Invertebrate Paleontology and Evolution Clarkson

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Palaeontology

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Contents

XV

xvi

Port One General %laeonto!ogical Concepts1 Principles of palaeontology 1.1 I t ~ t r o d ~ ~ c t m r ~ 1 2 Ocuurrcticc nf irlvcrtcbrate fossils 111 Pharlerozoic rocks Hard-part prercrv~tion 'loft-part prcscrvarion 1.3 llivlciotls of itivcrtcbratc p ~ l x o ~ ~ t o l ~ g 'T'axonorny The spcuc5 C O ~ I C C ~ ~ Noi~lenclarurcand ~ d c r ~ t i t i c a t ~ o n s s d specie5 of h T~sonomir~ 111er;rrzhy Usr of st$1t1stac;11n e t l i o d ~ ~

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

2 Evolution and the fossil record 2.1 Introductiori 1.2 Ilanvln. t l ~ c cpecit.5 and tlatural sclcctioil Inllcritanuc a11d the qource of varintiorl W1lt.r~ drwc variation cnmc from? Sig~iifica~~ce of allclcq Mutation 'iprr'id of ~ u ~ ~ t a t i n t l f thrnugh popiilat~otls Ilutir>i~ Allupatnc rpcci,~nnii klctcrorllrnny T c ~ t ~ m~crorvolut~o~ldr)'i pntturnc ng Ar~alyndnthcr rr=fvrviiccq

8

mollusc^ r ~ t n ~ i . ~ r n c no n ly ~ l l i ~ ~ t ~ o t l tr 8.2 C:P.IWl hc.lrloll 8.3 S o n ~ c .~cpcct\ thcll t r ~ o r ~ h o 31ld~yo\%-th of lo Co~lcd 3

Ccphallc fringtc Erjrollrticnt and co~ptarlvc ~mictlircq Tllnras Pyg~Jl~l~rh Appcildagc? TrlIobltc track< anrl tra~ls Walkmg tnnvenrcntr in nrtlrrc-)pad? 13iffcrtiit kindr of tnlohirc trallf 1.1ft. attitudcc. h d b i t ~and c c n l o ~ y F,c dycl~ ,ind ontogcllv CPa+~rfi~a t~on E~olution C;elicral pattcrrl o f c v o l ~ ~ t ~ o ~ l Microcvulu tion Faunal yrovincrs Stranpaphic.~lL I ~ 1 I 4 Phvlurn Chclircr,itn Clav M c m ~ t c ~ t : ~ a t a Suhcla?~ Xtphost1r.1 Subcla5s E t ~ y p t c r i d ~ 1 1.5 L'l~ylurn Crust.xcc;i U~L~ho~rayhy Elookr, trcaciqcc dtld ryir~poria I~~divirlual pctpum .;~nd otllcr rcfcscnccs12 Exceptional faunas; ichnology 12.1 Introduction f 2.1 Dtlrgcn Shdlc f;iunLl

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

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3HX 388

392 39 7 400 400 4C)(1

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

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xiv

Contents

12.7 I c h n o l n p Clnu~ifiratinno f trace cocrils Morphologic.11 m d prcccnratlonal rlascitication 13chnvto~~r.~l cI.~c~ifirntioi~ Phylngcilcric cla5sific ~tiola Urcq of ~ c l l n o l r , g C;crlinicrit.ly ctlvironlllent Crratigrapl~y Fossil l>chav~our Fiil~lio~~pli~~ Rook . l r tl~c hrytitlil~gc>t'tlic p r o j ~ c r;IS to IIOW I T y n c , to cscape from the min. 'rhcrc. in a dusv &rr ,111Iil prr~t-chcd n-it11 thc ti-~urtlj cditiotl, and r havu casc wort two g s n t arnniniai teq (probably 7'i!r111irrs nri t l ( \ . ~ r t i~logctht,r u.itl1 IJ;~vid Harper, Pcrur had lrvud iirdtioris of y e d s J ~ O Thcv cvc~tctl hccihl~clLl~\i~. Iligly, Liz H.arpcr, Dick JctT'i.rics. r ~ a t ~ o t i S~~s;aii which has c n n t ~ n u e d ~11inE now, and I all1 still glad to cscapc to t l . 1 ~hills on a f r u h sutrlrncr mom]l L g t - m ~ : ( ' ( ~ 1 ~ A l ~ t l t . h ~ * ~ r h ~ ~ i ~ I Oorhcn Ilavc . L ~ F ; I ~ i~ I ~ o back I!I,\~L! 111i- ,9r ,111 ~ ~ likc. .lhovC 311, r o rccord r r y hc;jstfclt coir~ctl~ing the ~ ~ C I I T I C S S A I I ~ C I I V C I T I ~ of a~jcierit lrl scirntific ri~.~:~h< trut rid st:ll\~nrrfricnd Ct-cilia Tnylor, lifc, a t ~ do f the rncthods avallablc for ~ t s r r > Irly 0 ,111 tlw \ u p ~ ~ o.und l~clpfl~l rt ~uq;crtinttr ~ h c 112s study. If ro, 1 will havc achlcvcd \\,hnt 1 srt orit 2 do. ,? . L h t ~

