t2 Biogeographyof the Late Paleocene Benthic ForaminiferalExtinction

ELLENTHOIYAS

ABSTRACT The latestPaleocene benthic extinction thus was corn- During the Late PaleoceneThermal Maximum (LPTM) plex.and factors such :is changes in deep-seacirculation, benthic fbraminif'era at rniddle bathl'al and greater increasedCaCO. corrosiveness.increased lemperatures, depthssuffered extinction of 30-507cof speciesduling a decreasedoxygenation. and changesin the patternsof lew thousandycars. Extinction was less sevcrc at neritic high productivitymay havecontributcd to its sevcrity. to upper bathyal depths, where temporary changc-sin launal compositionprevailed. Preextinction dcep-sea Environmental Setting taunaswere cosmopolitanand diverse.and contained heavilycalcified species. Imrnediate postextinction fau- The late Paleoceneand early Eocenewere the warmest naswere more variable geographically, exhibited low di- epochsof the CenozoicEra: polar ice capswere almost l'ersity.and were donrinatedby thin-walledcalcareous certainlyabsent. and shallowseas covered large parts of or agslutinatedtaxa, possibll, because CaCO, dissolu- all continents.During theseepochs, major changesoc- tion increasedglobally liom neritic to abyssaldcpths cured in climate and in the carbon cycle on -elobal just bcfbrethe extinction. Thcse assernbla-ges were dont- scalesof rnillions to thousandsof years. Long-term inatcd either by long-lived taxa sLrchas Nuttttllide.r warming of the deep oceansstarted in the mid-Pale- truerrtpt'iorby bulirninidtaxa. the latteraccompanicd by oceneEpoch (Miller et al. 1987a;Shackleton 1987; a-uglutinantsin somcareas. Kennettand Stott 1990;Zachos et al. 1992.19931 Seto Faunasdonrinated by N. tmentpri rverecontnton in the 199-5).Warm-water pelagic marine organismspene- Sor.rthAtlantic and at lower bathl'al through upper trated to polar latitudesby the end of the Paleocene abyssaldepth in the lndian Ocean.and might inclicate Epoch (Stott and Kennett 1990; Premoli Silva and oligotrophicconditions as well as incrcasedcorrr)si\L-- Boersma 1984: Boersmaet al. 1987: Aubry 1992; ness.Buliminid-dominated launas might indicatehigh Berg-{ren1992). Thermophilic vertebrates occurred in ratesof depositionof organicmatter or lor.l-oxygencon- theArctic (Estesand Hutchison1980: McKenna 1980t ditions.Such llunas werccommon globllly alongconti- Markr.r'ick1994). with vegetationand soil types indi- nentalrnargins. and locallv co-occun'edu'ith sedimcnto- cating warrn clirnates(Kemp 1978: Nilsen and Kerr logicor planktonictaunal indicators of highprocluctivitr. 1978:Wolf'e1994: Basinger et al. 1994:Manum 199,1). In the bathyalcentral Pacific. however. buliminid-domi- Clay mineralassociations in oceanicsediments at high natedfaunas co-occun'ed with planktonicfaunas sug- to midlatitudes indicate high humidity and intensc- gestinsoli-sotl'ophi'. and thev could reflectlow-oxygen chemical weathering(Antarctic. Robert and Kennett conditiorrslesultin-u from sluggishocean circulation. ox- 1992;New Jerseymargin. Gibson et al. 1993;New idationof clissociatedmethanc hydrates. or rlarming ol' Zealand.Kaiho et al. 1996).Oxygen isotopeclata sug- bathl'al irbyssalwaters caused by a chan-gein deep-sea gest that latitudinal temperaturegradients were ver)/ circr.rlation.Alternatively, they could indicatethat thc shallow (Shackletonand Boersrna198 l: Stott et al. tiaction ol'organic matter rcaching the seafl(x)rin- 1990: Barreraand Huber 199l: Zachoset al. 199,1: creasedas a rcsult of decreasedoccanic oxygenf,tion. Braloweret al. 1995a,1995b; Lu andKeller 1995b).

214 The Late PaleoceneBenthic Foraminiferal Extinction 215

Ratesof speciesorigination and diversitywere high fbr and Miller 1992 Lu and Keller 1993. 1995a,1995b; ten'estrialvertebrates and flora (Hooker 1991;Collin- Canudo et al. 1995: Aubry et al. 1996: Thomas and son 1983:Reaet al. 1990:Winget al. 1991,1995; Maas Shackleton1996: Schmitz eL al. 1996).superimposed et al. 1995).as well as fbr pelagicoceanic organisms on a long-term decreasein values that startedin the (Romein1979: Boersma et al. 1987:Boersma and Pre- middle PaleoceneEpoch (Shackletonand Hall 1984; rnoli Silva 199l: Corfieldand Shackleton1988: Mc- Shackleton1986. 1987:Corfield et ai. l99l; Corfield Gowran 199l; Aubr1, 19921Berggren 1992: Corfield and Cartlidge 1992 Corfield 1995).An isotopeexcur- 1993:Kelly et al. 1996).The latePaleocene and earliest sion similar in magnitudeand estimatedduration oc- Eoceneepochs thus constitutea time of major innova- curredin the toothenamel of herbivoresand in carbon- tion of the biosphere(Briggs 1995 ). ateconcretions on land (Koch et al. 1992.1995: Stott et The unusualwarmth of theseepochs has beenconl- al. 1996).and in terrestrial-derivedorganic matter in a rnonlyexplained by high atmosphericpCO, concentra- New Zealandmarine section(Kaiho et al. 1996).The tions.probably ciiused by'plate tectonic processes such 'uvholeatmospheric-oceanic reservoir thus undervu'enta as massivevolcanism in the North Atlantic Volcanic rapid.negative shift in carbonisotope values during the Provinceduring initial openingof the North Atlantic. LPTM. the durationof which hasbeen estimated at be- decarbonationof limestoneor oxidationof organic-rich tween 50 kyr (Thornasand Shackleton1996) and 200 sedirnentsduring the beginningof the India-Asia conti- kyr (Kennettand Stott l99l ). ncntal collision. and high hydrothermalactivity along Massbalance equations show that this carbonisotope nridoceanicridges (see Thomas and Shackleton1996 ercursionwas so largethat it probablycould not have tbr a review). Proxy data for atmospheric7rCO. levels beencaused by transf'erof terrestrialbionlass into the tenttrtivelysupport higher values than todav (Cerling ocean-atmospheresystem or by eruption of volcano- 199l: Freernanand Hayes 1992). although this rematns genic CO.: it was so rapid that it probably could not debatable(Stott 1992).Clirnate modelin-e indicates that have beencaused by a changein depositionor eroslon at suchCO. lcvelstropical temperatures would be much ratesof carbon in carbonateas comparedto carbonin higher than deduced fiorn oxygen isotope data. and organicmatter (see Thomas and Shackleton1996. fbr a mechanismsfor highly incrensedheat transportfiorn review). This leaves f'ew explanationsopen. and re- krr,i,to high latitudesat the low latitudinaltemperature centlirit hasbeen speculated that this unusual,negative gradientsremain r.rnexplained (Sloan et al. 1995:Sloan carbon isotopeexcursion could have been causedby andRea 199-5). massivedissociation of (isotopicallyextremely lightt In the latestPaleocene Epoch the world warmedeven oceanicmethane hydrates as a resultof the deep-ocean more fbr a shorttime (Kennettand Stott 199l: Pak and warming(Dickens et al. 1995:Kaiho et al. 1996). Miller 1992:Thornas and Shackleton1996). which has the Late PaleoceneThermal Maximum been named Benthic Foraminifera TLPTM:Zachos et al. 1993).This eventoccurred rvithin paleomagneticChron C2:1r.in nannofbssilBiochron The latest Paleocenebenthic fbraminif'eralextinction NP9 (= Biochron CPt3),and befbrethe last appearance has been describedfrom land sectionsand at oceantc datur-n (LAD) of the planktonic fbraminif-er Mr.,rrr- drill sites. at depths ranging from outer neritic to .ot'ella veluscoen.si.s(Aubry et al. 19961Berggren and abyssal(appendices 12.1 and 12.2:figure l2.l). This Aubry 1996).Oxygen isotopedata indicatethat inter- extinction was rapid: it afl-ectedbenthic fbranlinif'eral mediateto deepocean waters globally warmedby'1 to faunascatastrophically worldwide. and it occurreddur- 6" C over lessthan a f'ewthousand years (Kennett and ing or close to the beginning of the LPTM and the Stott 199l: Thomasand Shackleton1996). but tropical ne-eatir,'eexcursion in carbon isotopes.just before the to subtropicalsurface to neritic temperaturesremained Paleocene/EoceneEpoch boundaryas most commonly constant(Stott 1992; Bralower et al. 1995a.1995b1 Lu defined(Thomas 1989.1990b: Stott and Kennett1990; and Keller 1995b:Schmitz et al. 1996).This rapid Nomura 199I in combinationwith Setoet al. 1991: Pak warming of the deep ocean has been explainedbi' a andMiller 1992;Bralower et al. 1995a,1995b; Canudo changein deep-to intermediate-watercirculation. dur- et al. 1995in combinationwith Ortiz 1995:Aubry et al. ing which the dominantsources of deepand interrnedi- 1996:Thomas and Shackleton 1996: Kaiho et al. 1996). ate waters shifted from high to subtropicallatitudes This major break in deep-seabenthic foraminiferal (Kennettand Stott 199l: Pak and Miller 1992:Thomas lbunas occuned between the Paleoceneand Eocene andShackleton 1996). epochsif Viewedat coarsetimescales. whereas plank- Coeval with this short-termwarming was a shifi in tonic foraminifera i.rnderu'enta major extinction at the carbon isotope values by about -2'/rr, in surfaceand endof theCretaceous Period (Cushman 1946; Beckmann deepwaters of all oceans(Kennett and Stott 199l: Pak 1960:von Hillebrandt1962). After the late Paleocene 215 THCI'1AS

described [:1::':ii;i,,T::il]li:,:llljH;'il...,1jp,;T,i:::"""'as inappendices r2r and r2 2: nun,bers (driil sites)and rerters Thc paleoge ogr.phl rf thecon rine n r: in rhe,atr' pureue eneEpoch is aficrZach.s et al. ( I 99.r).

benthic extinction faunaswere predominantlyof lclw (ODPI Sites 689 and 690 (Wedevent. Fau_ and Miller 1992)probably do not cover the time inter_ nal comparisonsmay be more difficult because of dif,- val directly after the extinction f-erencesin taxonomicconcepts becausethe rnagnrtude of differentauthors and of the 6lrC because anomalyis much smallerat thesesites than of the differencein sizefiaction being studied, at others,suggesting that its minimum value (and varryingbetween 63 and 150pm in most thus ."r.i, (opp.n_ the detailsof the benthic dicesl2.I ancl12.\. fbraminif-eralextincrron) oc_ cured between High-resolution samples. faunal data and carbon rsotope Most authorsconcluded that a change analysesare available fiom progritm in cleep_water Ocean Drilling circulationplayed a major role in the extinctron.that The Late PaleoceneBenthic Foraminiferal Extinction 2|7

