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THE EVOLUTION of the CEPHALOPODA Bv L

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THE EVOLUTION of the CEPHALOPODA Bv L THE EVOLUTIONOF THE CEPHALOPODA BY L. F. SPATH,D.Sc. [Rennrr.rruoeno*r BIOLOGICAL REVIEWS, Vot. VIII, No.4, Octonr.n19331 - CAMBRIDGE AT THE UNIVERSITYPRESS. THE EVOLUTION OF THE CEPHALOPODA Bv L. F. SPATH, D.Sc. (ReceiaedFebruary zz, 1933.) (With Thirteen Text-figures.) CONTENTS. PAGE I. Introduction +r8 II. Distinctions between Ammonoidea and Nautiloidea +2r (r) The initial chambers +2r (z) The coiling 42+ (3) The siphuncle 430 (4) Other characters 439 III. The ammonoid ancestor 4+2 IV. " Bactrites " and recapitulation 445 V. The primitive Cephalopoda 450 vI. The supposed cephalopod Volborthella 4s3 VII. The phylogeny of the Cephalopoda 455 VIII. Thb Dibranchiata 457 IX. bummary 459 References 46o I. INTRODUCTION. Tun development of the ammonites cannot be discussed independently of the evolution of their Palaeozoic forerunners, the goniatites, and the earlier types of these, again, are very intimately allied to the ancestral nautiloids. Although some- what diffident of casting so wide a net, I feel that the problem of the evolution of the Cephalopoda must be restated. In the past, what in my opinion are erroneous views of the development of the Ammonoidea have been popularisedlargelythrough the enthusiasm of workers who were attracted to the subject but had little practical experience of ammonites, and who too often were inclined to treat these fossils not as the remains of natural organisms but as material for speculation. This article, however, not only criticises the prevalent view that the ammonites, like the ancestral goniatites and Clymenia, arose from a straight nautiloid (Bactrites) and then hurried through the early stages of coiling before uncoiling again towards the end of their career; it also discusses various views put forward by workers on the primitive nautiloids. Yet with the purpose of making the article intelligible not only to the specialist but also to those who have only a slight acquaintance with the subject, I have attempted to illustrate in the text-figureS at least the more important features of the different fossil Cephalopoda referred to in the following pages. snorres n fq peqsrlqnd eg ol crlselueJ oo] tou sB/K slloqs perrnbce ,tltuapuedap -ur pue ,{lpaleodarr{crr.l^\ spodolegdec ssel-llerls Jo uolssecrnsIEnUI}uoc e Jo uol}ou 'passncslp (9z6r) s(ualuur?rqcg uale ag o1 lecrtaqtod.,tqoo1 ag otreuos ,tq pareprs -uor ueeq aABq',ro1eq o1 parraJe: 'sprueurdlcaqr uo s^tell (fr6r) s(.^A.eloqosalI{AA 'ller{s 'putur aql Surueqt8ua:ls;o .,{e.,vrlecruouoce uB .{1e:aurst 1r .{ur o1 ,,'slcnpord Ienxas snourunlo,r dlqeqord aqt eceld o] urooJ eJour Surure8 ;o dlrssacau eql 'uorldrosqe 's11eqs 'd ruo{ ^,(q aq} Jo eprsur ar{} uo paleut8uo,, uol}Elnuue slq} (ozt 'rc6r) uuztuepand ot Surproccy (;so,IarolluO,, lo sLuJoJuleuec Jo uollelnuue 'aldruexa 'se eql Jo esec oqt ur JoJ .,tren,{ew sarnlea; eldurrs ,{1}ua.redde5o uot}e}ard -Jelur eql pu€ lsregtro pue'npaqlnqloA Jo arnleu eHl ,,'sap4tng,, Jo qJuocolo;d '1sr8o1o1uoae1ed er{} are qcns I qooq-lxet eq} .