcifctrcc. \<. (1"'X 1 ) Iihvlog~-:iy . I I ~J t l ~ r cc o n i r n u r ~ ~ h r ~ ) w < ~ c ! n ~ r i-r s .I roinpr!nt?nc ~ ~~ ~ S+ ~ I ~ l~l t lD.U. l ,~ ~(1OH~ I) ~Ttie Magncsi.iil L~lnr'itonc (Upycbr brc,llvc c r 7 ~ i l r ~ l ~ l n ~ t i ~ * ~ i Pcnm.~n) rccf cun~plcxcr m i nurth-raqt tnglnnd, rn 111 P ~ ~ l ~ i ( ~ i ~LT / ~ i i +lirrrr~1>cor1 ! ~ ~ ~ ~ ~Rr$ : Cr~iiycrt~~!i~ I> 13. 1 1 , 5tr~1ti>t1l~t~j1cr1c>. dllrrdtb (cd. I>.F. Toonlev), h o c ~ r vn i Spti/fcs:> ( L ~ D.E.(-i. S 1 3 t i ~ p:and P. I<. C r ~ n v t ! > c r ) , Ernnotrnc l ' ~ l ~ , c m t u l o ~ ~ *, ut .~: dM u l ~ r ~ ~ l Spcrial u~\t~ Hl.~rk\vcll.Ocftlsd, 537-4'. Publicadun NII. -30. Tl~lcs. O k l , ~ l ~ c ~ tSEPM. ~ l . ~ . pp. kI.~qi~r.I ) . A . T (I1+*.')) i~r,ii/r~ty,~i/s -/~I*IPI t11r L J p p t l(>l-R. .-lr,~frrt~/i~itr srl~~r..<.r,rr!rC1~-i/t>~lri.i~11r) i!f 1/14. Cirf1;r1~ ~ l s t r i c l . Thoma<. K.IF.K. 2nd l i r i L \V -E (IC)'43) T l i c q k c l c t n ~ i .ii,~tFir~~i/. IR,1n2 , 7'1- 1 3 . I f . i l n c o n ~ c l ~ ~ p I i t S c no!c ~ c r v rpncr: a finite S C ~of ( x - p t ~ ic~ l ~ ~ i g n cEl~oErir~or~ . 47, \Innnyaphs (tnnnrrmnlr. prtlr-ccl~~rr rr+trcd to it1 text) 34 1 -60. Harpcr, !,).A T. n r l t l Ivst<-~nh r ~ ~ ; ~ l ; ~ ~ t ~ ~ ~J tT nI I ~l ItI ~~I !q!!/' i s ttltr s , (Aott!qii (11 ,\Jtlrll1~, Er17lty). fll7ii P t ~ ~ ~ r , l r i o l lnp-.~lSr)cucrv of tl:jiv. J.-L. (14X-I) A t ~ . ~ l y \c-I,~d~.;ti~lur c ct Trilob~tcs:~ : t l A n t c r ~ c n No. 67. (;rn!ogcal F n c ~ c b of Plrtlcnrn, pclint LIL. vuC. h / i , l ; , l 17. ( ) ! - ( I . (L; ~ l u rl,idlmi) ablc, well i l l u ~ t r ~ t ~ ~ T ) 1 I 1 ~ I I , ~ ~ 1 tC'. I . I-I.,Aucllr\--CI~.~rlc~. M.C;., I5nl;wtt. M.G. 1.t Licylcr, A.M., C;ock\. I..I<M. a17d R ~ t ~ h ~ cKhK, ,A!. (I1l7H) A gtlidC t t i \tr:~t~g.~llllii. pmrcdurc. (I')t,Hj T h r c n i i i p n ~ i t ~ r ~ d rn~ d ~ t n l r r u r c r>l' Lowrr t;er~lr,~qc.tlSncicn. nl' ILlmclnll Syrr.i.rl I
w
:
[(lt--
Evolution and the fossil record
t>\'cr ~ l l i
11C)t PC' l i i ~ \ t y ~ ~111 ~c Inttr !I :\t~cinr
I~II~F
111 11I?I11\ , L71
111~1
cytolnl
nr!:.lrll\rnF t3I,li I c - 1 1 ~ ~ it1 -
i r ~ - t r F>rniip tllcny ~ ~ o l ~ z c t = ~ cyccpr ~ ~ l innlcculnr h i o l o p Arc tranxfornii~~g and .1i4diiig ltu T h c rrcc t h i ~~ynthc+i+, nncl nclv v i r w ~o n tlic Iiaturc. o f tlir 1 1 wlf-rtylc. ~ gciic Fcun to bc r h,inyi tig our wlit~lcconccptinii ot tlliclltlv d1y Amon<-t .ill the sricllcCF ~ 0 1 7 c ~ m~sirll ~ d nrg.inlc ho\\' organisms rv01ve. .111rl St.llllc.!, cx-oluhr>i~it is n n l ~palncontolnp thnt har thc Tn thc following tcst. tlicrcfurc, I present fintly .I TI<-cdudlicrt ~ i t ~ r q uper\pccrivc c of ~colo@cnlurilc. It i q thc rock qunplificd account of classic neo-Dar~vinl;ln evoluAltllnuyh rccord nlonc th3r providcc nn Ilisrot-icnl pcrspcctivc tionnry thcory. .iirncd p.~rticul.irly ~t xtudents nf thr. f,irht.r trl for tllc ~tutlvnf cvnlonon.ir\. cvcnt+, ,111d thi< ciinc Enrrh rcicricc who rnny only I T J V ~n Jimitcd backd i n i ~ n ~ i ncould ii ncvcr l ~ a v ccnmc frrmi 3nv other groutid in b i o l t ~ ~ T y .l i l ~inuludc~Foinc ~rlfonr~at~crn \vt-rc. nlaiiy source. Ac~-orrlin~Iy. thc input of pal,~coiitolog-yttn or) thc impact oi ncw i n h r n ~ ~ r i ofrom n mc~lcc~~lnr tll.~t,1111~~1+ of tiiric cvnlurinti t h c o ~ Einq hccn In unrfc-r~cni~rlrng thc hi$- grnctic?; o n c v o l u t ~ n ntlicory. This section i r not i i t > i ~ prL,-c ~ to^ o f iik. in inrc.rpristlrrG p,lttcrrls of ci-oltttrori irlrcrlcicd tn be o o m p r r h r n ~ ~ v[lor c , does it pretend I j,lnvlbl I l l 1 (c.g. adnptix-c r.~di,leinncl ;\nd lirrcc rlf cic.i ~iioclrni h d q tn he ~dinirtcclt11.1r rhc For~il record i< incomwhcrc v i r , . , Simp~nu(151431, Maynard Smith (1'375, FrlttlltBVC J plete, ~11dto iiitcrprrt i t can in \ornc. inllyllWh 'Ilke trying to rc,lrl .I di.ln7 wilth h ~ l thc t pagc-5 13oh7haiisky rt al. ( 1 9771, Garnlirl and Vilics (IOX7), lir>nt rhc 11 m~qcing' (Shrlilon. IOSX). Yct it 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~c<, hullding upon thr nriyrlal F P C C ~ C F had contcptloilr of C~IJTICF r)an171n. Mnr~vyt.ar%AFcl the ct1clc'. Crrc)] fint rcal n~ulttdi~crplinnry nrnnlgnm of dat,l u7ar pubh n r m tvl Ilrhcd 3 q Ft,~11tt~totr:tllr ,IrLldt.rrt .Cyrt~/tr.siv{Huslcli., TI1115 the ( 1912). From tl~ichigl-rlv r u c c c ~ r t i ~attempt l a t lvrldT h c thcnry o f cvnlurmn liiikq togcthcr J nlliltipltc~t (Y-F;. l l l t l ~ i t ~ gtosether iiifnmi.~tiou from v:\nnus sources, thc of hioln@i_calphenomei1.l and i\ undcrp~nneilhy a ~ndic,-~tcd .I 1x0-Tl_ranwniansvn t h e i t t;rkt.r it< ilarne - ~ n tdl ~ i qig t l ~ cevidcnuc of thc gcoIo~c.ilrecord. I t rcmainq ,I prncntc in thcnr)., n o t n Frovrn f i t . Llr thc irnnirncc t i ~ l t \ r 3 ! r \\
to .I
Darwin, the species and natural selection
27