't o1'the deep to intermediatewaters of all oceans benthic extinctionevent and 55 Ma to the NP9/NPlt) '::rcciat leastfbr some time at subtropicallatitudes. biochronalboundary. I derivenumerical ages fbr ODP -l that decreasedlevels of deep-seaoxygenation re- Site 690 assumingconstant sedimentationrates be- .rnc fiom the increasedtemperature of thesewaters tween the benthicextinction and the NP9NP10 zonal ::.Jrl causolfactor (e.g.. Miller et al. 1987b:Thomas boundary.and betwcenthe benthicertinction and the -.9 1990b, 1992'.Katz and Miller l99l; Pak and top of Chron C25. For samplesfiom DSDP Sites,525 '.l lcr 1992,\995 Kaiho 1991.l99zlb: Nomura l99l: and 527 andODP Sites689 and 865 I assignnumerical ,irorri et al. 1991,Ortiz 1995;Spei jer et al. 1996b). agesaccording to Ber-ugrenet al. ( 199,5)to daturr levels r-.1-r\.however. suggested that the patternsof f'aunal in Thornaset al. (1990).Shackleton et al. (198.1).and , .nctionvary geographicallyand that their complex- Braloweret al. (1995b).Finally. I revisethe numerical ' . -'.lnnotbe explainedby decreasedoxy_senation only agesat thesefbur sitesuntil the benthicfbraminifcral :,:.. Nomura 199l: Coccioniet al. 1994:Thornas and carbonisotope curves agree with that fbr ODP Site 690 . ,,.'kleton1996). (Thornasancl Shackleton 1996) in orderto correctopti- r:r this chapter.I first erplore postextinctionbenthic mally for .",aryingsedirr.rentation rates. especially fbr the :',:rninif-eralfhunal patternsfbllowing the extinetion dissolutionzone at Sites525 and 527 and fbr the inter- .:rg high-resolutiondata liom the sites mentioned val with very low sedin.rentationrates just abovethe ex- - re. Then I use literaturedata (figure 12.1.appen- tinctionat Site86-5. . -:. ll.l and 12.21to try to evaluateoceanic envlron- .:rt;rl changes between late Paleoceneand early Selectionof Literature Data Sources -.ne times.to deterrninethe biogeographic pattern of : :rtinction. and to speculateon its possiblecauses. Appendices12.1 (DSDP and ODP sites)and 12.2(land sections)list sites fbr which data are available that straddlethe extinction level: hence only a lew sitr's Methods fiom T.jalsrnaand Lohmann( 1983)are included(figure l2.l). Data on sectionsin the NorwegianGreenland Srnple Preparationand StratigraphicFramework Sea and Labrador Sezrwere nclt included becausein High-ResolutionStudy ':. n.ranyof the sectionssedirnentation probably started . . iirs chapternew data are presentedfrom ODP Site afterthe extinctionevent (FIulsbos et al. 1989:Karrin- '-5. and a f-ewadditional sanrples from ODP Sites689 ski et i.rl. 1989). Data on agglutinateclfaunas were .i 690 and DSDP Site -527lfigures 12.2 and 12.3),as mostly not included. e\ren thou-ehthey suggestthat , :rparedto data in Thomasand Shackleton(1996). changesclccun'ed in agglutinatedfaunas at the end of \.,rrples fbr benthic fbraminif'eralfaunal and isotope the PaleoceneEpoch (e.-s..Geroch and Novak 1984: . ..:lrsis were taken fl'om DSDP Site 527 (Walvis Kuhnt and Karninski1990: Kaminski et al. 1996).be- :,.Jge.South Atlantic), ODP Sites689 and 690 (Maud causeexact data and precisecorrelations are difficult in < .c. WeddellSea). and ODP Site 86-5(equatorial Pa- the absenceo1'carbclnate and thus of isotopedata. Data , 1rt. They weredried overnightat 50" C and weighed. fiom the North Searegion are included becausc detailed :.:n soakedovernight in distilledwater. Most samples stratigraphicdata are available (Gradstein et al. 199:l). , 'rrggregatedreardily and could be washedover a 63- Ferv paperson neritic faunasare included because ,:: screen.Benthic fbraminifera fbr fhunal analysis benthicfaunas at thesedepths are more geographically :r'e picked fiom the >63 prrnsize fiaction. fbllowing variableand planktoniczonation is con.rmonlydifficult: lh,rmas ( 1990a). All specirnenswere picked and therefbre.it is not possibleto detenxinewhether ob- ': ,rLrntedin cardboardslides. All samplescontained suf: servedtaunal changecan indeedbe correlatedto the -.ient specimensfor analysis(>250). Taxonomy is as late Paleoceneextinction in the deepsea. For instance. :: Thorras( 1990a)and Thon.ras and Shackleton( 1996). paperson neritic to brackish water Paleogenefaunas ,,:rdlargely fbllows van Morkhoven et al. (1986).The fiom the northern North Atlantic were not included -,rr'Ciesrichness was usedto calculatespecies richness (Berggren1974a: Murray 1989).Data on neriticsections r a sample of 100 specimens usin-s rarefaction in E-eypt(Sperjer 199,1; Speiier et al. 1996a.b)were in- Sanders1968). cluded becausedetailed carbon isotope data are avail- I use the integratedmagnetobiochronologic fiame- able (Charisiand Schmitz1995: Schmitz et al. 1996). '.r.rrrkof Berggrenet al. (1995),which is basedon the Data trorr.rSpain (Molina et al. 1992 Ortiz and Mc- :eomagnetic polarity tirnescaleof Cande and Kent Dougall I 991) weretentatively included because plank- 1992)and modifiedby Candeand Kent (1995).For tonic fbrarniniferaldata suggest that benthicextinctions ODP Site 690, I use the biostratigraphicinterpretction were at leastclose in time to the extinctionin the deep ,i Ar-rbryet al. ( l996). givin-ean age of -55.-5Ma to the sea. Data fiom coastal New Jersey were tentatively 2I8 THOIYAS

bi/triserialspecies, 7o N. truempyi, "/. (infaunal morPhotYPe) 10 20 30 20 40 60 80

o

-o

(100) G aveli n ell a beccariif orm i s, 7" numberof species 5 20 30 40 50 55.35

55.40

55.45

o 55.50

55.55

55.60 Figure 12.2 Contparisonof the spcciesrichness and relati"'eabundance of significanttaxa at the time ol'the late Paleoceneben- thic lbranriniferalextinction. See text firr clerivationol'thc' age moclel. Datafirr DSDP Sites 515 and 517 and ODP Sites 6lJ9 and 690 allcr Thotttrs ancl Shacklcton 689.and 690: data fbr ODPSite 865 lra\e not 1ct bcen published.

included(Olsson and Wise 1987;Gibson et al. 1993: 12.2 and 12.3).This percentageis a minimutn estimirte Gibson and Bybell 1994) becausethe clay mineral becauseseveral nodosariid species are rare, so their recordat thesesites may be a meansof correlationto rangescannot be determinedwith accuracy.Some no- ODP Site 690 (Robertand Kennett1994). dosariidsare absent in the intervalconesponding to the LPTM, but reappearhigher in the sectionat varyinq levelsthroughout the lower Eocenesection. Results At both Walvis Ridge sitesthe extinctionoccurs just above the base of a level with severedissolution. al- H igh-ResolutionStudies though benthic fbraminit-erawere well preserved: At ODP Sites689 (paleodepthll00 rn) and 690 (1900 planktonic tbraminifera show severedissolution and (Thomas 1996).No low- m), DSDP Sites-525 (1600 rn) and 521 (3400 m). and fiagmentation and Shackleton ODP Site865 (1300-1500m) benthicforaninil-eraldi- eringof theCaCO, percentageof the sedimentoccuned versity dropped precipitouslyat the extinction level. at Sites 689 and 865. but faunasat these sites show with 35 to 5OC/cof speciesbecoming extinct (figures more etchingand are lesswell preservedjust abovethe The Late PaleoceneBenthic Foraminiferal Extinction 219

numberof species(100) BFAR (nr/cmZky) buliminidspecies, 7o Cibicidoides, % N. truempyi,% 20 30 40 50 60 o 5oo 1000 1500 0 25 50 75 0 10 20 30 10 20 30 35

40

45

CJ

c)