{q Sur8pnf le.reue8eq} o} pel}}asreadde '8uture.t .,reu qcrq.n sturod uo uorurdo5o acua8relrp l{cn{u lills sI aJeqJ . srqt lseSSns leql euole lerrateur er{t Jo 8ur1ep InJtqnop pue arn}eu 1ca;redrurer{} }ou sI }f }ng 'suorlcssurq] 'xuletu ur arnlcnrls leurelur IBaAeror IIeJ dlquqord plno.,ra er{} Jo d1rur1p1s.{rceql 'suerurcads'peqsn-rcun 'a'!'pa.trase-rd 'pecgIJcBS Jo ]unocce uo dlqe-rnoleJlsoru oql 'sJalceJer{c Jr uelg snorlqo ssel aq} go uorle8r}salul palle}ap erour ro3 gSnoue luepunqs Jo elqBlrns sr teql IBrrelB(uou ]sourlB sr eJal{l tBq} PUB-sEunEJ e}IuoururB elrssecrns uaeunoJ slr qlra-porrad }{nep ar{} Jo eloq,a oql ueq} uol}Ernp -ra8uol dlqeredruocur Jo serA-souoz el:rtaruo81n;rqnop o.{a}'t{lr1y\-ueruola0 Je,^.oT sII{} '8ur11eddu'paaput leql repeer 1ecr8o1oa3-uoueql uodn ssardurro1 d:essacousr 1I 'sl 'pap;ocer e8peyvrou{ }cexe Jo {JBI ;no ueeq e^€q sprouourtue lprg eI{} acuarl.^a uerrnlrs reddq eql Jo do1 eqr ro uzruoleq re,{aoTaqr Jo eseqaqt o} u,^aopouroc e.^a uerll6 'sruJoJpolceuuocsrp pue pelelosr uo esle .ro sotcedsu^\ou{ dlereldruocul uo 'Sutsodurt peseq ,{1a8re1ere ,(ag} sesodrnd lecrqde-r8r}€rtsroJ }uellacXe roAo,t\oqtnq eq detu seuoz elrleruo8 snoJeJruoqJ"J Jo suozrJor{ a}ruoururz JrssBrJI Jo Jaqrunu 'suorsseccns (s{ror oqJ paldnr-ralurun pug o1 souroreq U llncsrp erour eq] eI{} 'seuneJ raplo er{J crssernf e8pal.u.ou{rno uI }sIXalprs sde8 }ea-r8}BI{t reerl ol ".ll;o pesud-lns eq deru euros q8noqr 'urvrou>1dlq8no-ror{} ssel re; dlureual sI crssernf aqa 'seune; elruourure olrsseccns dueur os Jo uorlducsap eq] roJ serllunuoddo lenba pro$e lErraleur leco,rrnbeun puB pooS Jo esuePunqB PuE dqderSrlerls Paugar pu? reelJ qclqa ur uorleurroJ snoocelarC ror{}o due sr areql rer{laq.l.rInJtqnop sr }r te^ 'pallos ag of urzurer llrls suelqo.rd .(ueru ,lroq SursrlBeJure I pue uorlerederd ur sr 'sdr1 qderSouour ar{l;o ged qlue^ela eq} ,^aol{ ,sre8ug ,tru }e slrssoJeseqt ;o .,iro1s 'eldurrs eql e^Er{ o1 ;1es.(u pe^orleq I pue .(la.Lrleredurocelord plnorn >lw} eq} }eq} 'rynog luapuuoc sei\rI ary {o oaprouoluuv aLft{o qdorSouow,(ur ;o ged 1s-rgoq} otor.{a to8e tueq11 l s:ee,(ua1 ,,'ctsszrnf,, ol ,,snoec?traJJ,,uro{,,teuecol6,, o} ((aueJ '.,tes 'e}ErqaueA -olld,, tuo.r; sdunf Jrer{}Jo epn}ru8eu agt esrleerslsrSolooz ,Lrag 'slerurue -ur Jo o]BJqeuol Jo suolle1nur elrssocons Sunrasqo Jo esee eql Jo 4zeds 'serJn}uec or{/'t esoq} Jo rusrurrtdo epceg aq} eJ€r{s lou op I uaou{un }uase;der (sao8 ol elq€{ sr le^relur paprocerun dre,ra eJour eql uerro}srq lecrSoloa8 eq} lJeq JoqunJ er{J'relncr}red ur slrssoJrno Jo 8ur1epeql pue 1e-raue8ur aJuorrs lecrSoloa8 'd'p Jo ernl€u a,rrlelncadseql Sururecuoc (toL tt6r) oreq.&\eslepres a^eq J ]Br{1v\ ol acuereJar e ,(q olcruB er{} ecqerd ol Pe^\ollB aq .,teurJ uosBer ourBseq} roc 6* apodopqda3ary {o uolru1oaaaqJ t#/wuu),Q/ 420 L. F. Sparn scientific society. Many observers, bewildered by the seemingly endless r-aria- bility of one small groupr missed the essential uniformity of the ammonoid (or, indeed, cephalopod) stock as a whole. But these examplesmay at least partly explain why the problem of the origin of the Ammonoidea and their relationship to the ancestral nautiloids is still debated. My own views outlined in ry23 (p. 6S), are more or less diametricallr- opposed to Hyatt's Q867-rgo3); and it is attempted in the present article to shou' that some of the opinions