' v r r n'l~rsl~ rvolutio~l ~ r v li,irlgcr Ii.lvc t.~kcriplncc CCIV qliecles to EIII;LII ,Irc,li only \i.llcrC c~r-I111nd ;ill:.< liot penllrt thcit ~lircctc~hsrrv.~t~oti. Tllcrc 14, I7ccome addptcd to i t 5 utvn cr~\:irnnrr~cnt. I~ll\vect.r.n o ut11t.r ~ F ~ r o~nr l i i c lc~icoinp.isccc ~ $0 L>nnv~t~ \vas p,~rti~-\rlarlv it~tcrccrcd i l l ~clcctrvc rrllltli ai-rtl ,tee-ord~ ~ v i t ht l ~ rc v ~ ~ l c oicr>inp~rLitivc ~~cc brredinp of ai11tllnl.i;md plant5 ur1dc.r rfc>~nc.cricnnot~. ~~lrrnmr., hiuclrrmlrrrv , I I ~ L ! p h v ~ i o l o p ,of ~ C I I C ~ ~ CHI r rt-al~zcd[h;rt tlnr prcscllt g c ~ varicry t nf r l c r ~ w* t i ~ c f ,~.td c vrolr~,g-v,ant-l t r f dit- rcl.~tir)ticl~ipc hrtu7ccn c.imlr had bcct~p r o d l ~ r e d111 only ,I fc\\ thnu.;nnd crqiil#nl, prrcc~vctl b\. ~ . ~ Y L ~ ~ I O I IW I Vh ~ r r , ~ the‘ < ve;ln f r a n ~oldy one o r ;it must a fcw .inct'str.il hpr.;. tr~drnccIn .;chrrc ot thrw ficldc I.; clrcun~st.inti:iI. Hc cancltided t h ~ t.I g c a t pstcrlrial firr 'tic\c-rnt ~ I l c brolyht j ~ toqcthcr ,lnt{ ~ntcrprctcd~t hriilrlr up wltli nlodificat~on' irlttst in all . ~ ~ n i i i i ,3nti ~l~ rlr .r rhporv of tioriii~J,~blrc o r ~ < c y u ~ ~ ~ and l c c ,.I\ plant5 I Z , I ~ J C I ~c ~ ~ i I1. l ed cpreiicd up wch ~ r t ~ t l l ~ i ~ l r)rl!vl~,~nckv 11aq s,ilJ, 'Notliij~git1 i>ir,lnp ~rnnkr, L > ~ ; ' ~ C I TI Il Il i~r . Id C ) ~ I to tllc 1,rli~'ftl~nt ~ 1 1 7 ) i l . l ~ < S ' I ~ Te~ x i r p t in 111c Iiy11t o t ' c ~ r ~ l i ~ t i r ~ r ~ . ' proccsscf, t l i o t i ~ lprob,iHv ~ nil ,I clinvrr timc ~c:rlc, Ille rc,ccrlt Llcllr ,~:tcrnptc r o Jiccrcdit c v u l ~ ~ t ~ n11nd n npcr.~ted11) t h r \r~ildstarc; i l l r9rhc.r \\,orcls 'il.ltI!\. ,clf-wled *crc.,~t~oei >rrc.r~ri.itr'IIAVC' I>CC-II \ O ~10- u r ~ l~ e l e c t ~ o n-l'li~rs ' bruudmg cspcrimcnt\ .irc rhr lll~~ntlv CI1sparc11cd h j 1.lu~t(1 OS?), Ki1~I1t.r1I W 1 ) foundnnotl nf cl,u;s~ral yrncnc5, whcrt. rhc mcclin111tl Yt.lnlev ( I c ) X ? ) t h ~ trlo f~rrt11c.rrolnrnc-nt niciri o f hcrcdiq rs rindrrstuod 111 term\ o t - ~ r cct'rcct~. n p r d l d Iicrc. I'lnnmii n r a F also r o ~ ~ u c r n ewd ~ t hp,llncontologv 4ltliou!:F1 Cliarlrc Llanvij~I < y c t i c r . ~ l rcgarrlcd l~ ,r< hut hc f o ~ ~ t idint i l tllc hcqil rcsor~ln r ~n:ly changu. He I1.d Iic1pct4 IO find c v l d r r ~ c r :lilt J I I I ~ T I , .~iid A ~ F p l n ~ i th.lci ~ L l l ; ~ n p -olrCr ~ l Ion? pcn- nt' gradntint~nl chnllqc h r h ~ . r c n, ~ n i t n ~~pccicc. l of I\ of tirr~c~ n tlint d n c n t\pc\ crt '\prcic\' h.ld ;Inwil 'ir~tinitely i ~ ~ i i n c r o utr,~r~'riticli~,~l s l ~ i i k ~~'~ 1 r 1 1 1 ~ c t i i i c !tnnl prk.-culftlnt: O ~ L ~ ST. IIC'\C T ~ I TIVLIT~CTI ~ ~ ,11111 ancc5ror5 tn d e s i e ~ ~ d a r land t ~ nf \ t r ~ t ~ ~ r r a p l i i c . ~ l l ~ Ihnnltl h~nlcclf ~ J c i i t i t j c ~I lI I A I I Y dl tT~,rct~tpoint; LirrangcdS C I I ~ Ss l i a \ v ~ ~'clrqcrn ~ g t with ~nocIitic.~~tor~'. 1v41cl1cil~~lil 1~ rrqnrrlr-cl '1.; cv~dciiccfor thc o n ~ n I n Lct. ht did not I;rnr.l w11,1t lic I1.1rf cspcttcd. cjt' ~nodrrn EPCCICF jj-or~l prr-cYi\tIIiy ,111~1 morc m.lnz*in as~lirn~cl that the inrpcrfcctioiir i ~ tl~ci rock ~ ' n n i ~ n \,iiwrrrnrr. ~c 411 tllcqc .Ire acccpterl tr~d,~y, sccr3rd 2nd tllc lirrlitntianr of ktrnwlcdpc . ~ that t rlrnc tllwr~chr l - f i n d nnd ~ t 1 J ~~ tCl I T . I U ~ I I OIllttivn.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%ipti
an\?ll th.it ,111 luv~nqclrgatllsiilr\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.tcr<'tvrrc inl~t.trtcd,n\ 111 hlc unknown, 50 that l i i c views oli tlli5 wcrc ~prcul,lt~l cl;i\sic prvrtlilntc th.ir thc 1 1 ~ . ~ 1 ;trf tltc s~mffc had ~ t l di t i ~ i l f f ~ ch~r~tlir r ~ ttllenry to hc fccn to v.lorl: Ilt.comc lo11gc.r in rc\ponuch t t ~.I ' n c d ' tc reach Tlic pioneering work of thc Aunt npcrntlny to prtducc. dpparcnr ity ccccntial tcr thc a m p l l f i c ~ t ~ oof n I S J ~ V I I Irt~eon 'F 'L~nrarc-kranc h a n ~ ~ - \ ' 1J~nv111, ). ctn thc t ~ t h c rhand. p m v ~ d c d.I lopc.11 ~ t i drc\r.il~lctlico?' f i ~ cvnlunoiir .IT L ' I I ~ It-iiic ~ ~ fhdt ' : h,+s \tr>nci thc tcct a( time ,1nt1 Inheritance and the source of variation p r n v d d c ,1 Fr,lrlins: pc)int fhr I.ltcr d c ~ c l o p n l ~ i i t ~ . ~ F I Iin: IS The f~dlt ~ d co f r > c ~ n ~m~ iq~~~xvt~rk r' q 1350 WJS In the cells of;ill euknrvntes (a11 O ~ ~ : I ~ ~excc'pt vlnicee, bactrna dnd C ~ ~ I I O ~ J CtJ1crc ~ C ~ IIC, ~ ) ()I! t / 1 t a [ l t Y T f ? l O f ~ ~ l l ' i / l ' . T h c 1narn prjlrltq of clic dicr>n. arc rtniq,rl~rtc~nvnrd. hodleq. the chromosomes, wl~iclinrc t1-iaJc of prnt c ~ n;111rl deosyribou~irlri~acid (DNA). I . A ~ ~ i m acpcrin l rc1,rcxlucr ariorrx r'lpidlv than ii ~ r r r d r dt n m.iint.1111tllcir nurnbcn. Anlrnnl popilsmooth endoplasmic reticulum lntioni. lio\i~c\~cr, thnligli flr~ctu,~tins, tcnd to rcn~.linst,lblr (Hc-rc hc ii.,e< ~ntluc~rccd I?): tl-tc En~lisllmnn M.iEth~i\. tvhn I ~ a d w r i t ~ c noil t11i;i vacuole ,/ mitachondrfon subgcrt rIl~irrgILL^ :~trd(~vitl~lil nlcltlb~r< oftlie qamc qpccicq) tbr IrlJrcs, it'thc ch:lrdctfr\ tli.jt indiv ~ d u : ~bcdr h ,\rc c t ) 1c. t r . ~ ~ ~ ~ ~ tn~i tlic' )t t ~timt c I gcncratlon. 3. Wir111n s p c c ~ cnll~ .inirrl:~I\v~n', 2nd th~uvarl:ltlon Ir ~rllicri rcrl. 4. It1 rlte ~tnlgglch r lifc t l ~ c > w~ndividt~;~ls host tittcd to currivc 111 ,i particular ciiviroii~nciit ntc thc (>TIC$ tu IIVL.nnri tlr tc-pl.cbdticc. T h c othcrk Arc wc.cdecl out in tlic intcliw rr>talpctltir>ii. T11c hvour,~bler h , \ r , ~ r t c n < tth,lt ~ c ~makr >llch sliwivfil porsible ,Ire it~hrritcdby tiirurr yvilcrG~tionr, .111d thu accuintrlatic,rl nC ditkrcrlt th-otir.~bIcc t ~ ~ r a u trm Imds to tlic ~ c Ir,ltiorl p o f ~ p c c ~ c~sv, c l lddaptcd rtl p ~ r t i c r r l ~ chnvircl?lrrlcnt*. r Tl~ic19 tvh.lt r).lncitr c~llt-d'n.lt~lr.11\clccrlon0.