55

3a JC 60

Figure 12.3 (a) Speciesrichness at 100specimens, (b) benthicforaminiferal accumulation rates (BFAR). and relative abundances Lrf(c) buliminid taxa.(cl) Cibititloicle.s spp.. and (e\ Nuttallidesrruentpt'i at equatorialPacific Site 865. \urnericalage s accurdingto C--andeand Kcnt ( 1995 ): agernodel atier Bralou er et al.( I 995a.I 995b ) extinction level. At Site 690 similar slight dissolution abundancesvary strongly between sites.Tappcutina scl- occursin the faunas,and CaCO. valuesdecrease just rnensisreaches up to 40% at the Maud Rise sites.up to below the extinction level (O'Connell 1990; Thomas l5c/cat the Walvis Ridge sites.and up to 25Vcat Pacific t992). Site 865. At all sites.peak occurrencesof T. selmensis At all sites.Gavelinella beccariifonnls has its upper- occurdirectly above the extinctionlevel, whereas those most occurrenceat the extinctionlevel of severalother of A. ctrttgorten.rl.soccur at the top of the LPTM interval. taxa (seebelow). Our data confinn the observationby Despitethese similarities, postextinction faunas are Katz and Miller (1991)that this speciesis most com- most notable for the major difl'erencesbetween sites. mon at higher-latrtudesites: it never reachesabun- especially in the abundancepatterns of buliminid danceshigher than about 5% at equatorialPacific Site species,Ntrttallides truempti, Oridorsali.s ttntbottatus, 865. Other speciesthat havetheir highestoccurrence at Cibit'idoidesspp., and abyssaminidspecies (Thomas this levelinclude several large agglutinated species with and Shackleton1996; appendix 12.1). The postextinc- calcareouscement (Dorothia ox\:('otto.Tritario ltura- tion faunas diftbr much more from site to site than the rtensis. and Trita.ria pttleocenicci) and calcareous preextinctionfaunas, which arecomparable to Velasco- speciessuch as Arugortictt,elasr:oen.sis. Bolit'inoide.s type f'aunas at all five sites (see also Berggren and tlelit'rttulu.s.Bulimina mitlvruven.si:;.Bulimina thatrcte- Aubert 1975;Kaiho 1988).These postextinction f'aunal nis, Neoflabellina reticulctttt.Neoeponitles hillebrandti. differencescannot be completely attributedto the pres- Neoeponicleslmntu, and Pullenia c'on'elli. enceof stratigraphicgaps and comparison of noncoeval Immediatelyabove the extinction level, speclnlens f'aunas.because faunas at all five siteswere coeval with are smaller and have thinner walls than below. even the short-term carbonateisotope excursion. Data were specimensof speciesthat straddlethe extinctioninter- collectedby one researcher.so thereis homogeneityin val (e.g., Oridorsali.s umbonatus and Nuttallitles the databasewith regardto taxonomicconcepts and size tuempti). At ODP Sites 689 and 865 the changeto fiaction. Postextinction f'aunas differ rnore by geo- smallerand thinner-walledtaxa occursa f-ewcenttme- graphic region than by depth; f'aunas fiom the two ters below the highest occurrenceof G. bet'ctrriifitrni's' Walvis Ridge sites and the two Maud Rise sites are possiblybecause of bioturbation. Tappanirnselmertsis quite similardespite large difl'erences in depth, andAragonier aragonensis are at all sitesmore common At the Walvis Ridge sites.postextinction faunas are abovethe extinctionlevel than below (asalso observed donrinatedby lV.truempt'i, O. urnbonatu's,and abyssa- by Boersma 198,1b:Speijer 1994), but their relative minid species:the latterare especiallyabundant at the 22O IHOITAS deepestsite (figure 12.21.Abyssaminids (.Abt"tstttrtitttt 738 (Lu and Keller 1993),but the benthicfbrarninif'eral pottgi. Abt'ssrttrtitttrclutrdrorrl. and Clinttpertinupluni- recordis limited. lmmediatepostextinction faunas are .qrriz) increasein relativeabundance postextinetion at very poorly preservedbecause the extinction occurs MaLrdRise. bLlt much less than at the Walvis Ridge r,r,'ithina layer rvith very strongdissolution. I observed sites.At Pacific Site 86,5these species have very low verycommon small. bulirninid species (including I .ie1- abundancesthroughout, with a slightfurther decrease at ntensi.sand ,8. .simple.r)just abovethe extinctionlevel. tlre extinction level. Nuttallitles tntemPti declines suggestingthat the faunalpatterns at this site resen-rble stronglyin relativeabundance at andjust abovethe ex- thoseat Maud Rise. tinction level at the Maud Rise sitcsas rvell as at Site Detailed clata on faunarsu'ith age control by thc 865. in stron-qcontrast with the situationin the South LPTM carbon isotopeexcursion are availablefirr the Atlantic.At the WalvisRidge sitesthe postextinctionly'. Zur-nayaand Caravacasections in Spain(Canr-rdo et al. tnternpt'ispecirnens are smallerand havethinner '"r'alls 1995: Ortiz 199-5;appendix 12.2).In both seetiotrs than preextinctionspecirnens. but are much more nu- stron-gdissolution occllrs at the extinctionlevel. as indi- merousthan in preextittctionfhunas. cated by thc presenceof dark gral'. larninatedshales' At the extinctionlevel the relativeabur.rdance of the Faunasjust above the extinction level of G. beccati- buliminid species decreasesat Site 527. doesnot ifttnnis aredominated by the agglutinatedspecies HaTr- -eroup changeat Site 525. but increascsstrcxgly at both Maud lophnrgrrtoitlesretrosept(t At Zunaya. abovethe dissc'l- Risesites and Site 86-5(figure 12.2). Above the extit.tc- lution interval there is a thin interr"alr,'u'ith commot.t tion levelthis groupis dorrinatedby snlirllspecies such Bulinrirtcrtu.rpumettsi.; (Bulitnirttt spp. Assernblage): as T. seltnensi.s,Bolivinrtrrft'^s sp. cf . B. decoralc, and higher N. tntempt'i is dorlinant. At Caravaca ,V. Bulinina sirnple.r.At the Maud Rise sitesSiphttgeneri- truemltviis not as comrnonas at Zumirya,and it is ac- noicla.sltrevi.sltittosu is contmon(up to 40%) acrossthe corrpanied by Anornttlinrtitle.scctpitcrttt.s and Oritlorsuli's extinctionlevel, at theWalvis Riclge sites it is nluchless ttmbortttttts.The Bulinina spp. Assenrblagen.ray have conrnon (up to l-5%).and at Site 865 its abundancers persistedsontewhat longer at the someu'hatshallower similar to that at Walvis Rid-ee.br-rt the specieshas its Carar.'acasectiolt than at Zut.naya.but fbr this part of the lrighest occun'enceat the extinction level. Bulintirttt section age control is not as precise,and short-terrn ,semitostcrtcris abundant at PaciticSite 865 iust above changesin sedimentationrates nlay obscurethe pattern. the extinctionlevel. then decreases in abr-rndancebut re- In the Tawanui Section(Neu' Zealand;Kaiho et al. mainspresent throughout the lou'erand middle Eocene 1993,1996; appendix 12.2) the benthicextinction oc- Series.At the Maud Risesites this speciesappears cl.rly cursin larninated.dark gray.organic carbon-rich shales. in ZoneNPl2 (51-52 Ma. Berggrenet al. 1995).In the Age control is provided bv recogniticlnof the LPTM lcrwermostpostextinction specirnens of B. setrtit'rtsttrtrt 6lrC excursionin terrestrialor-uanic matter. Within a at site u65 the costaeon the lower part of the test.typi- .l-cm intern'aldirectly abovethe hi-ghestoccurrence of cal lbr this species.are unusuallvthin and the speci- G. bet'cctriift,nrlr.itaunas are dominatedlry Btrlintitttt rleus are srnall;possibly the speciesori-einated at this ttr,tltcntensisand Prtteglrtbobulimirtcrpupoides. Above time from a noncostate.trihedral Bulitninu specles. the Buliminu spp.Assernblage there is an -S0-cm-thick At Site U65the predominanceof buliminid species interval with r,'erystrong dissolution. in which faunas appearsto have lastedfbr a shortertin.re than at the aredominated by a-eglutinanttaxa (Rhubdutntrtirr.rspp.. Maud Rise sites.At this site the intervalof bulirninid Cottotntchtunntirtudepres.ra. and Reoplta.rregularis\. clominanceis only l5 cnt thick.representing about l0 Higher in the section faunas arc lnore diverse. with and kyr in our agernodel. where as it is about-500 cnl thick. conlmon Cibic'idoitle.Espp.. Arnntttlinoides spp.. representingabout 100 kyr. at the Maud Rise sites. B. Iurpanren.si.s. Above the buliminid-dominatedinterval at this sitefall- We can concludefiom thesehi-sh-resolution studies nasare dominatedby Cibit-idoirft'.sspp. up to a level in that postextinctionf'aunas are characterizedby low di- Zone NPI I (zonalassi-snment in Braloweret al. 1995b; t,ersity.and that geographicditterences cannot be at- 52.U553.61 Ma in Berg-erenet al. 1995;fi-eure 12.3)' tributedto insufficientage control. difl'ering taxonornic where the relativeabundance t'tf N. trLtemp.r'rincreases concepts,or differingsize fracticltls str.rdied. With this in to l2-25c/c.At noneof theothcr sitesis therean interval mind. we can try to evaluatethe lower-resolutiondata dorrrinatedby Cibicidoirle.ispecies (up to 307c). Ihe availablein the literature. taxonomyof theseCibicitloitles species is still underin- lar-ee.flat shallow-waterEu- vestigation:they resemble Literature Data ropeanspecies such as C. pntprius (Brotzen1948). A detailedrecord of planktonicfbraminif'era and iso- Beckmann(1960) was the first authorto recogni/ethe tonesis availablefrom southernIndian Ocean ODP Site imoortanceof the benthicfbraminit'eral ertinction at the The Late PaleoceneBenrhic Foraminrferal Extinction 221

end of the PaleoceneEpoch. based on his study of the studiesdemonstrate that the extinctionwas a very short- hathyal LizarclSprings Formation (Trinidad). His clara term event, so contparisonof faunal events between tas given in Bolli et al. 199,1)show a very str.on-{in_ sites is problernaticfbr data that are so disparrte in creasein highestoccurrences as well as a decreasein chronologicresolution, a-qe control. taxonomic concepr. lowest occurrencesof benthicforan-rinif-era in the Lrpper and size fiaction studied,as presentedin the appen- Paleocene Moro:.ot,ellavelct,st'oensis Zone. close to the dices.However. some overall conclusions can be drawrr Paleocene/Eocene Seriesboundary (Bolli et al. 199.1). fion.r these diita. especiallvin contbinaticlnwith the Benthicfbraminif'eral faunas in the overlyin-s,:lzone hat'e high-resolutiondata. ,rnruch lower dil'ersity.There are more lastappearances In the first place.the extinction\\'as se\ere at midclle ,)t agglutinated. noclosariid.and rotarliidtaxa than of bu- bathyat through abyssal depth. with extinction of lrrninidtaxa (figure 12.4). Another early recognition of speciesran-eing between 30 and -507c.At upperbathyal ihe irnportanceof a latePaleocene benthic fbrantinif-eral through neritic depthsthe ertinctions were much less 3\tinction event was by von Hillebrandt(1962). who severe.but faunalcomposition changes r.vere highly sig- lr)ticed a strongdrop in diversityof benthictbramini- nificant.These corrpositional changes were temporiiry ierirlfaunas toward the top of his M. t'elct.scoen.sisZone and many speciesreturned to the sites (Speijer 199:t; Zone F) in the Reichenhall-SalzburgBasin (figure Gibsonand Bybell 199,1). i l.-5). In thesesections. however. rotaliid taxa appearecl Second. preextinction Paleocenefaunas were re- he ', the ntoreeotnnton \urvi\ r)rs. markably cosmopolitan,extending over a wide depth The irnportanceof the benthicfbraminif-eral extinc- range(Brotzen 1948: Berg-uren 1911a. lgllb: Ber-ugren i.n was recognizedby many later authors(appendices and Aubert 1975).The larter authorsdistinguishecl a 1.1 and 12.2)in studiesof land sectionsas well as of continentalshelf fauna (Midway-type) and a lou,ercon- DSDP and ODP sites in all oceans.High-resolution tinental slope and abyssalplain fauna (Velasco-tvpe).

firstappearances, last appearances, % of totalspecies % of totalspecies 20 20

P7 P7

P6b P6b

P6a P6a

P5 P5

P4 P4 = P3b P3b c0 benthicextinctiori o c P3a P3a o N P2 P2

P1c P1c

P1b P1b

(a) F-igure12.1 Lorvestand (b) highestoccurrences of speciesbelonging to the four mostimportant of taxaas a percenta-ce -sroups of thetotal nunrberof speciesintheLizardSpringForrnation.Trinidad(Beckurann1960:Bolli etal. lgg.lt.Thehorizontallinc showsthe levelof the bentliicextinction. 222 THOIYAS

