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Radiolarians and Tethyan Radiolarites from Primary Production to Their Paleogeography

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Radiolarians and Tethyan Radiolarites from Primary Production to Their Paleogeography Chapter48 RADIOLARIANSAND TETHYANRADIOLARITES FROM PRIMARYPRODUCTION TO THEIR PALEOGEOGRAPHY Patrick De Wever CNRSUM 1761,Laborûtoire de Géologie Musé m Natiowl d'Histoire Naturelle FlançoisBaudin, JacquesAzéma, and Eric Fourcade CNRSUM 1761, Départementde CéoloqieSédimentairc UniveÆitéPierre et Marie Curie Park, France I. INTRODUCTION Paleogeographyis establishedusing mainly plate kinematicsand unfolding mountainbelts, but basinalpaleodepths are definedusing specific facies, among which radiolaritescan provide key information. The commoneqlJatioî rqdiolqrites = deep oceanic âasir is obviously too simple to be correct; and to infer from the geologicalmessage (e.g., radiolarites),the original,environmental, biological sig- nalrequires a goodknowledge of the successivefilters that have changed it andthe originatorsof thesesignals (e.9., the radiolarians). Thebiology of thegroup, as far asfeeding and ecology are concemed, must be takeninto account;in effort,the present-dayradiolarians hold the key to the past. The resultsof such studieshave made it possibleto documentprecisely paleo- The Ocean Basinsanil Mareins, volume8: The TethysOcean, editedby A E. M. Naim ?t dl. Plenum Press.New York. 1995. Itttd'ltlhrrltl ltl|dkrlllrllrI thtrl(.k | )(, t\(,vo,{,t /r/ It||rthhr'lruN l)irlhyrrcr'yon l!rhysMcs.z.ie ()l ccntralcapsulc in thc lilc ol radiolariansThe rrrr|,s.llrr.rrrri, r.csulr ol.rhis srr(ly ir rhcidcnri lllustrillcsthc ilttl)(nlirtlcc lhc Iicuti.rurrtl l.crrli.n.l upwcllirgs skclctondepends on the centralcapsule (Hollande and i, t'c.cc., arr'ingrhc M.*,;nl Wcstrcss thc li rn tion ol lhc siliccous c(fualiontkl/.)/.r.ik,.r is the only pafi of interestto the micro- ubulklun!uutt.i(ttt, hccausc: thcy cxisl ilt itn ufwclin! lirr,iunrct.1960). 1'his siliceousskeleton cnvilonntcnt.lhis vcrysinrpliliccl statcnrcntrùrst bc rnrtlificd by lukilgsome othcf pulconk)logist. pafantctcrs. with seawaterand suchas dissolution and diagcnesis, into accorttrf. The skeleton,included within the cytoplasm' is not in contact Radiolariansarc ntarine protozoarians ,l.hcy during the cell life Skeletal growth is disconttnuous; . witha siliccoustcsf. naveexlstc(l not cxposed to dissolution sinccthc Canrbrian and conrprise ,l.hcy altematewith stases'Periods of silica precipitationare a largenumber of specics.,r, olienare founrl phasesof rapid skeletogenesis In rocksothcrwisc unlirssilifèrous. physiology than on the environment(Matsuoka' 1992)' andtheir strutigraphic valuc is vcryhrgh, espc_ ttrnr" d.p.nà"nt on the cell ciallywhen they arc thc onlyfbssils present. They were disrcgardcd lbr stratigraphie purposesfbr a longtinrc, bur havebeen srudied thoroughly:;ince the 1980s.Thcir 0. BiologY uniqueassociation with ophiolites allowsdating of f"rloA, of active,oceanrc spreading(and crucial times of l, Reproduction conrinentaldrift). Becausc of thedifficulty rn extrac hng radiolariat'rom siliceous rocks, nev- the first zonâtionsof the Mesozoicwere pro Many aspectsof the reproductivecycle of radioladansrcmain unsolvedl