t h ~ iirw t
I
All thi\ ccchrn
~ I ~ L13,1nvin'\ I L ~ I carli inqk~ncc,, u - y c d thdt n ~ n v i nhad 11ut rually c l ~ c r \ v k ~ ~~ u ~T J ~
C T I ~ I C F ,M l v ~ r for t
actually accumulated. only huw d ~ o s ca i i i r t ~ ~ Icw l< fittvd to their ~ ~ ~ v l r o r ~Cnlcd ~ i i tco~,unrivc. ~t I11 t l ~ i ~ they \~c.tq: 17rlt ~ii1~1>und. fi>rt l l ~ ,n105r S C * ~ ~ I I I I~C ' c ~ k ~ivwof tlic r h c t ~ r y , Ir pr~,
rough endoplasrnic reticulum Eukoryote cell structure, with orgonellel.
Dutwin, the species and natural selection
29
Tltc DNA ~ric\lct-~tlc f i ~ r r .I~ Ir>ily, ~\ n~,rC t' nuclear R N A it r-ctnln? thc orpniz:ttion r d t l l c on?L , ~ c ~111 \ , PNA, .lrc . ~ d ~ i ~ ~gu.lninc. iic. ~liviii~rrc n r ~ c d 1i1,il 13NA. but \vllrii nuclcnr ILNA is rcprcrc~..;t~rnl-r.rr t t ^ pnscthlr- cc~mhin.ltinn~111 rn,!jor n i u t ~ ~ for p l ~~ ~ ~ n l r r ubliaol r cl~y.but thc u ~ n l c l I ~ : ~thrv h cnn cu~ct,coding !I;x ~ ~ i inrhlch d\ arc produccd arc o r ~ a u l z r dand llullt ,I, , m,lkr protrinr. I t h . ~ rhrcn F I I ~ ~ V I - It h t up into f i i ~ c t ~ 1o0~~ 9~1d1 and % whoIc 13odlcs scn1alnc ($1 I < nf t l ~ r r cb x c c acting t n ~ c t t l c rr-nde fim n r ~ enf t h r Irl,un tarkq for ~ r ~ r h c c t ihlolo,~ l~r f t ~ rthp x ~ r ~nnrl ,icrd'i, : ~ n d rhcrc : ~ r c tnnrrl. thdi~ tiinlre. Some p r o q e u has alrcndv hccn 111adc; 1 1 ic I ~ L Q J ~I C~ ~o ~ F I I cc~rnl~lr~ntinn\ ~Ic ill thi3 'bacr-mplct' [low krrowu t h a t fame penes Are r ~ t ~ ~ l i ' t l l ~d, tl ~l 'd, \ . . t m ~ r tn i m~kc all rhc htnci,l-uc ,ln-unu ,ICIA\, a114 to concrnlcd orllv with the yntlwsir of rnntrnalr, '1.4rhrtri try iir tlic nzht orrlr-r m iiinkr- n specific o t h c ~ r Are 'regllarnr' Srilrs. w-h~rh tontrul dncl yrs'tc ti. 0rgs1112c the c t ~ u ~ p n ~ ~ i. ~ i cnidcdirrct thrir iiuildiny. Irc r\ntt1c~i7cd,illlt in rhr iinrlrtls, but 5kirh gellef releaee cheiiiic;r~prnducts \\.hlch st,lrt a t ~ t n cin~rhc ~~'11111.11.r-vr~pl~lsin. ,md this wlrolc ho5r of c o n ~ p l c sre.lcrioilf. In ~ O I I Iklildt ~ bi , O I > 1 1 1 1 , ,111 tli~,infim11.1non h , ~ rtn bt. tran~fcl-rcd of dcveloprnent the priies ,Ire switcIlrrE on nnd otf oi tllc ~ ~ u c l ct~o rthew r slrcc c ~protclu f .;ynthcw. F n l l i ~ w i n~~h l c p-occ5s of sw~tcliingolr of stnicniral gcricy rs FIVCII I V ~ I C M .I rr,murription', ,I fu~rthrr ~ r ~ ~ l ~ c(mcwcngcr ulc wnsnr gciie receive? a n JpFropnatc 7 tllnnl~rc. 14 t~jnnerl.ivhir 1 mcwcr oiit of' rhc t ~ ~ ~ c l e i i c , Slriiplr orcnrusrlrs such A S bactcr~aand f ~ m p have I tlir~utghprrc.5 ln thc nuclear u~crnlir~ir~e.cets of;l$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 1. <, form, n.i I \ I Z ~ I ~ I T C < hv F C ~ hi^ iiififc-t~~iit .q~pp,lr.liict~ of cell tiil~crit\n a ~cvact copy of tllc ~ l i r n m n c o t u cnr~! , - '--J-7--, :.b. vcgctdhlcr polvil 111 r ~ i hnnd pimr ~ n i l r ,rcqycc: -4.. geirllc5 111 the parcnt rell. l'111.i pmccqc, knorvn 1 :L.?' rivclv. T h c ~ n h c n r c d~ciicttt-trrntcnnl 111 a i ~ vo r p n - ~ n i t o s i s(Ftg. 2.2). is ct?;ctivcly thc c:uiic for .)I1 t ; ;- ",;d-~ . ,- 7: I ~ T T Ikntnvn , nq tl~c* genotype, t s smc-~cctIlpnn b!, thc vnnintic cclls in niiy nnc hodv. *,1. 3.. ; 5: 2 rtnJlronnlrrlt ( p r r l l v h l ~ tl~rfrugh rciirrrr gcricq) t r ~ W h c i ~n ccI1 i< nnt d~vtding,thc c h r n r n c ~ r n n ~a c ~ -P--. ---....crcntc n dcvc.lnp~-d~t~ri~virJt~.tI. Thlc il~divldu.~l, thc I,lrgcly inv~viblc:t h i ~ir knnivn 25 intri-yh.lsc. A< 1-0 . prodt~ctoi botli Ilcrcclit~,111il c n u i r o n ~ i w ~ rr ~ t tlir . dlvlrinn ~ C ~ I I ' hinlo$ical F , F h i n l n ~wvralc thc rhr phenotype nlnsnii1es. c ~ c halrr~,~dy dirridr-d Into tr+m c+hrnn~,trl Interphase S o n i c t i ~ n cF I~I I ~ I C qC17CS t-017rr01 ~itt' tc) t h r broprrrph,~w.t.rlhcrc two ccnrnolcc moyc to nppnn~-r~trt>t~ of a pnlcq nnd a ' ~ p i ~ l ~ f l(>T c ' r r n t ~ i n t J i ~ c . ~.ippc> d~ : n A t niet.