parent difference in provinciality between late Pale- + agglutrnants + nodosariina oceneand early Eocenefaunas might be thoughtto re- + buliminidea sult at leastpartly frorn the lack of a monographon the a^ + rotaliidea o- early Eocenebenthic fbraminiferal taxonomy compara- o (1975) :O ble to that of Berggrenand Aubert for the late o PaleoceneEpoch. so that thereare greaterdifferences in !^ 5N taxonomicconcepts between authors. This cannotbe the C full explanation,however. because greater difTerences between early Eocene faunas fiorn diffbrent locations have beenobserved in faunasstudied by one investiga- (9 L to < tor (Tjalsmaand Lohmann 1983:Kaiho 1988;Katz and ,on"luonnirroeor"not rogozl Miller l99l). Finally, postextinctionfaunas were low Figure 12,5 Numbersof speciesof the four mostinrporlant of diversity groupsof speciesin the Reichenhall-SalzbLrr-eBasin (von andhigh dominance,in starkcontrast with thehighly di- Hillebrandt1962). The verticalline showsthe levelof the verse.low-dominance preextinction faunas. The postex- benthicextinction. tinction community thus hiis erstructure typical for fau- nas afTectedby a major environmentalperturbance (e.g., Pearsonand Rosenberg1978). In theseperturbed com- although severalspecies (e.g., Bulimina ntitlv'atensi.s munities the speciesT. selntensisand A. (rragorren.ri.sare and Con'phostonn midv'avensr.r)occur in both faunas comrnon to abundanteverywhere below shelf depths (e.g..Kaiho 1988, l99l: Nomura 1991;Kaiho et al. (appendices12. I and 12.21Boersma 1984a). 1993;Sperjer 199.1: Coccioni et al. 1994).The Midway Otherwise. f'ew thunal observationsare valid on a Ihuna typically contains more Cibic'idoirle.iand espe- worldwide basis.The postextinctionpattern as seenby ctally Anonuilinoide.s species, whereas the Velasco Tjalsma and Lohmann (1983) and Katz and Miller fiiuna has A. velu.sc'oensis,Gyroidinoide,s spp.. N. (199l). i.e., dourinanceby 1/. tntempt'i with survivor truempti. and agglutinanttaxa such as Dorothia o.tt'- speciessuch as O. umhrnatu.r.appears to be typical for contt, Guudn'ina lnev'igata, Trita,rio lnvanen.si.s.and the South Atlantic and possibly Gulf of Mexico; Tritcr-riupuleocenico. The common speciesN. truenrytti abyssaminidtaxa are common below about 2-500m in generallyis consideredtypical fbr the Velascof'auna this type of assemblage.Thomas (1990a. 1990b)sug- (below about -500n). Tapparina selmensisoccurs fiom gestedthat this associationmight be seenonly in studies the shelf (about 100 m) down to abyssaldepths. but is usingthe largesize tiaction (>150pm) becausemany of rarein bathyal-abyssalfaunas below the levelofthe ex- the buliminid taxa are small. High-resolutiondata in tinction.Both Midway and Velascofaunas are charac- Thomasand Shackleton(1996) and this chapter(figure terizedby extremelyhigh diversities.the occurrenceof 12.2) show that this is incon'ect.and that faunas are many rare taxa (especiallynodosariids). and thick- dominated by N. truenrpli and abyssaminids even walledcalcareous taxa. thoughthe >63-pm fiaction is studied.as was also ob- Third. there were geographicdifferences in faunal servedby Mtiller-Merzand Oberhiinsli( 1991). assemblagesin spiteof the cosmopolitanoccurrence of Patternsin the North Atlantic are more variable.Most many species.Gavelinellu becc'uriifrtrntis, fbr instence, North Atlantic data are from the Goban Spur-Bay of appearsto be more common at high-latitr.rdethan low- Biscay Sites,and postextinctionfaunas vary from buli- latitudesites (Katz and Miller 199l). althoughit was minid dorninatedto N. truentpti dominated.Possibly, alsocommon in theTethys (Speijer 1994\. Arugonia ye- the two typesof faunascan be differentiatedby depth: lascoen.sisis extremelyrare at both Maud Rise sites. Buliminid-dominatedfaunas appear to occur at rniddle whereasS. breyi.spinosais abundantat these sites. but bathyaland shallowerbathyal depths (DSDP Sites5218, showslarge fluctuations in relativeabundance (Thomas 549; Reynolds1992; Boltovskoy et al. 1992),whereas 1990a. 1990b).The geographicdifferences have not N. truentpt'i-dominatedfaunas occur at lower bathyal been well evaluated;the availability of more oceanrc throu-ehabyssal sites (Schnitker 1979 Pak and Miller drill sites in depth transectswould help in separating 1992;appendix I 2.I ). All thesesites. however, show in- depth frorn geographiceffects. and in distinguishing creaseddissolution and occumenceof unconformitiesin Paleocenebiogeographic provinces, possibly similar to the lower Eocene stratigraphicrecord. which makes thosein the latestCretaceous Period (Widmark 1995). faunalcomparisons more diflicult (Aubry et al. 1996). Fourth. this high degreeof faunal cosrnopolitanism Bathyal land sectionsclose to the Bay of Biscay endedwith the late Paleoceneextinction (Kaiho 1988; (Coccioniet al. 1995; Ortiz 1995) show buliminid- Thomas 1992:Thomas and Shackleton1996). This ap- dominatedpostextinction f'aunas, although the patternis The Late PaleoceneBenthic Foraminiferal Extinction 223 corlplicatedby the occurrenceof agglutinatedtaunas in noides species, and Cibicidolrle.s.Shallower faunas an intervalof severedissolution. In New Jerseymargin (100-300 m) containmore buliminids,but on the shelf DSDP Site 605 N. trtrentpti-dominatedfaunas occur Cibic'idoide.sspp. and especiallyAnomalinoitle.s aegt'1t- abovethe extinctionlevel (Hulsbos 1987;Thomas un- llacr.s dominate.Sections in Spain (Ortiz 1995) and publisheddata). Shelf faunas(Olsson and Wise 1987: Italy (Braga et al. 1975; Di Napoli et al. 1910: Gibson et al. 1993)in the region show common Epis- -500-1500m depth) show stronglv increasedpercent- tominello,Pulsiphonina. and Tunilina, and cannoteas- agesof buliminidsabove the extinctionlevel, including ily be comparedto the deeperfaunast coastal sections Bulirninct semicostattt.Faunas from the Reichenhall- generallyhave unconfbrmitiesat the extinction level Salzburg Basin (von Hillebrandt 1962; 200-800 m) (Gibson and Bybell 1994).The postextinctionfaunas show increasedabundance of rotaliid taxa.but detailed fiom Trinidad are buliminid dominated (Beckmann stratigraphyis not availablefiom thesesections. We can 1960:Bolliet al. 1994r. concludetentatively that westernTethys faunas were In conclusion,we can speculatethat the northern similar to North Atlantic fhunasfiom the samedepth. Atlantic abyssalto lower bathyal faunaswere similar In the IndianOcean, no simplefaunal pattern can be to South Atlantic faunas (N. truempv-dominated). extracted.particularly because many DSDP and ODP whereasthey werebuliminid dominatedin the upper-to siteshave records with poor recoveryand unconfbrmi- mid-bathyalreaches. We cannot be certain.however. ties. Faunasat some lower bathyal-abyssalsites (ap- that we are indeed comparingcoeval faunasbecause pendix 12.1) seem to resembleSouth Atlantic faunas tlissolutionand unconformities are common. (N. truempt'i-dominated),but abyssarninidsare much The bathyal faunas in the North Sea (depths less common or absent.possibly becauseno lower 750-1000 m) were dominatedby noncalcareousagglu- abyssalsites are available. Other taxa in the postextinc- tinated taxa through the late Paleocene and early tion f'aunasare mostly lon-e-termsurvivors such as O. Eoceneepochs (Charnock and Jones 1990; Gradstein et Ltmbonatu.s,although B. sentic'o.statomay be present al. 1994).These agglutinated taxa underwenta period abovethe extinctionlevel. Faunas at upper and middle of severe ertinction close to the Paleocene/Eocene bathyal sites(appendix l2.l) have very different pos- Epoch boundary.after which extremelyimpoverished. textinction faunas.dominated by Lenticulinn species low-diversity. ag-tlutinant faunas remained (e.9.. (Mackensenand Berggren1992) or by Anonutlirnides Charnockand Jones1990; King 1989:Gradstein et al. species(similar to fhunasin the easternTethys: Speijer 1994).Direct postextinctionsediments indicate stron-q 1991). Such f aunas dominated by Artomalinoides CaCO. corosivenessand dysoxicto anoxicconditions specieshave been interpreted as indicatinglow-oxygen (Charnockand Jones1990; Schroeder 1992). At some- conditionsor high productivity (Speijeret al. 1996b; what shallowerlevels (100-500 m) calcareoustaxa (in- Nomura and Kennett.personal communication 1995). cluding G. beccuriifonzl.i) were present durin-e the At all thesesites. however. core recoverywas poor and PaleoceneEpoch (King l9u9). Thesefaunas were re- the existenceof the LPTM 6lrC excursionhas not been placedby extremelylow-diversity agglutinated f'aunas fully documented.High-latitude ODP Site 738 (1350 similarto thosefrorn deeperregions. The timing of the m), however.where the carbon isotope excursion is pre- disappearanceof the calcareoustaxa is difficult to es- sent(Lu and Keller 1993).has the buliminid-dominated tablishbecause of the biostratigraphrccomplexity of the patternof the Maud Risesites. North Searegion (Gradsteinet al. 1994;Berggren and Data on the Pacific Ocean are available from few Aubry 1996).The local North Seadisappearance of G. sitesonly at bathyaldepths. Northern Pacific ODP Site becc'uriiJonmsprobably did not predatethe global ex- 883 (Pak and Miller 1995; 1000-2000m) has a poor tinction ol C. beccariijormis at the LPTM by very much upper Paleocenerecord and an unconformity, but its time becauseit can be correlatedto the lower part of di- postextinctionfaunas appearto be of the N. truentpt,i- noflagellateZone D5 (King 1989).which hasbeen cor- dominatedtype. Faunasat DSDP Site 577 (Pak and related to just below the middle part of nannofbssil Miller 1992;Kaiho in press;1800-2100 m) havecom- Zone NP9. Palynologicalstudies indicate the occur- mon Bttlintirtusemic:ostulrr above the extinctionlevel, renceof a peakwarm period somewherein dinoflagel- althoughbulirninid percentages are much lower than at late Zone D-5(thus Zone NP9; Schroeder1992). which Site 865. At this site. however.a 0.-5-rnsection in the might possibly be correlatedto the LPTM in mid- criticalinterval was not recovered(between Cores 9 and BiochronNP9 (Aubry et al. 1996). l0 in Hole 577), and the sectionprobably contains un- Tethyan faunas are known fiom land sections only confbrmities(Aubry, personalcommunication 199-5). (appendix12.2). Sections in the easternTethys (Speijer Sites577 and 865 (describedabove) thus have bulim- l99ul)from a depthrange of 500-1000m show postex- inid-dominatedf'aunas. Land sectionsin Japanand New tinction faunas with common N. truemot'i.Anomali- Zealand(500-1500 m: Kaiho 1988:Kaiho et al. 1993. 224 THCI'1AS