posedonly duringlate l9?0s, first with partitionand sporogcncsis' theCretaceous, then lbr theJurassic, antl crthcless,two typesof multiplicationcan be recognized: laterfor the Triassic. interpretedby Kling (1971)as a f, tlimorphismobserved in someradiolarians is oi gcncritlionsin thc rclllo- rcxu l dimorphism that could result from an alternation (1977)'Inultifliclliu) l)y ductivecycle (as in foraminifera).Accortling to Hanson ll. THE SILICEOUS pLANKTON: in nsw cclls CYTOLOGY, BIOLOGY, flrsion may occurbefore the secretionof a new skclcton AND ECOLOGY OF RADIOLARIANS 2, Nutrililtt Â. Oeneral Châracteristics wlth prcy or llacliolariansarc floating predators They await ritndonl contact Itadiolariansbelong to the class in the environmentThe captureis by meansof filopodes' of Actinopotls. They arc characterizedby tlrc whh nutritivepanicles plcscnccof a capsularmembrane, which within the ectoplasm'They are omnivores' and feed on a wide physically separatescctoplasm from enckr 0tt(l prcy is ingested lllsrr. They incluclesymbiotic algae polycystines, organisms: silicoflagellates, tintinnids' other protozoans' in their cytoplasm. wrttr thcir Vût'icty ol tnicroscopic siliccousskeleton, are forms as large ând active as copepods' etc' the only radiolarianss./. preservcdu, iorrli, presenlly. tlirrtorirs.algac, bacteria' and possibly nllx)ngpolycystines, the Nassellarians (zooxanthellae) also contribute to râdiolarian nutrition Radio- are the mostcliversified, but Spumeltariuns iyrrrbiotic irlgac sccntto be for relatively long periods the most abundant(Lombari and Bowden, l9g2). Int'ilrn c,uttui,ting algal symbionts are able to exist thealgae' glllt,uttu|1,ur"nt, nutrition,as long aslight is availableto sustain "*tclnal seemsto be ancient' becausediver- ll. Cellutar Organization 1'h$ struiS inlcldcPcnrlcncewith the symbionts is the llty,,t nr.ii,,lariun*,tndphytoplankton, notably the acritarchs-dinoflagellates' A relationshipexists between the centraicapsule (basis lirr thc brokrgicirl lltttl('lltt(trlglnrrrl gcologictl time (Anderson,1983)' classilication)and the skeleton(basis of the paleoniologicalclassilication;. Thcl. lorc. thesctw(, typL,sol clussilicutionsirre not tulirlly iidcpcn.lcnt. ll, lleokrgy Iladiolar.iansarc all marine, and most Iivc irulutc,lintlivitluals, our sorrrr. s1'tccicsarc colonial.Sontc ol-these ". tltdiotarirtttsilrc prcsentin all the oceansand open seas' cobnies arc as big as 5 nr and includcnlrrrv lixr'lrtrivclytttittittc, spccintcnswithin a conrnton.gelatinous lnlss. F-oIi",rtate.l inttivi.lu ts, lhc cytrr plirsrrris suslaincd by thc skclctalcrcmcnrs. l. ll,'ttlirt' IltÈ.crilttitl eirpsrrlc.eonristing ol.encklplasnr (!tnd its inclusi{)ns), 'l'llcy to ltlachto floating ., nuctcusillrl |{ tt r | | r t I t t t I t t t t r tttr'||llr\ivf' lhrlllilll!|rl pllllisltIs irtcablc r.. irx.l)rir\r,rs lIllrtc(l hy rlrc cirpsuliu nrcnrbranc.An inclividualtlrirt l]irs hr\l ('vrfytl',1 lntl llr(.ccnlritl ||h|e{lr|||||x'i||||||||||||1.||!||lt||||x.|||||l|||vv{,||iC||||l|||v(|||(.||ls(|.1()|l|2()()to350m cir'srrlcis itblclo rcgcncril(call thc Inissi'l.l tlx'(olltftll trl wllielt vrllicswilh l)itrts.lllt\ f iIlrrrlk'r tttrlt lrr' tr'lrrlr'rllrt I'tl' tlltllol{'N' ,l,]::']':|''',.'l|]/t|n/lIl'.|l'1|l,|t{|t('|||ll|ll(1l|n'|ll|f|l|(.l|!nh|w'|s([l|di'i||iiU|sw||i.