~ph.lwthc ntirlc.i~-tiicn~hr1 9 /--*\ pnrtic~ilal- c l ~ . i r ~ c t ~ A r. . s o ~ i ~ cgericas .ITC h ~ h % , ~ {thcm. pleiotropir; thar ~ u , tl~r'y i k h t .;vl-cr,ll cli.lnctcrc vnnlsJlcs . ~ n dthc chror;lmolnc< 11nc up .1Jn11g t l y -\;. . uncc tht. v n l c gciir- prod~rrtqmav uwJ 111 d~ffcr- m c d ~ nplanc, ~ ~ ~ 3 ~ 1cpllthng 1 111 h 1'0 FO that c.v c n t biocl~cmic~il p.1rl1w.a~~. A p l ~ c r ~ c ~ r n c\n~oli ~ i ;c ~ h t chrcrt~r~ttd bccortlcc 3 Jnughtc.r cl~rcrlrlor~ir &in1 311 1111111010y04 Cornc? x ~ , i t ltlicir ~ gcrlc. Tlic c ~ i i r ~ ~ i ~ i o IcI oU ~ I In I ~cC TI$ p r r . Herr n p t n tliert' ; l r t f011r ph;lr~fbut qoriic k '. & . / a l w ~ y cCOllGtdrlt ti rr tlic I ; ~ C L - I C < I. n l i ~ s ~ i i . i ti lh~c ~ c, I ~ L ' thccv arc ' i ~ ~ n i f i c n n tdlft2rrnt lv frr>iii tlicir rnltnl 33 pair< clf slirr~rirt~r~ntic<, 1%hilt ill the' f;uit flu' countcrpnrt%.1 )unng prcydlnsc nf tlle fint inciol telophase Drtlst>pElillrrhcrc arc. t i l t ~ sp , ~ ~Tlic n . tmc~ubcrsof c . , ~ c - l ~ dlvikion. w l ~ t n t h C t-hrnnlo~c>~iics pair tip, thr pair arc. all homolognus. 1.r.. somc<Jrc IC~C*IItic31 (XX) 111 I ~ C ' o t l ~ c ccs r they .lrc c ~ ~ l ~ ~ i i paiw g i i l gnitfir q.inw lcngth w ~ t hi t v honln d~h~it~li!.ir /?CY). T~IC ~ C S C ~ I T O I T ~ U ~ ~ +L?-~dI m C
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Darwin, the species and natural selection
.-..'I
, .
interphase
"'..
.
,.: "'
centromere
prophase
metaphase
telophase
!c~8:?.2 The rtnges of mitotic cell drvision in a wkorptic cell. For explonaAon see text (Modified from Kershow, 1983.) p m c t c r froni a rillglc p.lrc.nt cell. Each iriarcri,rl war exchxlged durir~gcrossing over. c ha)?half the nu~nbcroKp.rrcnt chromo~omes: T h c ~ c panlctes containing unpaired chromo'1. ~'lronrotornc\ arc uupnircci, ,1nd none srr iclentl- \ornrs; arc caid to he haploid, and the chromosnrnc , ' : l l . l n ~ ol the p ~ r c ~Cl t~ I ~ O ~ ~ ~ SIIICC ( ~ I T genctic ~ C S nuntbcr 1s H. 'Ir
I..
32
Evolution and the fossil record
~nterphase <--
-L...
prophase ,--,
--
--\,
7 -. .. %.
.
,--
" .
q,
./I
!; s
C Z s
., ., -
/'
'\
.,..-.I--
<
.
'
rnetaphase
anaphase
\
," chiasmata
second snaphase
telophase
gametes
Figure 1.3 The stogas of rnerotic cell division in a eukoryot~ccell. For explanation see text. (Modifiedfrom Kershow, 1983 1
hn~vever.arc not ~lccccwrilyi d c ~ l t ~ c fur ~ b ,tlwv 1n.l V[rl~cric ~ lire* p icrtil~zcd17v cpcn n tllc I~c)riiolug~~:i,us CX~FC ill a I I L I I T I ~ C Iof ' C X ~ T ~ S S I O I ~ Sknucvu '. ;rs allelc~ rhron~usomc.\Kri)~iicllifi-rc~~ t p;lrcrl tf coin? tnger11t.r ar~ctpar up. In t h i s proccs\ of recombination ,I nv\\ eacli nf tvIilc11 Ir rl~ghrlv ditfcrcnt cherzuc~llvJIwill lead to thc dcvcloprllcnt nt' diff>rei~ces111 tl gc~lorypcrcwltz horn JL~T.IJVt-stqt111y~ C ' T I C \cieriving . l~~~ I.dI1L' fkorn d~tlL,rcrrt
Darwin, the species and natural selection n.
111
thc heteto~ygouscondition
Evolution can thus he cnnudcrcd as haccd upon frcquenctes tlunugh i~atural
rrrnt kltidc. Ilr .und rl?. 1911ind~uidi~;~l cliatigcc in gcne I*'ir><~trg R R will Iln\.t. brown c y ~ c ,w h ~ r c tlic . ~ ~ ~~T~<-~iotl.
>I
at' 17 will hc blllc' ~ v c d ULIZ . thr c r j n n h ~ n ~ Rr or r R o*iII.il\c7 lc.1d tt'r Isri>\v~~ C ~ C S .fur t l l ~ I . I,. rJ :,. A,.I: 11 Ir uwtllinant nvcr r. 2nd ~ n . ~ < irc k c cffrci. 11r i,>inultiplr \~~, afict from tcrls to thousands of base pain. and It-,, ctc. - arc all cclnqcqucncc\ of dipluidv. III chrernosomal mutations. wl~rrl~ a r c the tnoqt inon, ulally plants increase thctr porcnti.11 vari1l11lbr1 by polyploidy, 1.r. dupl~cationof chromorstrcrne kind, may ~ n v o l v c inversion, rearralipe~ l i ~ cd r , ~ l o ~ l ln l ggreater gii~lnbcrof l o c ~L)r allclr<. ~ ~ i c r l taddition , or dulction o f long rrqucnceq nf DNA. Thcve t n l d to llavc larger-scale phenotype I'ld
'9,
I,'
,I.YSO~
itrrir
33
34
Evolution and the fossil record ~ c n l r may , hc rl~rcLrt ~iiohilcg c ~ ~ c clurnunts t~c (vari- is not I~kclyrh,lr inutntmlrs xvill rprcad v r n (1st 111 a owlv nnilicd >urnping genes', 'transposons'. n r Insgr pnp~il.~tinr~. c v o t ~ l i thcv arc dom~n.lnt. z t ~ d cvrn 'molecular prasitcs'). \\ liich drc DNA r n u t a t ~ omtCr ~ ~ ;lrc high. Hut 111ost t ~ ~ t i p~o rp l~~ l , ~ < ~ ' i ] ~ l C l l C " S .1h1~ f < l I?FO\'C :1111?1gC \ I T C I R I < I F C I ~ ~ ~0C r< 51.0111 nonq ~ r c .not cvcillv d~ktnbutcdthroughour thr'lr one c f l r c ~ n ~ n \ n ntcl ~ c .rlj~>tlrcr.TIir\. <.In altvl. thr qpcc~cc; raijgc. T h c v tcnd tn 1.r rubdlr.lderl intn wlielc gr.rtc.tlc str~ic.tlri-c.uf ,111 org~ilic~rl throt~gh rc1,itivcly m , ~ l l units ot ~ C I P L I ~ : I ~k110\1711 ~ O ~ , ,I? rc.n-ranyr.nlci1t o r c x p : ~ n \ i uc~l r~i t < TrrlctIc nlcjteri.ll. garnodemes. Esaralplcs c,f si~clinatural yop~il.lr~orr Scmc tcilcl to att,~chthcmrc.4vcr tcr parricul.lr wcr; y r o u p i t ~ garc r:~bhitw,lrrel15. cchoofs ut'tirhcr, par~ri othcn 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