1996) show buliminid-dominated f'aunas.although rronly beenattributed to lctcalf'actors such as basinr..- cornplicatedby dissolutionintervals with ag-ulutinated striction.which certainlyrnay have contributed. but ti:- taxa. severedissolution occurred synchronously with worlc- Data on the neritic thunasdo not indicatea relation- wide dissolution.In the easternTethys (Egypt). Speile ship betweenthe benthic extinction (thus the LPTM) (1994) observeddissolution layers over a wide dep'l:. and changesin sealevel. The cxtinctionevent occumed range.Benjamini ( 1992)reported sr.rch layers fiont tit. during the secondorder supercycleTA2 of Haq et al. north centralNegev (lsrael).and Berg-erenand Aubr. (1987),and might havebeen ccleval with part of short- (1996)tiom the Khieu River sectionin Georgia(N\\ term third-ordercycle 2.3. in mid-nannofbssilBiochron Caucasus).Similar dissolutionintervitls occur in lan.- NP9. Spei.jer(199/+) and Speijeret al. (1996b)did not sectionsin Spain(Tethys and Atlantic sicles:Coecioni e: observesignificant sea-level efl'ects in neritic seetious al. 1994:Canudo et al. 199-5). in Egypt. Europeanand North Anrericansections have Dissolutionis more cLfficultto trace in DSDP an.: many Llnconfbrrnitiesin their upper Paleocene-lower ODP sitesthan in land sectionsbecause poor recover\ Eocene stratigraphicrecord (e.-e.,Olsson and Wise is a problem.A thin laver(<,50 cnt) of sedinrentmav no: "n, 1992; Gibson and Bybell 1994; Berggrenand Aubry be recovered.and thereforemany sitesare listed as 1996).su-e-eesting that sea level was relatively low at data" in fi-ture 12.7 and appendix 12.1.We catlnotb.' the tirne. In the northernSpanish sections the benthic sure of the tirll geographicand bathyrretricextent trl extinction event has been correlatedto a lowstand carbonatedissolution. but it was extensive in antl (Pujalteet al. 1995).lt lrasbeen proposed. however. that aroundthe Atlantic Ocean.as well as at leastin partst,l sea-leveltrends in the North Atlantic reflectedthe de- the Indian Ocean,along the Tethysmargin. and alon-c velopmentand collapseof the IcelandPlume under the the Pacificmargins. We haveno dataon the Pacifrcdeep North Atlantic Volcanic Province,and thus were re- basinfloors. which were probablybelow the CCD clLrr- gionalrather than global (Nadin and Kusznir 1995). ing most of the Paleoceneand Eoceneepochs (e.g.. a: shown by red clays in Cole LL,I,I-GPC 3. Rea 199-1l. We cannotbe sureabout the exacttirr.ring of the initir- Discussion and Speculation on CaCO, tion of this increaseddissolution. but it may have Dissolution, Productivity, and O, Levels started slightly befbre the LPTM (Thomas l99l: Thomasand Shackleton 1996: Kaiho et al. 1996). CaCO, Dissolution The occurence of such widespreadCaCO., dissoltt- tion during a periodof warming is unexpectedbecause The global data set on the benthic fbrantinif'eralextinc- rising ternperaturesalone would resultin ciecreasedsol- tion event is incompleteand requireshigh-resolution ubilit5rof calcite(e.g.. Broecker and Peng 1984).There- studiesto describein detail the eventsthat occurreddur- fbre. the occurrenceof widespreaddissolution strongh in-ethe LPTM. Data are neededfiom sites located on indicatesmassive addition of carbon to the ocean-at- depth transectsin order to discrin-rinatebetween depth mospherereservoir. and the patternof dissolutiotr(rnore and geographicefl'ects. Most infbrmation is fiorn sitesat severearound the Atlantic. less in the Pacific)resembles lower bathyaldepths (figure 12.6a).with f-ewdata liom that predicted to result fiom n.rassiveburnin-e of fbssil abyssaldepths. Geographically. most data are fiont the fuel and defbrestation(e.g.. Walker and Kastin-s1992). North andSouth Atlantic oceans. Coverage in thePacific The occurence of the extremelynegative carbon iso- Oceanis extremelypoor. data being atvirilableonly tion.t tope anomaly in this interval of dissolutionsuggests middleto lower bathyaldepths (fi-eure 12.6b). Coverage that the addedmass of carbonmust have beenisotopi- is slightly betterfor the Indian Ocean,but recoveryis cally unusuallylight. The amountof biosphere-derived poor at rnanysites. In addition.taxonomic coneepts ft-rr organic carbon neededto causesuch a large rsotopie the postextinctionf-aunas. specifically for Bulinrittrrspp.. anomalywould have to be reflectedin widespreadde- Cibitidoidesspp.. and Anonralinoicle.t spp., needs to be structionof land biomass.which does not agreewith unifbrrnizedbetween researchers befbre global patterns paleontolo-ticalevidence (Thornas and Shackleton canbe establishedwith any degreeof confidence. 1996).Therefbre. widespread carbonate dissolution as- Combining lithological and paleontologicaldatir in sociatedwith the 6lrC excursionslrpports the specula- this incompleteset. however. sug-qests that a ren.rarkable tion thatdissociation of methanehydrates (and their ox- globalepisocle of carbonatedissolution occurred at least idation of methaneto CO.) may have been involved partially coeval with the LPTM (figure 12.7). In the (Dickenset al. 1995;Kaiho et al. 1996).This specula- North Sea region calcareousspecies occuned only in tion implies that the temperatureincrease of the deep the shallowestneritic areasof the time (Kin-e 1989: waterspreceded the carbonisotope excursion slightly. Gradsteinet al. l99zl).This severedissolution has com- in agreenrentwith Thomasand Shackleton( 1996). (-)(-)AeC FF> .9roturgcaG 66-^9E u\v- 'L: -o_o_oti6P€*€ss ZA oc:+;0Jlyo);;i o-c uso -t

Figure 12.6 Distribution of sites listed in appenclicesI l. I ancl 12.2 according to (a) clcpth and (b) geographic location.

: .tuthorsol theoliginal papers und grlen rn appendicesI l. I and I l.l.

n no data il no dissolution Z slightdissolution Z dissolution

OOA(JE !t> :::!O; => c(u(c?-:*>--o.5dftfi

za arlgoal;i er3a-; <=> 5=r?30 t

Figure 12.7 Occurrcnceof'clissolution across the intcrr,alof the benthicfbranrinif-eral extinction by (a) depthand (b) geographic location:sites listed in appendices12.1 and 12.2.depth categories as in li-eurc12.6. ercascddissolution ilnd etching of nriclotaunas.but no clcar scclinrcntar-r'er iclencc-. 226 THOIYAS

Suchan increaseofthe ocean-atrnospherecarbon reser- this oxidation(e.g., Mullins et al. 19851Hermelin and voir is in contradictionwith Stott'sinterpretation (1992) Shimmield 1990:Levin et al. 1991;Schmiedl 199-5: of data on the carbonisotopic composition of organic Wishneret al. 1995).There is considerablediscussion matter in foraminif'eraltests. although such data cannot whethersuch benthic faunas reflect the low oxygenler'- be easilyinterpreted (Goericke and Fry 1994:Hinga et els or the high food supplies(e.g., Corliss et al. l9U6r al. 1994).Stott's data (1992) might be explainedby Linke and Lutze 1993: Miao and Thunell 1993; Lou- many factors. such as species-specificeff'ects. diage- bere 1994: Rathburnand Corliss 19941Jorissen et al. neticefl'ects. effects of varying temperature.or salinity. 1995).Recent species of Bulimina areparticularly com- More problematicfor the methanehydrate hypothesis mon in areaswith high ratesof depositionof nonde- might be the precisetiming and sequenceof eventsdur- gradedorganic matter (Corliss and Chen 1988;Caralp ing the LPTM: dissolutionappears to havepreceded the 1989;Corliss 1991 ; Mackensenet al. 1993).Additional 61rCanornaly. factors of importance might be the lack of metazoan predatorsunder low-oxygen conditions (Wishner et al. 1995).and somethin-walled taxa have beendescribed Dissolved Oxygen Levelsand Productivity asbeing opportunistic taxa rather than directly linked to What caused the benthic fbraminiferal extinction. and high productivityor low oxygen (Alve 1994;Goodal what can explain the biogeographicpatterns observed? I 994). This is a difficult questionto answerbecause there is no We thus cannot determine fiom the benthic f-aunas agreementon the interpretationof Recentbenthic fau- alone whetherthe buliminid-dominatedpostextinctiort nas (see e.g., Gooday 1994, Schnitker 1994. fbr re- faunasprimarily reflect low-oxygen conditions.hish- views). There is no modern analog for faunas domi- productivity conditions, stressedconditions. or hi-ch nated by N. truem1tt'i because this species became CaCO.,corrosiveness. Similarly. we cannot determine extinct in the late EoceneEpoch. Its modern relative, whether the N. truempll-dominated fauna reflects the Nuttallitles wnbonif'era, is common to abundant in occurrenceof CaCO,-corrosivewaters or low produc- CaCO.,-corrosivewaters, and specificallyin Antarctic tivity. We needto considerevidence from co-occuring bottom water (e.g., Bremer and Lohmann 1982: planktonicorganisms and sedimentaryand iscltopicevi- Schrniedl 1995).Others. however. have describedN. dencein order to evaluatethe most probablecauses for untboniferaas a typicallyoligotrophic species (Loubere the extinction.and comparedata on dissolutionwith 1991,1994; Gooday 1993,199.1). Possibly, the waters thoseon benthicfaunas (figures 12.7 and 12.8). in which N. urnboniferais common are CaCO. corro- It appearsprobable that the occurrenceof small and sivebecause of the presenceof abundantdissolved CO. thin-walledbenthic foraminif'era following the benthic derived fiom oxidation of or-eanicmatter, leaving little extinction primarily reflects increasedCaCO., corro- food fbr benthic faunas.thus cornbining oligotrophy sivenessof deep watersbecause both dissolutionand and corosiveness. Some support for the view that .ly'. thin-walledtests occurred at almostall locations(figure truentpt'irnay havebeen an oligotrophicindicator may 12.7).and the associationof thin-walledtests with cor- be seenin its Eocenedistribution pattern: This species rosive waters is well documented(Boltovskoy et al. had its highest abundanceduring the middle Eocene 1991).Proxies for high productivityor lower dissolved Epoch(e.g., figure 12.3;Kaminski et al. 1989;Berggren oxygenlevels are not ascommon as for dissolution.and and Miller 1989;Miller et al. 1992).It then declinedin thus neither can be seenas the dominant f'actorcontrol- abundanceand becameextinct during the late Eocene ling the occurrenceof the thin-walledtaxa (hgure 12.8). Epoch,at a time when globalproductivity rnay have in- In addition.ostracodes. which are complexmetazoans, creasedconsiderably (e.g., Hallock 1987;Hallock et al. becamethin-walled during the LPTM at ODP Site 689 1991; Aubry 1992;Brasier 1995;Thornas and Gooday ertthe sametime as the benthic foraminifera (protistans) 1996). did. Ostracodeshave been shown to form such "de- In the Recentoceans, low-diversity faunas dominated graded" tests in hydrochemicalregimes unfavorable by small,thin-walled specimens belonging to the buli- to biomineralization.i.e., in CaCO.,-corrosivewaters minid group commonly occur in low-oxygenenvlron- (Steineckand Thomas 1996). rnents(e.g.. Boltovskoyet al. 199l: Sen Gupta and Increasedcorrosiveness thus probablyplayed a role Machain-Castillo1994: Kaiho 1994a).Such environ- in the extinction.It hasbeen hypothesized that the ben- ments usually occur below a well-developedoxygen thic foraminif'eralextinction was largely causedby de- minimum zone,where oxygen levels are low becauseof creasedlevels ofdissolved oxygen (Thomas 1989), and enhancedproductivity and oxidationof organicmatter, a parallel has been establishedwith the Cretaceous and whereCaCO. dissolutionmay be severebecause of anoxic oceanic events. although of lesser intensity the high concentrationsof dissolvedCO. resultingfiom (Kaiho 1994b).The LPTM benthicextinction occurred The Late PaleoceneBenthic Foraminiferal Extinction 227

no data Bb high O2nowprod low O2ihighprod l3

10

=Fd(g(!(g =>o!f,=> =33-oi6i6 bobbai 3€=3: 5E:=l/

Figure 12.8 Occunenceofbenthic l'aunassuggesting either lovn-oxy-een conditions or increasedproductivity by (a) depthand (b) ::rrgraphiclocationl sites listed in appendicesI 2.I and I 2.2.depth categories as in figure I 2.6.