expressedby Hyatt fifty years ago and still adopted by certain *'orkers were based on mere conjecture. Thus it is still commonly asserted that (r) the recent IYautiluscan be traced back in an evolutionary seriesthrough spiral gyrocones (Fig. 5 c) and curved cyrtocones (Fig. 5 b) to the straight Orthoceras(trig.5 a). This series is held (z) to be characterised by the absenceof a calcareousprotoconch. Then it is stated (:) that an Orthocera.sin which the (supposititious) protoconch, i.e. initial chamber, had become calcified (instead of being shed) gave rise to a straight ammonoid shell (" Bactrites") with ventral siphuncle. Next (4) this is said to have produced the loosely coiled " Mimoceras" and the involute goniatites which in turn gave rise to the ammonites. Assertion (r ) was a fallacy even in Hyatt's time ; lie himself statedthat there were no such serial relations in time. Assertion (z) was based on an assumption, dis- credited many years ago and now known to be quite untenable.With regard to (3), it can be demonstratedthat every rich Orthocerasfauna (and sometimesa single species) has some forms with marginal siphuncle. (+) " Bactrites" (recteLobohacttites) is a secondarily uncoiled goniatite becauseonly series going from curved or coiled to straight can be demonstrated, never from straight to coiled. When previously ridiculing the notion of a straight Orthoceros stage being " omitted " or " skipped " in IVautilus and the }lesozoic ammonites, I stated my conviction that the first goniatite, namely the Silurian Agoniutites, u'as nrerelv a slightly modified I'{autilus (e.g. Barrantleocera.i),but my immediate concern was a disavowal of the usual palingenetic methods. Since then I have been prornpted to discreditHyatt's " laws" more vigorously,merely as a result of practicalexperience. T'hose who work on other groups may at least be interested to hear s'hat constant handling of cephalopodsfor many years has-rightly or wrongly-suggestecl to nre, as opposedto the theorising of amateursat a distance. If it be objected that special- ists like the late Prof. J. Perrin Smith (tgSr) came to different conclusions, I can only reply that in my opinion he constantly overlooked the fact that by hereclitv an ammonite was an ammonite, and that like other organismsit had to grorv and thcre- fore necessarilyhad to pass through more primitive stages(see Spath, r933 6). The principal conclusions arrived at in the present enquiry are summarised in the final section. Their briefness may invite categoricaldenials, but rvhat is u-anted is patient sifting of better evidence, if such be available. Much of the material for this article has been taken from my Catalogue of the Triassic Cephalopoda in the British Museum, now in preparation, and I must expressmy grateful acknon'ledg- ments to the Keeper of the Geology Department, Dr W. D. Lang, not only for allowing me to make use of this unpublished information, but also for enabling me ,.',{Jessecau,,dllecluerlceur oq 'palcage lou ,(eur suorluJ5rpotu eql q8noqrle lou sI luarunS:e uleur s(JIo./y\ePuIqcS ler{} pu" olqerre^ .{1euer1xeare sqcuocolo,rdosei{l }Et{}}Itupe III^aclng dq uerrt8 suol} 'slleqs -Erlsnllr aq] qtr^\\sa:n8g leut8uo s.e]?od
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