\lrnultl 11c n n r ~ i i ~ ~C\I tlv : r c ~ hic.,~ > ~ 13c111g Cut C ~hr)t,c.Hut 111.1y I\c cq1 It
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l l ~ t l f l 1 ~1 t
Darwin, the species and notv~olsel~ction
genetic drift. hrcu~nesilrlportarlr in tl l c ~ c51n,llt pnpuiatinn~. I'his n <miplv ~n rvnlut!c7tzap ch.lt~::v tr~rtuYdt i c w bc clC,lrthat w n l r k ~ n dn f r c n l ~ t i n iir~ m prne f r r q u r n c i c ~ttirough r;~rldn~ I I cl ~ , ~C.n .cn r ~ ) rt,\~tn111\ cwcnt~nlfor syc.c-jr. Itt mnv r . hc .I l ~ l c n twrthout , selection bcirip i nVolvt-tl. Hcnt c c r*r.l~g,~pl~ir .I] rct~l,~tron. r.c. cnmc pnrr o f a pnp~llntlnn tain locnlrzed pop~iIarior~< iic~ct'~~cIt'cl j; c > I I I .ill 1~11r1,ll cut off frtlnt tlic rest h\ ,I hdmcr, 2 % norcd p ~ nr~ few r paln IiIay hecemc qurt~ ~ I I ~ I I ~ I C ~ ~ L L . r. But thcrc are ntlicr k111tIx n f iqrhnrlnn \~,hich may cvcn dlfferentiatt. ns new rpt.cic\ H I C ~ C ~ \ br r.yu.lllv c.fTcctivc I ~ ccrt.~llt I c.l\cs. Thrrc. 1% bccnuvc the folindcn o i t h e pclptilnt~cll~ l~np!>cnrl-l to lgc.11ict31~tio11. it1 \ ~ h t i l \pcci~'$ l ~ C C ' U I ~ ;1~1~1pt~d IC r:lT a pamcular wt of nllrles. 1urruhobjtat5in tllc sartlc gct~cralarea. I'hyioIn m y gcnc pool thc etTecn nf all t h r f;icrorr c r r w al nr ti~olo~,lval t~ctr~ niay n AISO IFOIJIC pnp~ila- ~ ~ d c r cmd f,lr are conrinu~llv lIIFCrat-tlll~,tllnr~ch ,to thdt tlicv ~pccintt',c.g. C ~ . I ! I ~ L = Fill t h r tii~ilng m.iiiv biologists brl~c-ve tIrat Inon r l u ~ t a l i o ~.Irr ~s IIlrccdirlg cvclc ttr mcrclv qizi- clifkrcncc5 prcncutml ;lnd ocillr a t a more ar lcr5 cpn5t;lril rarc. In: cclpul~tion.Ay.1i11. ~ I ~ C T IIIJV C L ~ c .I ~ l ~ ~ i p Gcnc lc f r c q ~ ~ c n cw ~ ci tr h n ~the gene pool n r c rclatril n r 'tic incun~pat~biliry; icrtilir,~cinnrn,~vtnkt plncc. thc chrornnsomc lcvcl to m u t n t ~ c ~ ncm\rlnr: , ovrr ~~LT,I~Y {>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 . :$, at Ica\t. sonlr of wllich may runrivc. WI~cre Thc ~ntcnsity o f hclcctinn unclcr t h c ~ elatrcr ell-cum1 1 1 , : r a i Y ~I J I ~ C ~ O ~ L I I . I I I C I I ~ I C hccn bur ivipcd 5 t ~ n c cis~ much LTCJ~C~.. At F I I C ~ nIrlc5 rhr'rr 1 ~ 1 , i j r 1 qrinic ~ ~ t . ~ ~ t ~ otllr p I iqvcirs c, nidv rcc0vc.r occur directional selcctio~itnw;~tcivollr cxtrvrrlc ir, 1 ,YFIIIL ,~q.~in. ~ !i(>ue w r , I L ]XI> p.iqscd t l v n ~ i q al ~ rcn.pc. in n popul.~tion can l>c wlcctcd nkqi~i%t, ~ i thc. ~ dmrn ~.xtrclnC\nrr positively %elected, S'iwnuu!: twt, dift:rt-nr spccicc frcrtn at) 0rigi11~1t l O ~ 1 7 l d l l d~ ~ ~ t n h t ~ pt rndp ~ ~ l ~ i t l o l ~ . Genetic drift: gene p o o l s One \~oulcithcrefi.,rc cxpect to h i d I r i tlic firwil ! o ~ ~ ! l u l n r ~ ir c ~ ivi c ~ ~r l ~ n it l l ~villcuntain otrly n rccord relics uC l o i ~ gperiod5 ~ ' ~ ~ c c ' \tnl)ility, Ic's irhp, .i%. ~!. n dcuidui~ls.ir~lplcof thc tot.11 qcrictic v.iri- drntvd by hruaks rcprcccnrlns tirirc., \vli~-ti~llccin~iofi ' - v i r l r l ~ r i itllc rvholc ~pccic'i.i.c. only .I fractlun W ~ very F rdpid. and t l ~ r spin l qtablc pc~iodq.T h n ir . ,i\; wcr.lll >rlic poc~l' n i thnt ~pccics. 7'hc yen7 uticn ~ v l ~ ir a t fn'ound, a$ will bc cuplatl~ctl.IT :Ilk: :11< pnpul.~rion.thc f c u ~ c \vill r bc thc ~ I l ~ l t ' xw n i e Icnptli In thc rwxt section. , I ~ J I c ~ dh\ivt~ I t o fit~trtgcncr;lnr>tl<. I11 a n y evolvi~ig5yctert1 thc lrlnrc nt~cmcnvirnn!I ~ I ~ rtlc1nsi< II (>lily n ~ i cc h r c ~ r ~ of ~ ~J p~ i or , ~ nrnt.l~ts ~ tllcre arc, tlit grratcr ~villbr tllc* prr\\thil~r!r ot :. ! I : I .r,~lirln~n 53tnplc c ~ i ; ~ l l ~1%' P.IISCL-CI ~ ~ , AC~WII to A~tErentiatiot~. Species thus unn be 5a1d t o arlw 111 ":, ,:,~rnctc\: , 1 1 1 ~ ;~llclcxui ,I [illy popula tinn rhnt scrpollse to the cnv1rclnmcnt5 ~ c t ~ l a l avail.~l,lc. lv Iiwr t~~~-ol~~hincrl in 5-rtilization arc. rhcrcfnrc It 1%well kilown t l i ~ rlatural t ~clcctioni 4 ~ p ~ r ~ ~ i l i g ,.I 1::.1 .p~r.i;~lcvolut:c~n,il?;proccsq, knt>wii AF ~t thc prcscrlt tirrlr (Ei~dlcr,Ic)Sfi). Evidcncc fru- thi\ tlation and species formation
-
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35
1)T
.