-iurin-erapid warming of the deep waters, possibly the continents(Bay of Biscay. New Jersey ntargin), - rlLlsedby a switchto dominanttbrrnation at subtropical whereas the South Atlantic sites are mainly along ..Ltitudesof warm and salinedeep to intermediatewa- oceanicridges and rise (figures12.1 and 12.6;appendix :L'rs(e.9.. Thomas 1989:Kennett and Stott 199l: Pak 12.l). We thus cannot determinewhether the differ- .,nd Miller 1992. 199,5).Warmer deep waters can be encesin distributionpatterns between the North and the :redictedto havea lower oxygencontent because of the South Atlantic are true geographicdifferences between .trong decreasein solubility of oxygen at higher tem- deep waters in thesebasins. or the difl'erencesbetween :reratures(Thomas 1989, 1990a:Kaiho l99 l). If the open ocean and margin. The lack of indicationsthat .peculationabout dissociationof methanehydrates is South Atlantic waters were especiallylow in oxygen (Dickens -'rrrrect et al. 1995:Kaiho et al. 1996).oxida- content.however, may sug-qestthat warm and saline :ron of these compounds would have used large waters did not dorninantly flow fiom subtropical re- ,rrrountsof oxygenand thus have contributed to the cre- gionsof Tethysinto theAtlantic. If Tethysfunctioned as .rtionof low-oxygenenvironments. There is no agree- a sourcearea. the more probableregion of outflow was nrentbetween the distributionof dissoluticlnand that of the easternTethys, in agreementwith carbon isotope 1ou'-oxygenconditions and/or high prodr.rctivitywith data (Seto 1995).On the otherhand. agglutinated ben- rcqardto depthand paleogeography(figures 12.7b and thic fbrarniniferal assemblagesdo support a Tethys- [.t3b). Atlantic connection(Kaminski et al. 1996). The buliminid-dominatedfaunas. however. did not It remainsthat faunal pattemswere not globally the rrccur Faunasindicating either high productiv- same.and. consequently, decreased oxygen content as a -rlobally. it1'or low oxygen contentswere most cornmonin the resultof global deep-waterrvarming does not fully ex- \orth Atlantic over a wide ran-qeof depths(with possi- plain the faunal changesand the benthic extinction. ble exceptionof abyssalsites). along Tethys. alon-{ the Thomas and Shackleton(1996) used carbon isotope \ntarctic continent,in the centralPacific. and at upper data to interpret the postextinction.buliminid-domi- to rniddle bathyal depthsin the Indian Ocean (figure natedf'aunas at the Maud Rise sitesas indicativeof a 12.8).Faunas dorninated by N. tntemp,r'lwere common high rate of supply of organic matter to the seafloor. rn the SouthAtlantic. and occurredin the northernmost This hypothesisappears to be in contradictionwith the Paciflcand at middle to lower bathyaldepths in the In- vier.l' that the Paleocene long-terrn carbon isotope dian Ocean.It is ditficult to interpretsuch patterns be- recordreflects a decreasein productivity(e.g.. Shackle- causeof the small numberof localitiesto compare:The ton et al. 1985:Shackleton 19t16; Corfield and Shackle- North Atlantic DSDP sites.fbr instance.are all closeto ton 1988:Corfield and Cartlidge1992: Corfield 1995), IHOIYAS a view that is in agreementwith bio-qeographicalpat- oligotrophy occur in the same samplesas buliminid- terns of planktonic microfbssils(e.g.. Hallock 1987: dorninatedbenthic faunas(Kelly et al. 1996).At suclr Boersmaet al. l9u7: Hallock et al. 199l: Ottensand sitesthe faunasprobably reflect low-oxygen conditions Nederbragt 1992; Brasier 199,5).Theoretically, one in the watercolumn. Such low-oxygen waters in the ba- would expect extrernelylow productivity during the thyal openPacific rnay have resulted fiom the high tem- LPTM. when temperaturegradients were shallow,so peratures(e.g.. Thomas I 990a:Kennett and Stott 199I t. that wind velocitiesover the equatorialregions were their fbrmationpossibly exacerbated by sluggishcircu- very low and upwellingand nutrientsupply to the sur- lationdr-re to theertremely lou'temperature gradients. It f'acewaters were almost null (chapter'9). In addition, may alsobe possible.however. that the buliminid-dom- benthic fbraminifera have higher metabolic rates at inated faunasreflect the input of a larger proportion of higher temperaturesand increasedtemperatures thus organic matter on the seafloorunder lorver oxygen con- leadto oligotrophy. ditions, as supportedby the strongincrease in benthic Thomasand Shackleton( 1996),however, argued that fbraminif-eralaccumulation rates at the level of extinc- global productivity might have declined.but that the tion at Site 865 (figure 12.3: Herguera and Berger carbonisotope records at the Maud Risesites suggested r99r). increasedlocal upwellins. and thus possiblyenhanced The peakabundance in buliminid speciesat ODP Site localproductivity. Along othercontinental margins pro- u6-5 is just below a peak abundanceof CiLticitloitles ductivity might have also increasedduring the LPTM. speciesthat was not observedat othersites. The Cibici- possiblycaused by changedpatterns of Lrpwellin-eas a dctide.sspecirnens resemble the Recent species C. result of changingpatterns of deep-intermediatewater v"'ueller.storJi.which lives on elevatedsupports above circulation.High abundancesof chiloguembelinidplank- the sediment-waterinterface and is mostcomnton in re- tonic fbraminif'erain lowermost Eocenesedin.rents in gions with active bottom cuffents (Linke erndLutze the Bay of Biscay and along the New Jerseyntargin 1993).If the earliestEocene Cibicicloide.s had a similar havebeen interpreted as indicatingincreased local pro- habitat. its high abundancemight inclicateincreased ductivity and an expanded oxygen minimum zone botton.rcurrent activity and winnowing, as also indi- (Saint-Marcl99la, l99lb: Pardoet al. 1997).Produc- cated by increasedpercentages of material in the tivity may also have increasedlocally in the eastern >63-um size fiaction. lor'vaccumulation rates, and in- Tethys,where typically a "flood of radiolarians"occurs tensivereworking of planktonicfbraminif'era (Kelly et in the dissolutionlayer (Benjamini 1992:Berggren and al.1996). Aubry 1996).At IndianOcean Sites 138.152, and162 The widespreadoccumence of the //. trtretnltti- the abundanceof chiloguernbelinidplanktonic fbra- abyssaminidfaunas can be tentativelyexplained as re- minif'eraalso increasesabove the extinctionlevel (No- flecting increasedcomosiveness of botton waters.as rnura and Kennett.persclnal communication 1995). In shownby the occurrenceof widespreadCaCO., dissolu- other easternTethys sections, however. benthic faunas tion (figure 12.6).These faunas. however. might alsore- suggestlow-oxygen conditions as well as increased flect an overalldecline in open-oceanproductivity dur- productivity(Speijer et al. 1996b). ing the LPTM. The patternobserved by Tjalsma and Local increasedproductivity during the LPTM in Lohmann (1983), that l/. tmempt'i-don.rinatedfaunas spiteof the decreasedtemperature gradients could have migratedto shallowerdepths in the oceansduring the resultedfrom an increasedinflux of nutrientsfiom the late PaleoceneEpoch. could be explainedby the hy- continents.An increasedabundance of kaolinitein the pothesis that organic matter derived fiom the surf-ace sedimentsdeposited during the LPTM indicatesen- waters becameexhausted at shallowerand shallower hancedprecipitation and chernicalweathering around levelsin the oceansbecause of the declinin-eproductiv- the world (e.g..Robert and Kennett 199,1;Gibson et al. ity. as also suggestedby agglutinatedmicrofaunas 1993:Kaiho et al. 1996).in agreementwith climate (Karninskiet al. 1996).This patternwas not global' and rnodeling(chapter 9). The higherraintall could haveled wasnot observed,tbr instance.at FalklandPlateau Sites to increasedrunofT and nutrientsupply to coastalwa- 698-102(Katz and Miller 1991.;. ters:the increasedinflux of fiesh watercould alsohave The questionremains open concernin-ethe srgnifi- increasedwater stratificationand causedlocal shelf or canceof faunascontaining common to abundantZlp- slopeanoxia (Malone 199l; van der Zwaan and Joris- lttrttino.selmensis, a species that occurs in postextinction sen 1991). f'aunasworldwide over a large depth range but was Increasedproductivity cannot be seenas the causeof common only at neritic to upper bathyal depths befbre all occurencesof buliminid-dominatedfaunas: at equa- the extinction.Although it might be seenas a part of the torial Pacitic ODP Site 865 planktonic fbraminif'eral buliminid-dominatedfaunas. this is not a satisf'actory f'aunasand coccolithophoridsindicative of extreme explanationbecause the speciesoccurs in N. tntetnpti- The Late PaleoceneBenthic Foraminiferal Extinction 229