|i'(||! llll(l(lllllllrrl llellv- lnlrtlttrtl tcl[t||rlhrlll('(t,|llxlllrlhrltlltlllllrfl'llll llllrrl'rhVlllll('ll( rll\rtrttfl{'lnlfrl lrrfllvlloll (t't l, Urelttrl rlerteltt | vtlllll l lllrll[ll t'ltlr'lllr ^LllvÈ, (lccpelwlttcrs (luring storllN), l{&lioluli rtscxisl itt ll ;r'crictltrltty ttccntts, liltrtt ltolcs lo thccqtlltl(,t, httl lhcit' 2. Iltulioluriun l)ixtributiotr lhundttnccvutics, ln gcncrttl,rtuliolttlitrrt rlislribuliott (lhcys lhc sllnlclilws it$th(tsc C()nlrollingplankttltt: wittct tltllstics,clirnrllc, lItitudc' tlcpth' hydrrtlogiccttnditittns As fbr other planktonic organisms,radiolarians occur in profusion whcre nutri (0,g,, tcrnpcraturc)and thc abundanccol llutricnts(trxnc than thc uvailabilityol' ment ls abundant. Becauseof their symbiosis with algae, radiolarians are mosr Therelbre,thc long-stândingclaim ol a connectionwith volcanistrt abundant lllicu in watcr). near the lower limit of the photic zone, circa 50_200 m (Fig. l). Radio h clroneous. larianslive in the upper part of the water column. They do not require a-deepocean, Radiolariansaccumulate in abundancein sedimentswhere productivity is hiSh but are found in deep-watersediments because they are able tà reach significant the suprajecentwater column, in divergent-waterzones (where cold, nutricnt-rich depthsafter death. In Wutcrascends into the photic zone), such as in pedequatorialzones, polar belts, and some continental margins where upwelling is active Quantityper m3ol waterfiftered In rec€nt sediments(Miocene-Present), the radiolarian abundanceat a sitc is 5000 10(m10000 )^ oonnectedeither to high- or low-standsof sea level (e.g., South Atlantic coast ol1 Afiica, WalvisRidge; Diester-Haass et al.,1992:Hay and Brock, 1992).Hcncc' heightof the sealevel is important only in the modifications it induc€sin currcnl the variations (silica abundanceversus nonabundance,high versus low scll associatedwith the upwetling. Moreover, radiolarian location can migratc with timel therefore, the local sedimentaryrecord also can migratc understandthis generaltrend, it is necessaryto take global phenomenainto In brief, sealevel \ariation and silica abundanceprobably are connected this abundance,but the prime factor is the existenceor absenceof upwelling Radiolarian associationsshow a latitudinal distribution. It also is possible to 7 tivtng speclmens surfacefrom subsurfaceassemblages. In sedimentsbelow upwellings (e.g,, NNI Discordedsketetons a mixture of fauna systematically occurs the cold- and warm-water offPeru. where Arctic watersare mixed with tropical ones;De Weveret cl., 995), or as surface and subsurfacewaters mix off Somalia (CatJletet al., 1992). in work on o the chancesof depicting tropical Tèthyan versus boreal fauna from radiolaritesare almost null, becauseradiolarite facies result from SILICEOUS SDDIMENTS: TAPHONOMY OF RADIOLARIANS Fourmajor processes control the evolutionof a biogenicooze from its initial to its present state: (l) supply of biogenic material, (2) dissolution ol material during settling, (3) dilution within the sedimentby other comPonents biogenic or not), and (4) diagenetic hansformation of the initial pro- Fig. L An exampl€of radiolarian death,an individualtest is at leastpartially dissolved
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