36
Evolution and the fossil record
Tclp1c.11 ~\.1111plc\C O L I ~ ~ tuikcrcr'. The w h o l e orytusr71 lr n sinqle rv\rc%nl and thc ~lm-pI~olol;.yar1c1 Lir~znnuo f itc cntnpui-Hrn im-llidr t h r rrsc o i rrb\muoim,ltcr\l\-tci mbhitr o r , ~ i i r i h ~ c - r t ~ z - r ~ .h.lctcri.1. ~ i c t ~ ~ ~The ~t c,l\c o f pnrtc h.we to bc compronuwc amonpt rc>rtyv,tll~ 1 IIL.b.lt ,ll ; T l l ~ * b l ! \ <<> th dein;~ndsand bascd upon Fcnctlc and phyic.11 ni,l 'inrFil\tn,~lrnc%l;lnr\~n' in ~aic)tli\II,IT hccn r.utr~~5~\.elv , lI>lL.. IIbj-0 .;turlri.d .r~-td i t . Hcfcrrc. thc Industnnl terial which tnay ~tsclfhc lunliiny. I , > t I l ~ * Il For csarnplc, the lenrer of .I tnlohirc'c cyt- .I. l-lcvtd~~tror~ In the. north of E~ylarld,the nloths o : g v l l t , 'blltl c of t l ~ c,111' inhabitir~t:thtv I~c~l~~-n-rnvc=rr.~l t m i i b ~tbf tree$ l ~ a da made of mlclrc (Clisptcr 1 1 ) . JL ic ~ h rc5t , 1 1 1 t I 07 r:.rlil mal's cuticle. Prrniln.li~lyt r i l n b ~ t u\Ircrc. uiinblc r p.11~\pCck.lrd c ~ ~ l uh,irrilc~iiizii~q ~ir 3 ~ 1 t thc h crlvirwsecrete ni.~t~ri+il orElcr than cnlcirc for any part r ~ f t l t s I * t r I I X ~ I C A I 1 t w r t 1114 T ~ I I c Z Ctllr~li ~ ~ ~ #I I I C O ~ I S ~ I C ~ I When O~~~. rxoqkclernii d u c to inh~-rcnt~ c r ~ c t Ii icn i ~ t . ~ t i r,lrl ~ i ~ ~7-1 , l , l [ l ' [ < I <' rccclrclk lvcrr tint kept 11) the carlv 15(51k t h c ~ c I ,~~II'~LLII although calcite hac the urrtur o f tr.~nsparciicy, I* riiotl~\1 1 l . d LIP ~ ' E ' J i ' r ~ O~'TEIC' p ~ l p l t I , ~ t ~die o l ~other : I1'L <+ I 1 1 1 1 \ ,!~\'C 1-iokcd our bv ~ r ~ - d ~ t Hut o n . the end o f thc ccrl- h i e c h ~ l g r tn niiiicr.il. psnducl ng douhlc i i i l ~ y \ L I I ~ Iti11111 7 difr't.rcnr depths, f i r 1fgl:httrnvc-llitlg t l ~ r n ~iti rn ~ l211, ~ tun. thr. wc~odli r ~ t i *nr.ir tire citrcc hnd brcorne pol( I I I ~ of Kt= Iutccl .lnJ thr. I~l.Ocll~, W J I ~ C I I 2rr w r y scnvtlvr to other dlrcction\ than thc t n u ~ r r ~ l oig; ~r l <-,?\I t r g x t l i kji1d1 u r ~ c l r ~ Jtr. i n I>.III I l c c . ~ i krllcd, cx7miny thc b1nckent.d Trilnhitc~ha1.c ovcrcnrnc thiq di\advnri t;yc fint 11 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. ~ - . ~ snin\n ~ ~ nt ol tlie vi.;u.ll susf~u r ) ) l ~Irclll3r I,\L,Il \I of t h t11c.w JlC'Jr WLTC t J 1 ~ 1 1 t l ~ cdirk-cnlni~rcd tbr~iis,thc and cccoiid h y r n a t l y rngctlirj~l\rnuc!ifir~tic~rl~ crysr:~lI,~tnc-c;thcv II.IVC JIW ~v01vc'd ICII\ dr>uljlr.. l l ~ ~ ~ l t ! r r p l r v.~rir-tie<, ~ ~ ~ c t ~ ~for i i t110 ~ nmore g than l"0. i \ t r.!~mad ~ n dgcr~cml rcnr~r. two kindc of ,tdapt~: t I r i l l ~ c , ~ n ~pc'c~att"ni~. ~ i w n l v i r ~:r gpcnrt-r variety o t ' g c ~ i cr-ton~ ~tratc+c-il ~ a v cbucn Jirtm~q~isl~erl. 'I'11~yarc d ~ t l k ~ l ~ ~ I . l t i . < !KC1 trollrri~:ditTcrcnt I-11nr.lctcrr. ~ ~ 3g l l l l l l f ~ ~ l l t ~ , l n l A rnnrc rcccnt cu.uilplc c-nnccnir chc r;dnIc. cnt WRY< 0 f F U N 1 V I I I ~d11d r e p i - o d ~ ~ f iOr1 I < , I t 1.t h,lt Gn1apnyo:nc f i r t c l ~ ~( C ~ ,; + ' ~ V ~ Zrtullolv~rlg.I 3c.ttcrc. drnuglit 11-1 1077. rhc av,~ilnhilicv uffqpring arc prudu~cdrapidlv. Ir~dtvldt~nl'i I ) N A t!I,l1 o f ~nlnll5ccil.i hccatnr. ~ ~ l i i crctlirrvcl. k Biyycr rccdq 5hort-livcd. rclyiilg upnn tllcit supenor reproiiul I .I I ~ I ~ C h: ~~ tlve nhilitics for spccles F I I W I V ; ~ ~ ' . K s t ~ ; ~ t r ~ p i O' ~ t ~ ' I~ou~c\.cr, wrrv irir,rrb : ~ h u n J ~ t ~2nd t . t l l o ~1;aryt.r hdjrlrc r l < I I bird? 1~1ths r r c r n ~ ~b~-.lk~ -r t h t r,71iId CI)PC u ~ l t htlltlfe thc other harld, produl-e fewer .off L l t i l r i ~lllllllpl~c ?car t o y ~ . i r~ t t dthElt~IICTL- 1\ 11t1 T ~ . J S O Twh!~ I ~ ~ n a l l - rpacc- T1-~iri s an a t t n c r ~ v econcept 2nd in con! rcsprcts still ~ ~ t e f ~Iti l1195. . hn~vc-\,ct.hccn hra~ll. 4 >plr\ o f t ~cnlr-cvc~lutinn\ l ~ o u l i lt l t b t bc rcvcrrll~lc. 111 tflc cu.implc. ~ ~ u t t t csbovcd thr \clectivr ~ d v a r l - criticized. s111ccso111c Fpecles have both high fccr~n I !l%I'. r<>171 tngc tni rhc grhnci\ clcar. but i n mnnv c,lrer. ~t 15 lefs dity nlld s~rpcriorcompct~nvrrlcw,;tud .ill por~rbl l i ~ < ~ 1 1 ? 1 , - ,(3 t*c\ 1)P c.l\v tr>uiidr.r\~;lr~d thc rclcctivc ,idv;l~~t.lyt'r nf pnrtic- conlb~nntionr; of tlle~c C I I C I - ~ O H I ~ F 1113)' nc( \I i i.rluiirl.~tit (Stcan~s,1977). Furrl-rer~nore.tlic coliccpl of rli iller c11.ir~ctcn. 1 1 11\ I It.,lt Scltction, thrn, i s srcrl .Is 3 pnman7 ~ g c ~irli trvu- 'Iluuted enerp' Is~ldpct'1s not lnvnnahlv vnlid. It ' I ! , ( ) t i < * Ill ~ L E ~ I OIt I I doc\ i11)t. 111nvrver, p r o d ~ ~ cidcal c Kcno- thcrrtbre not very r.lsy ro catcgorizc adnprivc n i ~~t T I I , I. I I - I ~ W I ~ccuirccq.