clominatedassemblages as well. At ODP Site 689 it is Productivitymay havedecreased globally during the nrostabundant together with an ostracodefauna domi- LPTM. but this efTectcould havebeen counteracted lo- nated by taxa related to the present-dayproponto- cally by lower oxygen levels in the water column. re- cyprids (Steineckand Thomas 1996).Propontocyprids sultingin deliveryof a largerfraction of clrganicmatter are opportunisticforms that live dominantlyin neritic to the seafloor.Productivitv may have increasedalong environments.They are confined in the deep sea to continentalmargins as a resultof increasedcontinental $'ood-basedand hydrothernal vent communitiesthat runoff and upwellingof nutrient-enricheddeep waters. fbrm in transient,food-rich environments.I therefore Speculatively,N. truentpt,i-dominatedfaunas may be suggestthat Z selmensis,originally describedfiorn seenas indicatorsof highly oligotrophicconditions on shallow-waterregions, was likewise an opportunistic the seafloor.and bulirninid-dontinatedfaunas as indica- taxon rather than a low-oxygen indicator,similar to tors of eutrophicconditions. The occurrenceof both nrodern taxa such as Stoinfctrthiu ffusiforml.s(Alve f-aunasafter the extinctionsuggests an expansionof the 199,1). trophicresource continuum. In conclusion.I suggestthat the latestPaleocene ben- thic fbraminif'eralextinction was a complex phenome- ACKNOWLEDGMENTS non.Factors that contributed to the extinctionmay have been the global increasein CaCO-,corrosiveness and This chapteris a contributionto IGCP Project308. The the global decreasein oxygenationof neritic through researchwas partially funded by NSF grant EAR abl,ssalwaters as a result of increasedtemperatures. 9.+0-5784to Jim Zachos and Ellen Thornas.I thank The averageoceanic productivity may have declined. Marie-PieneAr"rbry, Murlene Clark. Rodollb Coccioni. br-rtslu_ugish circulation and low oxygenationcould Andy Gooday.and Mimi Katz for their helpfuland con- have led to the sequesteringof or-eanicmatter in the structivereviews. and Robert Speijer.Birger Schmitz, cleepsea. 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D.. 1991.Early Eocene bi' (ierr1 titic Rcsults.v. ll-l:Collcge Station.TX {OceanDrilling Progrilm). otic and climatic changcin interior sestern North Arnerica. p.901 91.1. a.qrtr'. 19.p. ll89-1192. Thonras.E,.. Shackleton. N. J. and Hall. \{. A.. 1992.Data Rcpon: Car- Wrshner.K. F..Ashjian. C. J..Celfhan. C.. Co$in,s.N{. l\'1.. Kann. L.. lron isotopestraiigraphy of Paleogenebulk sedirnents.Hole 76lC I-evrn.t-. A.. Mullineaur.L. S.and Saltzrnann. J.. 199-5.Pclagic and rE\nrouth Platcau. eastern Indian (Jccan). In von Rad.U.. Haq. B. Lt.. bcnthic ccology of the lo$'er interfaccof the EasternTropical Pa ct al.. Procecclingsof the OccanDrillng Program.Scientillc Results. cific oxvgenminimum zote.I)eep-Seu Reseurclt. r. '12.p. 93-l l5 r. 122:College Station. TX (OceanDrillin-q Program). p. 897 901. Wolt'e.J. A.. 199.1.Alaskan PalaeogeneClimates as inf'erredfionr the Ttrlsnra.R. Cl..1976. Cenozoic fbraminilera fiont the SouthAtlanttc. CLAMP database.In: Boulter.i!{. C.. andFisher. H. C.. eds..Ctrto tl tlte Arctic, N,{TO ASI Scries. ll7. DSDP Lcg -16.ln Barker.P F..Dalziel. L \\'. D.. Initial Reportsof :oic Plunt.suntl Clintute theDeep Sca Drillrng Project. \'. -16:Washington (LLS. Gtxernment Frankfurt:Springer Verlag. p. 223 138. PrintingOlfice). p. .193-517. Zachos.J. C.. Rca.D. K.. Seto.K.. Nomura.R. andNiitstrma. N..1991. T.jrlsnu.R. (1.. 1983.Eocenc to N'liocenebcnthic lbrantinilers fiorrr Paleogcneand early Ncogenedeep \\'aterpalcoceiinography of the Deep Sca Drilling Pro.iectSit!'516. Rio GrandeRisc. In Barkcr. Indian Oceanas determineclfiom bcrrthicforatniniler \table cilrb()n P F.. Carlson.R. L.. Johnson.D. A.. et al.. Initial Reportsof the andorygen isotopcrecords. In Dttncan.R. A.. Rea.D. K.. Kidd. R. Deep Sea Drillin-eProiect. r'. 72: Washington(U.S. Govcrnment B.. ron Rad.U. andWeissel. J. K.. (eds.).5t;rtlrs.rirof Resultslhtnt PrintingOihcel. p. 731-7-55. SLienti.ficDrilling in the huliun Ot rarr. GeophvsicnlMonograph 70. T jrlsrrta.R. C. lnd Lohmann.G. P. 1983.Pultot't'tte-Etnt'tre butlnul Washington.D. C.: AmericanGeophl-sical Union. p.3-51 -185. tntl ultt'.tsulltt'rtlhit lorunrini.llnt.fhm tlte Atlutttit Ot artrl.\'[icropa- Zachos.J. C.. Lohmann.K. C.. \lralker.J. C. G. anclWise. S. W.. 199] leontologvSpecial Publication No. -1.90 pp. Abruptclimate change rnd transientclimates cluring the Paleogenc: \ rn iVlorkholen.F. P C. M.. tserggren.W. A. anclEdrvartls. A. S.. I986. A r.narineperspectire. Jtntnrul ol'Geolrtqr', r'. 101.p. lc)l-ll3 Ceno:.oicto.sntopolitut daclt-ttuter bertltit'lorunittifi'rzl. Birlletin Zachos.J. C.. Stott.L. D. andLohtnann. K. C.. 199-+.Evolution of carlv des Ccntrescle Recherchcs Erploration-Production Elf-Acluitaine. Ccncrzoicmarine temperatures. Paleot'eurtogrupltt, r .9. p. 353-3It7. tr4er.noirI l. :121pp. 236 THO\1AS

Appendix 12.I

Recovert,/ Site Location Ref-erence Depth Resolution r-pnr 6rrC Size Unconfbrmitr

Soutlt Atlutttit'

I . I oop 689 Weddcll Sea Thomas 1990a1 I I (X) Hich Ycs >63 Medir.rnr Thornasand ultcontoflnrt\ Shackleton1 996

1.2 onp 690 WeddellSea Thomas1990a: 1900 Hi-sh Yes >63 Good Thomasand Shackleton1996

2.1 oDp698 Falklancls Katz and 800 900 Lor.r' No > 150 Poor Platcar.r Millcr 199I

2.) onp 702 Falklands Katz and I 700- 1800 Low No > l-50 Poor Plateau Miller. lc)9|

2.3 ot)p7(X) Falklands Katz and 2l-50 2600 Low, No > l-50 Poor' Plateau Miller 199I

2..1 osor'329 Falklancls Tjalsrra 19771 17-10l7.l-5 Low No > 150 Mcdiurn Platciru Tjalsnraand Lohmann1983

3.I nsop20C Mid-Atlantic l jalsmaand ll7-5 3000 Lou No > 150. Poor' Ridge.west Lohrlann 1983: Miiller-Merzancl >63 Oberhiinsli199 |

3.2 osop-5 16 Rio Grande Dailey 1983: 1000-2000 Lo* No >l-50. Goocl Risc Tjalsma1983: > l-50. Boltovskovet >63 al. 1995

-+.I DsDp-52:l \\alvis Ridge Clark and Wli-sht 3-500-.1000 Lou, No > 150 Poor 198'11Hsu ct al. I 98.11Parker cr al. I 98-1

1.2 osop-525 Walr,isRidge. Thomasand 1600 High Ycs >63 Poor'/r.plrr Shackleton1996: ln core Boersrra198-l: Boltol skovand Boltovskoy1989

-+.3 trsop527 Walr'isRidge. Thomasancl -i.+00 High Ycs >63 Poor/rplrr Shacklcton1996 ln cofe Boersma198.1

Puci.ftcOt eart

5 oot'883 North Pacific Pak and Millcr 1000-2000 Lou No >150 Poor/ 199-5 uncontornritr

6 osop-577 Subtrt'rpicalNorth Millcr ct al. 1987b: I Sm-l100 High/ Partim > 150 Good/gap Pacific Pak and Millel l99lr rneclium unconlonnlt\ Kaiho in press

7 ttop865 EquatorialPacific Braloweret al. 1300 l-500 High Yes >63 Good/ lc)9-5a.b:Thomas. unconfbrrnifi'.) unpublisheddata. thischapter The Late PaleoceneBenthic Foraminiferal Extinction 237

Appendix12. I Low Ol/ .-t r.tinction Postextinction/ P()stextinction/ High Slrort-Ternr Long-Terrn Productivitl' Dissolr.rtion

r t ttriif orntis, IJ.rnidtraten.si.i, I selrnensi.s,B. sinple.r. Buliminids (8. .setnit'osttrttr). Yes Slight rt'mpti, B. tlttnelen.si.s, A. urttgonensis, S. brevisltirtrt.str S. elegurtta, N. truentpti. ltrtnitle.sspp.. 5. brzll.r1ruro.r'rr O. urrthttnutus

t curii.fbrtrti.s,R. turpentierue, T. sellrcl,sis, B. sirnple.r, Bulirninids (8. .serniL-tL.strttu). Yes Yes ipetensis,N. truentpt'i abyssaminids.S. brevisltirtosti 5. elegurtttt,N. trttetttpt'i

'.,trtriilrtrnis. Cilticidoidas spp.. Cibicitloitlcs spp.N truerrt|tti, No'l Yes nclli, lnliminitl.t O. untlxtnrtttts

-..L t ttri{forni.s. Gt ntidinoitle.s spp.. N. tnrenpti, bulinlinids. No'l rurtbottatus - ir1oirle.rspp. Lenticulinu spp . o

-.ttttriiforntis, P. cortelli. Ab1'ssaninids. Cihititloide.s spp.. No'l . l.1r(.)er?ri.r,Neoaponitles spp. huliminids. N. fruent1tt'i

.,,1L ttri4frtrrtri.s,P. crtrt't'lli, Ab1'ssaminids.N. truempt'i, N. lntenpt'i, O. unbotktlus, No'l 'f. seltnertsis,A. tra,qortert;is, --.rLtinants.Gtntitlirtoide.t spp. A. ttrug,ortrllsis B.;ttttitt'.:!Ltl,t

.,tcuri1fplrti5, bulirninicls. Ab-vssaminids.N. truempt'i, Abyssarrrirrids.N. truentpti, No'l ' .ttloitle.sspp.. agglutinants ,1. trntgurcnsis A. urugonettsii, bulirninids

't'Lturiifbnttis. P. tonalli, l N. truentpti,O. tortbonutus, No'l Yes . /rrit oalists,S. ltrevisltittttstt. ,\. unt,qurcn.si.s,P.proli.ut :t'Iit tttuIus, N. tnternTtti

,t t t uriilrtrnri.s, Ncoeponitle.s spp.. 'l N. Iruentpri,abyssan'rinids. Yes Yes'l Il. tri hed ru. 7'.se I ntettsi.s -:. Lrtinants

:.LLtari|lonni.s, N. truernpti. N. truempti. abyssanrinicls. N. truentpt'i, buliminids. No Yes Neoeponitlcs spp.. T..sehnen.sis,A. rtrtrgorten.si.s Cibititlolrle.r spp. --:lutinants. :'r, llolrlr,s spp.

i,tt coriifornri.s.A. t'elttstttensis. N. trtternpti, abyssaminids. N. trttctnpti. bulirninids. No Ycs :rtrern1tti,B. thuneten.si.s T..seltttensis Cilticidttides spp

'l : bat (trriifbrrnis. B. rnidttutcrt.sis. I N. lruentpt'i,A. urugonertsis. No'l lb.vssarninids -rlutinants. I'. tort'elli

: hct'crrri|fltrnris,P. conelli, '! T..selrrtensi.s,A. urugttnen.si.i, N. tntempyi. bulinlinids Yes Slight ',t,lrt.tt'oensi.s.B.rnitltovert.ti.s Ncrteporrrrlt'.tspp.

