Evolutionary Trends Volume I
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I Skeletal Biominerali zation: Patterns, Processes. and Evolutionary Trends Volume I Joseph G. Carter, Editor @ vAN N.STRAND Rt^lll9lf Chapter 8 cephalopod shell structure and General Mechanisms of shell Formation Klaus Bandcl Gcolosisch-Paläontologisches Institut und Muscum, Univcrsität Hamburg, Bundcssrassc 55' Di0O0 Hamburg 13, Fedcral Republic of Gcrmany (Gr€goirc, 1987) produccd by sPccialized Contents shccts and fibcn cells within thc pcrioiEacal goovc. Thc shaPc of these deoosits is detcrmined by thc muscular mantlc sdjaccnt to the L lntroduction. sräovc. Mineralized s[cll consisting of nacre and various in the fossil record. II. Preservation irismatic structurcs is added to the pariostracum by thc shell III.---' Structure of the embryonic sh€ll. 'nrantlc- lies bchind thc pcriosu'acum gland zonc' of thl organic primary shcll p 99^^ which Ä. r'rincr"tirarion Manv ceohalooods with an inGrnalized shcll also add covering thc primary shcu' p' lm B. Biomineralizatcs material of a variety of structures exterior of first scptum. p. 100 mineiraizäd shiu C. Mineralization pcriostracum using the muscular mantlc. IV. Microstructur€ ol the adult shcll. to the Ceihaloood shclls consist of thc primary shcll, which is A. Nacreous and associated sructurcs. p. 102 mineräizc{i inside the protection of the egg-capsulc, and the B. Lamello-fibrillar structure' p. 104 sccondary shell, which'is mincralizrd aftcr the primary shell C. Scotal structurcs. p' 106 The sccondary shclt is sccreted largcly after thc tube s-tructures. p. 107 has formäd. D. Siihuncular from thc cgg. Thc primary shcll is initiauy exterior to the shcll walls. snima.l hatchcs V. Mineräliz:tion attachcd to thc manüc 8nd ii beromcs mincralizcd only after function. 108 A. Rostrum P. rhc mantlc margin dcuchcs from thc shclt margin (Bandel Aulacoceratid rostra. 109 B. P. 1982). Thc sec-ondary shcll, in cons'ast' is sccrcted without rostra. 109 C. Bclcmnitc P. anachmcnt betwecn dic mandc and shcll margins. in thc scpiid cuttlebonc, ctc' D. Rostrum honrologi Durine evolution somc ceDhalopods abandoncd thc ances- o. ll2 tral miniralizcd chambcrcd shcll för a non-mincralizcd shcll, ond siphuncular VI. Shelideposits rvithin the chambers ss in modem'squids. Olhcr ccphalopods, likc Jurassic tube. Trachyreuthis (vainpyromorph octopod) (Figürc l; Bandcl A. Inu-acameral deposits. P. ll3 and cxtant sePiids continucd to p. and Gich, 1986) and Tcniary B lntrasiohuncular deDosits. l14 the cxtcrior ponion of thcir shcll. The jarvs, snd statoliths' mincralizc onlv vII.--'Ä. Niiniralized brood chämbcrs Lowcr Dcvoniän palacotcuthoirorph gcncn Boletz§a and of Nautilrr and aptychi' p' I l5 Mincratizeä iaws NaeJireurhis (Figie 2) possibly rcpresent intermcdiatc forms, B. Brood chamür of Argomuta. P. ll5 sccrcdne mincralizld shclls whcn young but later searetlng C. Statoliths. P. 115 oreanicihells onlv (Bandcl c, al. 1983). Äccordine to Bänäcl and Hcmlcbcn (1975) sccrcdon by thc shcll mantlä rcsults in 'mlc' biomincralizates such as nasre I. Inroduction in ."t ict tt c stt crural arangemcnt is vcry closcly controlled Prismatiö sructures, usually with basic carbonatc bv thc orpanism. Ccohalopod shclls arc