1 1 6 4
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Darwin, the species and natural selection 'chiclcic-d*compoticmt. It iiiny cvrn. nftcr ii~i~c-11r v o Il~rlonaryrl~vc.rgcticc,tilrii out to hc ~ ~ s c fsu i ~some l :ral gr~rcticc makc\ riw of htccdrilr: rspencatr-prv, to ,lrl c \ ~ c i ~ t ~di ,ei~l c c n d a n rnrg=~;;iI-nedin t h r t n ~ ~thc.~r h ~.tK.ct<.Molc~-~tl;tr ycrletrrs. u11 ~ i i u l t r ~ c t ihlriilicq c mav, in \VJYP not .IF vct firll\~ 1thl.r h.~rlti.conccrbtr.itrc 1tl~o11tlir ~ J C I I T C the undcrhtoud, I3c ncccFr,Iry fifrlr t h ~inniiitcn.tiicc ' 13fthc , ~ n t lulw)i~l i o ~ vgcrw-~~tlc 111,1tcnalI \ 5tmcturcd complcx ftrnctit>ns a i d intcmr.trc\ri- c l f rhc cell., nf ir+1ni7c.c! N r w t h a t trn\.c r n ~ c r r drhe h a w < rjwtaru;ln\. I t had 11rdcud I>ccti \uyycstc=d rh.~ttlic I \ ~ I L~.voIzrric~n ..I~ tl~con.. i r wnuld scmm apprn- Cntnhnnn 'rxplos~on' o f rncLa;.o,Ill 11fc i 1 1 . i ~ IIC' :tn ctrns~clrrtlic ~ ~ n l m r r ~ I C ~~ VI C C O Y C ~ I C111I rr'Idt~11to tlir ~iiitialcrncrgciicc 4 1 t - quch i ~ i t i l t i ~ t . r i ~ .I ~ ~ l ,gcr~cttc.~. ?r W ~ I C T ~tl~rw , I S arc rcvnlut~oai- Li~~~ilics'. W l i c t ~tnnrc i q klio\\+t)of their iiirl~tiot7c.i r , tC1c rht.m~~cal pntli\\b.lvr tl7.1t Ic.irl h n ~ nqct,cf t r ~ sri,~v pcrlrap~allo\v :h fustlicr ricvcli)pn>cnt of thi\ 11; il.loyrri, c ) r g n t ~ l ~ t ~ , ~t cr 5lt1ll c F,~r t c of ~ n~ c l ~ ccot~trol t ~ c ;~trJ~-cr:rilnti<~i~. ul11.(,lnil IiCr~rcthr. nl+r)+tt~:~clc) Ic\*cl. ' ~ I I c ~ c IlCl\' CDIlCCPrS l l d ~I c~ ~I C ) ~ \ V C ) P O F \ I ~ I C ' c L ~ ~ \ v I n t 11~1% :11r~..1(4v l ~ r v i i c ~ t a ~ ~ l ~ 15~ l i challt-ryec cd tn 11co-13nnvi1i1.;ni.Tlic lii.;t n Killiur.i'< >I7ui! lctr zintIcr;t.ir~~i~r.~ J~ithcrrtrclu\ir.c a5pcctq (1 ')7C))'ncutl-rh rhcory of e ~ , o l ~ l r i ~TIII~ l i ~ ' .p r n p i ) ~ ~ . i I ~ ~ ~ l i tllcurv, ~ t ~ ) n1;)r i t illrltvr r h , ~ trhc f:cc~~omt.t h ~ pt o p u l ~ t l o narc ~ irluclr niorc dl\ clxc t h ~ \~'Oiilrl r~ irrllrtir .~pl.mr.~tu+) i~ .I t ~ ~ i t - l1 i1 1 1 1 ~ t l~~ i ~nild d hc euprcted from natural ~electie)!~. T h c lc\.cl cnt' ~ I I +I YI +~~.< T I t11.1n f i v ~ n ~ r ,l \t i p p ~ ~ lvi ~ It l, ~dl(vnn.ltinli slio~ildbc much lower I ~ ~ T , ~'ucrf~il' L S W diKI I o I ~ ~ I L - I L .c~rnpkc.urt\ ( c . ~ . t'crcncrc chould qprrad rIrroug1-1tlit. p c ~ p ~ ~ l , ~ t;rnJ ion. lplll~rt ! #I/.,l L ) S 2 ) . 'hnnnh11' v.lt-~ants~ h n u l i lhc cliini~~.rred.5111icthcrc ,rI~i!,+ tllv tntlst iinport.~ttt ~IIC'IC O ~ ~ C U ~ V ; I ~ iIc ~~IrIt ~l ~ n l l yhrgl~d e ~ r c - rof cvidcirt ~ . n r i , ~ h ~ l ui t ~v ,w t I 11 I I I ; ~ I I Y C L ~ ~ A W O ~c,~n-v CE riit~rh morc 13Nh o i i r r G n n t l ~ ~ nrog do w ~ t hnaturdl qr1t.ctro11.[t .inccr, ! zhri nrrd tbt C O ~ ~ pror(~i~i\. I T ~ TIIIC~ ' s i ~ t.Iilrlt t9f crrccrs appcrir if hv rhanrr alont. T h l I~F 3 d ~ h ~ t d b 1ptl111t. .e l ,111dthe uze or cuiiiplcsrty ( ~ f c l l nri~rti~~l; c ca1.1- but cotivcscly it could he a r p c d thnr even ifvthn1 ~krrdnli p r i n i t ~ v c l t \ l w ~ C ~ I I I L -m ~ ow rnniiy differences arc ~lcutml.rlioqe ieiv tll,~tdu curl1tl:ln 111o\tn1~irnni.13~ 5i1cli tlncwcr~tralgc.nc,trc fcr n fclective advnntnjic arc smilable to ljt' ~ - I ~ u ~ r l ~ l c r i n 11 11;15 brcn c:illcd ' r c ~ l ~ ~ t ~t'c iur , i n)unkS DNA. hv tlic redonbra blc pow-cr of n;atucll k r l r c t ~ o n . o r ~ rcinsistr t of rnattv rcpctitiolis crf thc ~ a m c Ant>tlrcr c h a l l c n ~ c rlenviilg , Crmn the cur~crptn E tult nwht ~ t ' t i i rcxccq\ 1JN.4 r \ . ~ < t s ,is multi- multigcne E7;Trnrlles I S Dovcr'r (li)H3) tl~enwnf ii~olcr ~ l i ~ ~ l i t1.e. . < , F v u y r at' rcpcatcd cctlurrlcrs or cular drive. It is hnrrd on tJ;c undentailtiin~th,it I ~ I I ~ C cuprrs ol' chc w n c gc-rlc. Suc.11 mult~plc indivichlal membrr+ of a mu1tigcnc E~~iiilv sl>ow. y - r ~ t :-n r ' ~ ~ o n - r o rsrcliirllccs lr~~~ cvict too. 311d 111111~jrnilnrity within a spccles. Thc iiidrvrdu,il ~ n t l n b r n rlll>:t~of JU ~SPCC OF ~ C I WC C ~ ~ ~ I UCJ11 I I ~I)c . C Cn f n species do nor cr~nlve~ n d e p ~ r ~ d r n but t l y drc 1t,.,11 itr qulrc ~ ~ ~ ~ cx~ri,bblr. lclv 7uYrl1at doc.< thiq 1;1rgclvIiurno~cnizcd In n ph+nnnienui~krlon-n a5 IjNA, which ~t tint \igllt ~ p p c ~ton hc 'concerted evolution'. Accordtnq tu Uovrr t t n ~1s , I > . l l r ~ t .rrr~i,iIly . dn? Sor~lcf i ~ ~ ~ c ,ire t ~ trcbnucrn~t~\ due r o fiua tlon of n~~~rnciorlq wi t l i ~ a a~ popu1.i rmn ( 1 4 1 1 . .4 n~iiltlsrnt-hlrl~l\., for rsaniplc. cdn t,cc.~~~
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38
Evolution and the Fossil record
\pcclEll In( ch;~rrcttbn<- I r I tlllt~~TZSI.CL~CC i t 1 f i e q t i e r ~ ~dvu e to j i ~ ~ rq w r x e in wclurnct.. Fmt. maternal genes crtabpmucw-c. I T ~ ~ ~ ~ ~ ~ ~ ~ I .~ ~L i- ~nilt A d~ I to D T,lllhr I , eEect4 h5h the antero-posrenor ,)lid d u r < ~ 1 - ~ r ~ 1XYC, tr~il o f mirur,~l~rlct.~iniiMolel-ul.ir clnvr c;ln thlis fix J the egg, before ttrtiIizntto11. Scc~md,segmentatinn s.in he esr.1 v s r i ~ n tit1 .I tnult~yrtlcI f > t r ~ l l v,tiid , cuc.li tisrd varintrts. genes come illto opcrariot~aftr-r thc c ~ 15? Kprtiliz~d. ul~dcrctcloc c t ) n ~ t ~ r ~tt~l vl n l ~ .nrrcr i l In I. breed in^ p~up111~t1on. give a i d tiley define the numbrr drlrl pocltlon of t h ~ ~' I C Y T.ltld ni,ly it~glc'cilI ~OIIC c mc.lnf of o f a segolrnt. 1.c. lio\i~it I + ~011strii~ ted .111d \vl~,ltir frr>111tlic 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 u t11'rr-~l;itorv pc gencq rhnt We linvc alreadv s r r n that palaco~rtology.firith I I 1~1vrrlitfcrrilt f~lr~c-trans r l ~ ~ r i ncym h r y o ~ c n c x ~2nd c unique persprct~vr af gcolugcnl tlrne. wn pi1
