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THE EVOLUTION of the CEPHALOPODA Bv L

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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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  • Size Distribution of the Late Devonian Ammonoid Prolobites: Indication for Possible Mass Spawning Events

    Size Distribution of the Late Devonian Ammonoid Prolobites: Indication for Possible Mass Spawning Events

    Swiss J Geosci (2010) 103:475–494 DOI 10.1007/s00015-010-0036-y Size distribution of the Late Devonian ammonoid Prolobites: indication for possible mass spawning events Sonny Alexander Walton • Dieter Korn • Christian Klug Received: 17 December 2009 / Accepted: 18 October 2010 / Published online: 15 December 2010 Ó Swiss Geological Society 2010 Abstract Worldwide, the ammonoid genus Prolobites is Abbreviations only known from a few localities, and from these fossil SMF Senckenberg Museum, Frankfurt a. M., Germany beds almost all of the specimens are adults as shown by the MB.C. Cephalopod collection of the Museum fu¨r presence of a terminal growth stage. This is in marked Naturkunde Berlin (the collections of Franz contrast to the co-occurring ammonoid genera such as Ademmer, Werner Bottke, and Harald Simon Sporadoceras, Prionoceras, and Platyclymenia. Size dis- are incorporated here) tribution of specimens of Prolobites from three studied localities show that, unlike in the co-occurring ammonoid species, most of the material belongs to adult individuals. The morphometric analysis of Prolobites delphinus Introduction (SANDBERGER &SANDBERGER 1851) demonstrates the intra- specific variability including variants with elliptical coiling The mid-Fammenian (Late Devonian) ammonoid genus and that dimorphism is not detectable. The Prolobites Prolobites is only known from a few localities worldwide, material shows close resemblance to spawning populations mainly in the Rhenish Mountains and the South Urals. of Recent coleoids such as the squid Todarodes filippovae Where it does occur, the majority of the specimens show a ADAM 1975. Possible mass spawning events are discussed terminal growth stage suggesting that these are adults.
  • Geology of the Shepton Mallet Area (Somerset)

    Geology of the Shepton Mallet Area (Somerset)

    Geology of the Shepton Mallet area (Somerset) Integrated Geological Surveys (South) Internal Report IR/03/94 BRITISH GEOLOGICAL SURVEY INTERNAL REPORT IR/03/00 Geology of the Shepton Mallet area (Somerset) C R Bristow and D T Donovan Contributor H C Ivimey-Cook (Jurassic biostratigraphy) The National Grid and other Ordnance Survey data are used with the permission of the Controller of Her Majesty’s Stationery Office. Ordnance Survey licence number GD 272191/1999 Key words Somerset, Jurassic. Subject index Bibliographical reference BRISTOW, C R and DONOVAN, D T. 2003. Geology of the Shepton Mallet area (Somerset). British Geological Survey Internal Report, IR/03/00. 52pp. © NERC 2003 Keyworth, Nottingham British Geological Survey 2003 BRITISH GEOLOGICAL SURVEY The full range of Survey publications is available from the BGS Keyworth, Nottingham NG12 5GG Sales Desks at Nottingham and Edinburgh; see contact details 0115-936 3241 Fax 0115-936 3488 below or shop online at www.thebgs.co.uk e-mail: [email protected] The London Information Office maintains a reference collection www.bgs.ac.uk of BGS publications including maps for consultation. Shop online at: www.thebgs.co.uk The Survey publishes an annual catalogue of its maps and other publications; this catalogue is available from any of the BGS Sales Murchison House, West Mains Road, Edinburgh EH9 3LA Desks. 0131-667 1000 Fax 0131-668 2683 The British Geological Survey carries out the geological survey of e-mail: [email protected] Great Britain and Northern Ireland (the latter as an agency service for the government of Northern Ireland), and of the London Information Office at the Natural History Museum surrounding continental shelf, as well as its basic research (Earth Galleries), Exhibition Road, South Kensington, London projects.
  • Integrating Subsurface and Outcrop Data of the Middle Jurassic to Lower Cretaceous Agardhfjellet Formation in Central Spitsbergen

    Integrating Subsurface and Outcrop Data of the Middle Jurassic to Lower Cretaceous Agardhfjellet Formation in Central Spitsbergen

    NORWEGIAN JOURNAL OF GEOLOGY Vol 98 Nr. 4 https://dx.doi.org/10.17850/njg98-4-01 Integrating subsurface and outcrop data of the Middle Jurassic to Lower Cretaceous Agardhfjellet Formation in central Spitsbergen Maayke Jaqueline Koevoets1,2, Øyvind Hammer1, Snorre Olaussen2, Kim Senger2 & Morten Smelror3 1 Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway. 2 Department of Arctic Geology, University Centre in Svalbard, P.O. Box 156, 9171, Longyearbyen, Norway. 3 Geological Survey of Norway (NGU), P.O. Box 6315 Torgarden, 7491 Trondheim, Norway. E-mail corresponding author (Øyvind Hammer): [email protected] The Longyearbyen CO2 storage project drilling and coring campaign in central Spitsbergen provided new insights on the shale-dominated Middle Jurassic to Lower Cretaceous Agardhfjellet Formation, which is the onshore counterpart to the Fuglen Formation and the prolific source rocks of the Hekkingen Formation in the Barents Sea. Logs of magnetic susceptibility, organic carbon content, organic carbon isotopes and XRF geochemistry on the cores, together with wireline logs, biostratigraphy and sedimentology, have made it possible to refine the interpretation of the depositional environment and to identify transgressive-regressive (TR) sequences. Several key sequence-stratigraphic surfaces are identified and suggested to be correlative in Central Svalbard, and four of them, although not necessarily chronostratigraphic, also to surfaces in the Barents Sea. Due to the nearly flat-lying thrust faults in the upper décollement zone of the West Spitsbergen Fold and Thrust Belt, there is some concern about the lateral correlation of the sequences within Spitsbergen. However, some of the TR sequence surfaces appear to be of regional importance and are recognised both onshore Svalbard and offshore on the Barents Shelf.