consructcd of thc calcium in width, are also produced by thc shcll association with ii"..nrs d.2 -ic-ns at"eonitc and calcitc in closc (Bandel, contrast, biomincralizates pro- ""iräomtä their mantlc 1979). In äisänic inatcriall Likc all molluscs, ccphalopods usc to be lcss subjcct to various oüer duccd bv thc muscular mantlc appear mätlc o sccrctc thcir shcll. HowCver, bioloeicäl control, and thcv rcsemblc pattcms of inorganic jaws, statoliths,^oPqrculum hardDarts. such as thc radula' Similar pörly organizcd shcll smrctures are (aonichi) tissues othcr mineial erowth. and brood chafobers, src sccretcd by by noi'mandc cpithclia, such as thc mantlc ponions, thc somctimis also formcd iträn' Oi manttc. Tkec diffdrnt arm folds in thc casc of thc Ärg onauta shell' and thc lips in mantlc, thc periosracal gland zonc-and thc shell muscular thc case of thc Ncnrr'lus bcak mantle. are rcsoonsibli for thc sccrction of diffcrcnt pans of ttre sncit. St eti acposition begins at the mantlc cdgc-with thc in thc Fossil Rccord iormition of the p^criosuacuä, I hyer consisting of organic II. Prescwation gcncrally aragonitic composition' fossil Sk letal Bioninanlizalio : Potl.rns, Proccsras oad EtolttbnÜtTrcndt' Becausc of thcir ccphalopods 8rc commonly sltercd by a varicty ot dtagcncuc votunt t . togph G. c-t ttt, cdito,. vsn No§tnd Rcinhold' Ncw Yorl' 97 CEPHALOPODS .'.1r.. '. l-..:,:jl' ,i . ,:.-.J;.,;";,ri.'l.i , ... :' .: : , "r,i.;;;,.,:, ' ,:tn.., r,11!!rßrl3l§t,tttt!fl|!'§?L'i!r.!i+r:Jj i:: ]:1i: : 'ri... : -1";f.r: i...ij.ji.lj ilra -Ii $ ällijl ilä,ir,#ji#?#$itff ,:i:ffi 'r/#ffii"*ljTll:1Tü,:#klFiä:äffi Leich (1986, Fig.20). stäü:"t*il:1'Ti'"ä:iü,,iTHä,:}i{,"tr:d""xil ffi t3äf ,iHt'#lr;"ä,T'tdäffiiffiüäj*,"+ iäää äi,jv una", diagcnctic conditions' Nacre-can ot"^ t'a Eansformcd"cnaiir rnto calcrtc rods and nccdlcs which ä',:#i";ää;;;;ä lamellar sructurc in ürin section f, iffi **fälf*firt+lli'ft :u',rrffi raandel and Dullo, 1988' Text-fig. 9)' *"$:i:"f.{i '"fiää"iäri ääialinlsis-an*gäm"nt of alägonitic prismatic structures i*:r*ti"tllgliin'*x**xtff a.i,ää"äi"äJ' i#-anä öf thc crystal.-lites' yJä,'nx iBääi;rH:', t",:m:t',t''**,ffi1 #:ü! kiß"t+1",1"""'t.fH:J#§ixj;r"i'ffi""J"::l'll{-ü m**ä-tli:*ll*"i"+;.Tii,Fri:*;il:":ä"'H during-Tü! diagencsis' Jiri.nt,i. pothways leading from thc oriSinal shell lo must Dc ,r," lotiii ä" som'etimcs'very diffrcult to trace' but cephalopods' *tr[ffi1ffi**tr***H,r whe'n analvzing fossil ä#;,;;;ffi;iü oncs' üä iil".,üiL -Forstucrures can be mismken for Primary ;ä''rä"r;;;;. cxample, marinc aragonitic ccmens' ä'ffiä,*,".,',ll*"ffi,'?,lt"d*ll,ääl,"s,ii'iJTll; iüi.üä utraty necdlc'lilic and commonly grow syntaxl' oricntation as crystallites in the pnsmatrc air-*irft tnc sanic frgm äJ.':';i"";"-ä;;iv- äin'cutt to'disrinsuish. 11^91, Hff ::?lu"'i im:f.,ojtti,iir'l'-.$?li,,i'l"n;::ff*t'i"t' färmcd whilc thc ffiä,§#;ffir"flt """-"'*t f,r#;$]F*#"*ft*[g'l:ä'mffi ääiä.,;; ü;b"d bv Bandcl.and stanlev.(1988) In somc growth continusd under mannc :üii;:-tä.iti; ccmerit ii""."ir"üitf iq'?r.-lfi""l,i*:L:1,,::,*:':1"'1"{,gf*ä.i#j;"äL; *ri uoay su-rrounding it' The thc animal's death' --fi"-Ä-*tiondiüons after i'ttc ä'ää':ä;'äütinää-iäai*i'"ttl$^.d'::"§^3Yf I or aiugenetic strucrures can rcPlace nacre' äi liä-J""'i'e i: oic oit) rcctc in Pseudoriluccras ftom.rhe li";#ä'il;l'ä "!i' -i:i^* ""ir-näiäuctt $i*'#"}iix**:"1*i9""i",.