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

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P R ONE AT

Concepts

111

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

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of nur u~lrlerstandl~lg t l ~ c11istc)i-y o f lifr-, of pcrl1.1p~ rno5t significant of all C V C I I ~ F ID(>^ ~ I : I C C tlic about 343 Ma ago nt about the Gcprlnlliq ot' t h c ~ ~ I I I .(l l ~ l l t )~i11111ot1 , vc.lrc . i g ~h c Earth carrlr inti) C:nmbrlan Pcriod, For nt tlliq sta.yc tlrcrc \\:IF A q ~ c i t .II~:,yrrllh~lllt fi>rriijnqf r r ~ i .I lionrlcnr~ng ~ dlrc of den pmllfcration oidifl?sent kiilds nf lllarlttc ii~\vcrsl I I L lr.,. ~ l l t r t,iiid g;i\. I 111~11 t;lteci rouild \ t r b m t c ~ . 13un11g rh~scritical period r l ~ cpriiicipnl I, 51r1i 1 , I r y r p~rt~c.lcleof P!lanrrozo~c tltrle. nrc dictinpirchccl 1s). .I \.cry 1rh111tlic t i n t 30";I O F tlw ~ J , I I T C ~ * F l~i~ltm-y, Tl~ert. r i c h l ~ c r i t a ~ c the fossil rcunmnq of thu invcstcof c r ~ * r i ~ , l l{n>\ < \ I T T I ~ ~ O .~ ~ I I I \ I I I( ~ c t c rIjI ~ ~ L , I L 'h111rE~rntef that rvolvcd thro~~g!r succcssivc Ii~qtnncnl ,r~.:.tl ,rlc,rc', or L-ynt1ol7acrcn.1)t ? rock? 341 It 1 ,Ma pcnndq; their ~ t l i d vI$ the dornail~of in\rcrtchr;~rc i .1 ,rl l~ir. prcwnl,~l~lv onpnntrd hciorc cl1t.n. pal.wnntnlo~?; nlid rhe qul?iect of thi\ book. Tt.c\: rrnit~lc form< r, !iir hcc111 ~o hnvc dom~rl.~ted f itcrlth c , thc 11c~ut2ll01) Mn. nnd evolut~nll Lr t h . ~ t tllrlt, \v,~c vcn slozv. Nr\~enlieless, tllc rr\cib,ictc.n,~ atld phato~y~ltlrcric , ~ c t e r ~ a b werc I [ ,r~u!rlcr~tnl ch.111y111~ 111 thc cnvirniitllcllt. fur they #:I: jltf LK!grn i i ~ t o .Itrnosphcrc t11.1t war previ,irl LI~vtjiil ~ t \ L I dint nr~imallife eventually , IIIIC y~lv.~hl~. hlly \vll~.~r qnlilc. of thc cnrly Irvitly lbeaiirl,ln,cl~,ingc,.~ct-clcratrd, and with it all lirncr~e\. FIIJICS., ~ it~rc)nern r ~ d~-nucf~tot~cs o n l but 11 I I I I ~ L ' ~ . of IIC'W ~ c ) ~ + I \ > II ~ 'I~~~I C F I upened up to rlic WIIOIC nrr not cor.larno11 m s a i i d ~ t a ~ ~ e s . c ivlv:~~. Thi5 \v.ls n n t 1ir1t11coiiip~rativclv ~ t e Scdimcnt.iry ironcrorlc< iilny t~avc rich fo\sil l~ir. l 11 :~rl~1~~gc,~l t i d tl~crt* hissnn. a ,we 110 tbrcil ~nltnals rern;~ins. Occnsionally they nre f i ~ ~ m dFonw conme 111 IIO>L\III L ~ Y ~ L ~ I I I I P t)ldcr t11aii s h o ~ ~ t MA. mcks F U C ~ F L~cy%vnc-ker I I>~\ 7tbO A nr~dcvcn coirglo1nerntr5. A:wn tc>g~-thcr c!~eiilical 111~11tr1t1 t l ~ c t;nun,r tvr-ll. 'l'llr.\c ~ r rrninc.n~cly ? cig- halamcc i r uilrtnblc. T h r \cdirncrrt 1i1111 bc in r t w l n ~ cal cquil~briuni(>lily.iftcr dlngcnctic pl1!.cict7clrciilii ~ i f t u . i r ~itb r pn l,~contc\lr*,q. Il.~vc y T h c olclcior.~c i ~ , , ~ i -fti i l r ~ itrncc inqctlc) nsc pre~cn,cd. t l i e ~ c ( arc clcccribcrl in ccctirrn and thC h i v l , ~ c s r r ~ ~ b l .prcrciit in t h t rnck nlny tht* rturly of cucli ~-rlould\ yc< If llavc tlccrl ~ T J I I F ~ O I . ~ C C C ~ I I ~C ! I E ~ J I ) L - C . nbradcd. dnm- 7.2. with rcGrct~cc rt> h r ~ ~ l ~ i c ~ p r > c l \.I. fowl LOI cncloccs ,111 c ~ r i ~ i n . ~ 11ollou~FFBCL-, ,i'r For il~sc.~ncr lIy a ~ r and rnl1xt.d \I it11 eleinellcc nf n t l ~ c r d fizln~r. Even (hit k-~hcI1c.d ,111j1n.1litycrc ~ r i ~ n , ~preqeiit 111 .I hchvccn the p,Iir nf khclls tlf .I bivalve ur br,~tlilnll\ pod. tliir qpnrr may citlicr hc lcfi rrr~pr).or Lwcomc f>und, t1lc.y rnay not br p r c ~ r ~ \ ' r 111 C . I I I L ~ ~ ~ 6 1 d: ~ I nle11t- I T ] \ V ~ I I L -t~l r ~ - ~ ~ - c ~ i lv ~ i t c r ~ ~ . t i n ~ rtci~li~-, tjllcd w r h ~tclvnicnt.In thC 1.1twr C R C C . ~ ~ d i l ~ l c ~ i t re for core t