X I I C ~ h~~n7otit.il
17'
+4
111
Fossil record and modes af evolution
39
the rl.iturc oK cvnlunnnsn. ar~inlaland its ct~r'lronmcnt,arc pl>cr;rllv ,~ttrihutcd III partic~r1.1r. drqccrlr c)t' e\bi)li.ir?l: Itiic.yc5 to lrilcrocvolutivn. Althr~uglithc ycnc-tlu ' ~ n dl w l n t be rrt.~hl~~hcd and t h c I ~ i ~ t l of v v l i thro11qI1 ~ ~ I I J I P itit: proccsse? bv wf~icliqpccler d1I1;'rciitiatc art' well cnrood. -l-li lr I:, 311 Iliipnrrant c : ~ kt;)r ~.IIDCOIISOIcit~dcrstond, it is a n l r : rompnr~ti\-clurcccnrly that , ,\nd .it p t t ~ c n both t pllylctic c\,ulut~on,ind cnmfill1 confim3nrior1 of how tliey 1 3 3 ~dct~1:111y ~ npcr::111ri cvnlunon art. rccclvins nluclh .itScrltlrrn. ~ t c dt h r o ~ ~ gtlnie li h a $ bccn clrarly cccn Sirnprnn Of crliinl Ilnpolr.incc i- thc pcrcrption. glcnl~rcl :1',44, 1953). working ~rnainly \v?th vcrtcbratrr. ,-1111 tllc ~ C F I I rrc0rt1 .I~IIIIC'.t l i , ~ tratc'r cvoltlt~on 11nkc.d gcnctic- rhcorv a n d p,llncontnIoyc'al d ~ t ain n ,<'etaricrl tl~rnuqhtirnc 111 rolne rnrt;lrlccq rhnllgc tlliicly rnnnncs. a$ dlcl Cartrr (1954). I t carrlr tn h r 1 1 hccn vcnr r;rp~rl,hut 111 nt1ii.r caws t11c.r~-Iln* ~ ~ i ~ d c r ~ f,ito nt l di r q t i t r ~ l *1l1.1t spcrtr.; could anus \.cn. l~trlr e v u l u i ~ c ~ r ~ mnditicatii311 .~n. nt all, cithcr by litienge spl~rting(5pc:-int~on)~ 7 I)\' r rr.111dor>rlovcr long prn.l~ld~ o t rilnc. I t wrmr to I7r rhc ltlntron of thc ~vholc pnptilnrlm~ ( a ~ t n ~ c r l c ~ f l q ) . :Int L T ~ E I ~ ~ I D I I , I W C ] I ; I I I ~ L . 111 1 1 1 ~ 1 1 1~~ I I C ~ . I I I ( - C ~Unt'ortur~atcTv. howcvcr, tht-rc wrrc \*c.ry few < nut prclirrd . ~ ; tl rcg~il;frpnrc hist In f tr d i ~ d Instance\ whrrc fixst1 populations were ncttlnllv \ccn t 5on1cririic\ wc ran di.;ccn~~ . ~ pIiI d I I ~ I Cof CVOto chow 3 pndtial ~ n o d i f i c n t i ~ ntlrrougli tirnc. ~ I I I I I111 thC t i w ~ i lrervrcl, n singlr. qtclcli, 1 1 1 ~cud~ lnclcccl, many of dw clarsic- c.l~c$which bvcrr cup14 divrnlty , ~ n d Jcsccrl~l.~r~t\ qlllc!dv bt.ct>rnc pnsed t o rxhihit quch unbmkrn 3ent.c h a v c nor It17ti-d tor 11iCIn vnnuilr 11.1h1t.rtr. \i~cIi adaptive stuocl up well to mt-~dCni~ ~ i l n l y s iS~m . p n i i .ind 'intions JTC' CIIIC t l t ' c 1 1 ~ 'I T I O S ~~ - \ * ~ d cpndt t t r r ~of~ athcn .;.~sgi~cd, a7 11ad I ) ~ n v i n ,that the saritv of ti)*~11 : u n n r ~rCvt.:llcd h!. p.il,;cntitc-rln~.A l t h c ~ z ~rhc l ~ .cp.lcclnlens rntcnntldlatt. in mo~plroro,q S).bct\~c<'ll ~ I ~ T ~ I ~ I I FI C ~ ~ ~ I C ~I CI irliainp~ttc C C rt ,till ~ V L = E;\ I L I C C C \ F ~ V I I fnrni.; w a s due tn thc ~ n ~ ' o m p l e t r r nf ic~~ ud, t1ir1~1gh rn w m r 111Ft,lIliCC t??gmieiit~rv,guidc the rock rccnrd. Tllerc i.;. hc~u,cvel-,: i n a l t e r i ~ a t i v ~ hr. counrn~ , F c l ~ o l ~ ~ t i uch.lncc. l l n n ~ A l t h c ~ ~ tlic y l ~ v1t.w. cl~unciatcdby E 1 J r t . d ~,111d ~ GC-UI~CI 111 IC171. IT ( t i t h v pattrn,\ seen miy1it Ii.tr,e bc.t.11 prlhthat the rciclrri r n n b t takcn d t CIL-C V,IILTC, ,lncI t11v rd, tliey tvnulrl he utllikrlv trl 1-w iclut~rrncrl vcr? I A Cthat ~ rhrrc arc so fi-w tmnsitioi>nl writ< 1111 c71.1~rnl~tior1~ 011 1110111:rn d111n131~ done know11 ha* .;rliiicth~nghndarr1cnt.11 tn tcll r l s ahnut Ilhccr mrioli\ I ~ C I I\\'id1 P ~ b r cvplnred 111 pcatcr the nsturc. uf ni~crnevulunnr~arv praccssrs. 111 111 the hlldlw~np tcsr Ir 1, i t n p t m , ~ nrtr t nclic . ~ t Accnrtlitlg tn E l d r ~ d ~ ,ir~d c Crru~Jd,mart p , ~ l a ~ r ~ r i outret. l~rrwrvcr, that p a l . t c . r > r ~ t n l r ~ ~ i,~~t sr ~ d r c h n ~ ~ ct hdi n k i n s hnd bccn Jorrlinatccl by the idc,~ l>t~rich %Irew t ~ r k i ~ i.iry q i ~ i t r~iitl>rrtit~ I T I ~ G 11c-v. Cthat evaIu t~oriaryc h n i ~ ? ;niuss . ~ incvi t a l ~ l yhdve bcPn . I 11ariqr\ t l ~ a t,I scntbnclst ~.:ln prtxl~rcr'rhrouyll lc yeiic pool :l~rrocvrolut~ot~rctcr$ ti) s n ~ ~ l l - \ c - . ~ lcch a n ~ e ~and hence of the ch:lr:~ctcr~ of thc er~tircp o p t ~ l ~ t ~ o u . hlrl species. and chpcrra!ly tlic rr.r11riorrr1,1tlc~1 T ~ t mFl l ~ ~ r i lti1111 ld \ilolsld ~ I ~ V C J ~large VC niitnhcn, "I Ihnr ~ p e r to ~ c,I ~I I L Y V nrtc. M a c r r ~ r v t ~ l u t ~ con r i i , ueunlly rhe rntirt anccl;trdl. populatltrrl, snd noulri klther hand, conccrn? C V ~ I L ~I I C I I I ;ll1tx-c the OSCIIT nvrrall as 3 large part of tllu ancc~tr.11spcciec , , , r lrvrl. 2 ~ 1 t hthc clrlpns o f - ~ n . ? ~J ~ TO ~ Lr I ~~ S1 1 ~ 1g(*.npapl~icalrdngc. It srcrns a rcast~t~ahlc conccpr. .td,tpr~vrr.iC)i.~ti( ~ I I C T'IICSC t l I I~ ~ ~ ( ~ l ~ ~ t l and o ~ ~I-crtnidv x r v wlnc populntlon.5 may Ilnvc chalrgrtl I:\\v111 bc C O I I F ~ I ~ C Ti Ct >~turn. In J U S ~t h i ~n x v . Yct, ar ~TC'IICILICIV r~utril.C C I I I ~ ~ I ~ L I I mi$ < C T ~ C Fof y : ~ J r dln tcnnedia rc fortnc , ~ r c FCI~OIII \t-cti in
I