[,9e-rncarrBundenbach'From Bandcl 8nd l#üitrji$.|iffi1'J,td'*öj,1TäxTi'"".i§li,ää1,1 Ge"r-Y. BANDEL 99 all well-Drescrvcd Triassic, Jurassic and Cretaceous ammonitci so far sodicd (Birkclund, 1967; Bükclund and Hanscn, 1968; Drushchits and Khiami, 1969, Etbc et al' 1968; -Kulicki, 1979; Bandcl et al., 1982i Bandel' 1982' Khiini. 1986). A §imilar pattcm is found in thc cmbryonic shcll of bclcmnitcs and auläcoccradds (Bandel, 1985; Bandel et al., 19841Bandcl 8nd Kuticki, 1988), and this probably also @curs in bacridds and onhoccratids, which havc cmbryonic shells likc bclcmnidds in gcncral. Äs in Spirzla, the first chamber within thc ammonite ammonitelli is sphcrical, ovoid or lenticular (spindlc-shapcd) and it probably- surrounds the embryonic inncr yolk src. Sccretidn of thi cntirc ammonitclla was completed in contact wirh thc manüc epithclium to mineralization, bccause Fis. Skctch of the cut and polished cnd of thc -arc Prior 3. polottcd the ammonitellac dcvoiä of growth lincs, and thc frrst ohiaemoconc and rosEum of Belemnoteuthis their outer walls (Figurcs 4 and 5; was traclureo mineralizaiton affects only ?uidälc Jurassic, Poland). Thc phragmocone jB: mincralization of inner walls, of üc hcalcd rostrum 5A: first mincralization, iu;np lifc but thcn hcalcd. Ponions bcforc construction of telcoconch). This frst (arrows). Chambcrs closc to the 5C: shcil .-n-ä*"rüii^riirii" ttt iture as necdlc-shapcd crysta.llitcs aranged dcposits' From Bandel mineralization appcars äär #"räü-fiirea-ui siconoar! in sliehtlv sohcäilitic oricntations growing from thc inside an^d Kuliclii (1988, Fig.9). ourrrüds. tvhcn a sculpturc of tuberculcs is prcscnt' thc crystallites show thcir lpherulitic orientation bcst within devcloo which rcflccr thc ultrastuctural pattcrn of thc these ubercules, orsaniä shctl containcd in thc mineral marix (Bandcl 8nd Intcrcsdngly, s.rchacogastroPods initially mincralizc their iii"i"r, issS). About six costulcs arc pre-scnt.in €ach orsanic cmbrvonic shcll in a manner similar to ammonltes millini,crcr of Arthroplryllwt shell with thls lond oI anä such ceohalopods that havc an cmbryonic shcll similar in äi;ä;;i;, -d th"t.'feätures can bc uscd for taxonomic size to amrionitejlae. The Cambrian ancestors of ccPhalo- ;ffisä. ;iti-s;A reason, whcn thcir origin is takcn ino pods and gasropods alikc may havc dcveloped this mode of äccäunt It has-o bc undcrstood rhal lhc finc costulcs wcre ihell mincralization in response to a changc troq,-nce iJää u" üii"t"rion of thc minenl pans of thc shell, so i*r.r"ing, non-fecding larvac rcquiring 8 tight. shclt to ihcv rcpriscnt nothing that would bc visiblc on 8lrcsh shcll.' benüic iuvcniles in nccd of protecuon through mrncrarlza' ;;'iih;ili;;; äi grä**', omamcnt, nor microstructural tion. ihc vcry rapid growth of nccdlcs 0.2 microns in morphology. äiuä"r"r into rtic oieani; shcll may rrPrcscnt s primordial modc of initial mincralization in conchifcran molluscs. III. Structurc ofthc Embryonic Shcll' A. Mincralization of the Organic Primary Shcll' As in manv othcr molluscs thc structurc and morphology of thc cmbryonic ccphalopod shcU arc rclatcd to thc amount of ,irlr.