: lrt,ctttriiforrru.r,.l. r't,1a.rtr;r,rr.rl.r, T'.selmen.si.t,B. senti(o.\l(tt(t Ciltititloitie.sspp. N. lrrrerrlTr,r'i, Yes Slight . \(,nti(ribr(iug ag,tlutinants bulirninids (Il. sottittt.sttrttr\ 238 THOtaAS

Appendix 12.l (conrinued )

Recovery/ Site Location Ref'erence Depth Rcsolution lprlr 6lrC Size Unconformity

Indiun Oceart

8.1 oopJlJ Kerguelen Mackensenand 2000-3000 Low No >l-50 good/ Berggren1992 unconformitl

8.2 oop 7,18 Kerguelen Mackensenand 600-2000 Low No >150 Poor Berggren1992

9 oup 738 Kerguelen Lu andKeller 1993; 13.50 High Ycs >150. poor/ Nomura and Kennctt >63 lprlt in personalcomnruni- core cationlThomas unpublisheddata l0 osop2:15 SouthMadagascar Sigal 197'1 600-1000 Low No >150 Poor Basin

I I osop237 MascarenPlateau Vincentand Brtin 600 2500 Low No > 150 Poor 191I

12 osop2l-5 NinetyeastRidge McGowranl97,ll 600-2500 Low No >150 Good/lprivt Hovan and Rea not tn corel 1992

13 oop 752 Kerguelen Nornural99l ; -500-1000 High/ Partim > 150 Poor Nonruraand Kennett medium personalcommunl- cation l,t r:tov762 EasternIndian Nonruraand Kcnnett 1000-1500 High/ Partirn >150. Poor/ Ocean pcrsonalcomnruni- medium disturbed cation:Thomas un- >63 publisheddata: Thomaset al. 1992

North Atlantic l-5.1 osop-5:19 Bav of Biscay/ Reynolds1992: 600-1500 Lol'n' Yes/no >l-50 Good/ GobanSpur Berggrenand Aubry benthics unconlormtty l 996

15.2 DSDp:101 Bayof Biscay PakandMiller 1992: 1800 HiglV Partim/ >1.50 Good/ Schnitker1979: Saint medium no/ r.rnconlormity Marc l99la. 199lb; no Pardoet al. in press yes .+2(X) 15.-l rrsr;r''1004' Bay of Biscay Schnitker1979 3100 Low' No > l -50 Poor

15..1 r;sop5:1[J GobanSpur/ Boltovskoyet al. 600-2000 Medium No >6] good/post Bay of Biscav 1992 l6 nsor,605 New Jcrsey Hulsbos1987: Saint 1800-2300 Low/high No >150. Good/ margin Marc 19871l99lb: uncontblrrrrtl' Thomas,unpublished >6-l data The Late PaleoceneBenthic Foraminiferal Extinction 239

\ppendix l2.l (tontinuetl ) Low O2l :':.c-Extinction Postextinction/ Postextinction/ High T.rra Short-Term Long-Tcnl Productir,'ity Dissolutitttt

'l 'i. bet'carii. ortni.s, Cilticidoide.r spp.. N. truentpti. Cibit'idoitle.s spp.. No'l P conelli, N. Iruerrtpti O. Ltmborttttus

j. spp. No'l : ' beccuriilomrrs, bulirninids ? Stilosttntellu Lenti<'ulirut spp.

,'. bet'curii.forntis.N. truentpti. 7'.selntensis. buliminids N. truentpti, buliminicls. Yes Yes 3. tI uneten.si.s, agglutinands Cibicidoide.sspp

G. beltoriilormi.s, A. t'elu.st'oensis. ') T. .selntensis,A. onL,e.utert.sis, Yes'l ). welleri, B. thutetensis N trrterrtl'ti j. hettttriifitrmis, N. truentp.tI N. truentpti. Attontttlinttitles spp. No'l

.. lt t't t'u ri i.lbrnt i.s,N. t ruentpti, T. selntensi.s,Arugoria spp. Yes'l ..:!lutinants

r'. bectarii.fornis, P. toryeIIi, N. truenrytvi,Atromulirutitles Yes'.) ( il,irirl,tile.t.pp.. N. In!(tttlt\ i. durtit'trs,O. rurtbortatus bulirninids

G. bet.r.urii.fbnni.t,N. truentpt'i, N. truentpti, B. sentito.slttttt N. Iruentpti, buliminids. No Cibicidctitle.sspp.. B. thanetensis, Cibitidoides spp.. Cibicidrtidesspp., A. azr- 'l'. B. tlelitunilu.s .selntensis grrtrerisis,O. ruttbttrtcttus

G. be c c a r i i.fonnrs,a-e-ulutinants Bulinrinids.T. se Itne rt.s i.s Yesl Ycs

G. bet'turiiforni.s,C. lrt'plnlu.r. ') N. truentpti, ab1'ssaminicls. N. lruempti, abvssaminids. Ycs Yes (1.ntitlxut'ettsis, B. delitutulus, buliminids(8. senricostutu) bulinrinids.Cibicitlttides spp. Ne o e port i c!e.s spp.. G r roi d i ru t i de.r

Cibicidoide.sspp.. N truenrytt'i, l Cibicidoidesspp.. Aragonia Yes Yes abyssanrinids.G: rrt i tl i rto i tl e s spp. a rugortettsi.s. Ci bit' ido i de s spp.

No data T. selnrcrtsi.s,S. brevi.spirto.su, N. truentpt'i,buliminids. Yes Yes A. uragonertsis,B. Cihit itlt,itlt'r :pp. seilltcostutu

G. beccuriilortni.s.Cibititloide.! spp.. 'l Probabll'unconformtty N. trtrenpti. Cibit'idoide.sspp.. Yes Yes'l A. t'eI a scoe nsi.s, B. ntitlttuye n.s ts abyssaminids. T. seI ttte rtsis, P. tt tne II i, agglutinants bulirninids (B. setn it ostu) 240 THOf'1AS

Appendix 12.2

Section/ LPTM Resolution 6llc Size Lithology Country Formation References Depth

NorthAtlarttir' High Yes >63 Marl' calcarenite A.l Spain Llmaya(Atlantic) Ortiz 1995;Canudo et al. 600-1500 turbidites 1995:Molina et al' 199'11von Hillebrandt 1965 High No >100 Limestones/marls A.2 Spain TrabakuaPass. Coccioni et al. 1994:Orue- 800-1000 Ermua.west Extebaniaet al in press Pyrenees Low/ No >150 Maris A.3 Spain 'l'remp/Campo Molina et al. 1992:Ortiz ?50-500 (Ilerdian stratotype) and McDougall l99l ; medium von Hillebrandt 1965 10-200 Low No >150 ClaYs/silts B France ParisBasin Le Caivez 1970

500-t500 Low No >150 Marls c Trinidad LizardSprings Beckmann1960; Bolli Formation et al' 1994 Low No >63 Clayeysands/ D.I usA New Jersey; Gibsonet al. 1993:Gibson 50-200 claYs Vincentown andBYbell 1995 Manasquan Formattons Low No >150 Silty sands/clays D.2 USA New Jersey; Olssonand Wise 1987 50-250 Vincentown Manasquan Formattons 100-500 Low No >150 Sand/clays E.l NW North Searegion King 1989 Europe 750-1000 Low No >150 Sand/clays 8.2 NW NorthSea region Gradsteinet al. 1994 Europe

Tetht's High Yes >63 Marls' limestone F Spain caravaca(Tethys) ortiz 19951Canudo et 500-1000 al. 1995

Low No >125 Marls/limestone G Israel NahalAvdat (Negev) Speijer 199'1 500-700

Low/ No >125 Marl/limestone H Egypt Abu Rudeis- Wadi Speijer1994 500-700 Nukhl (Sinai) medium

Low/ Yes >125 Marl/limestone I.l Egypr GebelAweina Speijer1994: Speijeretal. 100-150 1996a:Charisi et al' medium 1995:Schmitz et al' 1996 Low/ No/ >125 Marl/limestone 1.2 Egypt Bir lnglisi= Gebel Speijer1994; Speijer et al. 50-200 Duwi 1996b; rnedium Schmitzet al. 1996 Yes

J.lltalvBellunoDiNapolietal.lgT0500_1500LowNo>l50Marls/scaglias The Late PaleoceneBenthic Foraminiferal Extinction 241

Appendix12.2

Low O./ Postextinction/ Postextinction/ High .::-ExtinctionTaxa Short-Term Long-Term Productivity Dissolution

. utccariifonnis, Cibici.doides, First agglutinants,O. umbonatus, M truempyi,abyssaminids, Yes Yes - - rnrinids,P. con'elli IhenB. tu.rpanten.si.s O. ttmbonutus,buliminids

. ueccrtriiJbrmis,N. truempl.i. Rare agglutinants.then small A. di.ssonata,N. trttemp.vi, Yes Yes ,::lutinants,buliminids, buliminids.N. truempt'i buliminids.O. umbonattrs -,tnalinoidesv'elleri

. ,:rbigirtosus,Bulimina trigonalis, Cibicidoidesspp., Ut,igerina . ? '! : ,,r'telli, agglutinants spp.,Hanzuv'ttia spp

.. ,telphidiunt,pulsiphottina, Buliminids, Cibicidessuc'-. Yes? ? ;.t,idoides, buliminids cedens,Bolitintt spp.

- . r'lnscoensis,B. mitlwayensis, N. truempyi,T. seltnensis, Yes? '! -.,.rnrinids,Cibicidoides spp. B. semicostata(buliminids)

..:.elinella spp..Cibit:itloidas spp.. T. selmensis,Epistominellu sp.A T. selmensis,Pulsiphonina Yes Yes - : 'tualinoidesspp., Ut'igerina A., buliminids, Pseudouviger- prima, Arutmalinoid€.tspp.. ina spp. EPistominelln

:...elinellcr spp., Anomalinoides spp., Agglutinants,T. selmensis Epistominellasp., T. robertsi, Yes Yes - :':cidoidesspp. N. truempti

;.et.t'ttriiformis,B. mitlw'atensis, Low-diversity agglutinants T. brevispira,Gttudryina hilter- Yes Yes ' ' inincr trigonalis, Cibicidoides mcrzni,buliminids - .htntmina ruthyenmurrd)'i, Low-diversity agglutinants T. ntbertsi,R. amplectens, Yes Yes assemblage ' - ::, tilrrythragtniuntpaupera, R' intermedia ,..:nrblage(agglutinant) (agglutinant)

: ;,t'c't'ttriifonnis,A. t,elascoensis, RareHaplophrugmoides (barren), Buliminids, N. truempti, Yes Yes : ,.t-r.elli,Gt,roidinoides spp. T. selmensis,buliminids. Anomalinoidesspp. A. aragonensts

: ueccrrriifolnls,agglutinants, M truempt'i,Cibicidoides Yes? Yes : ,,nelli, buliminids, Cibicidoides spp. 'ii

: ht't.t.uriiJormis,agglutinants, (?) Anomolinoidesspp., vagin- T. truempti, Cibicidoidesspp., Yes Yes : .,,r't'elli,buliminids, ulinids,Stainforthia spp., T selmensis,buliminids .- "icidoides spp. buliminids .- iit'irloitles spp.,lagenids, C. mid- (?) B. callahani, Cibicidoidesspp.. Buliminids, Cibicidoidesspp., Yes Yes ,;rgrr.sis,A. avnimelechi A. aragctnensis.P. v'ilcoxensis Artornalinoidesspp.

:,:rtnalinoitles, spp.,Valvulinerla spp., Anomalinoidesaegrptai(us, Vtlt'ttlineriascrobiculata, Yes Yes -. ,.tttstomoide.sctpplinae Lenticulina spp.,L. applinae Cibicidoidesspp., Anomalin- oide.sspp., B. .foraJraens i s

:.:gltiinants, N. truempti, G. bec- I B. semicostata,B. trinitatettsis N. truempti, Anomalinoides Yes? ? .;rii.fctnnis,buliminids, A. velusco- spp.,buliminids ,':ili. P con-elLi 242 IHOXAS

Appendix 12.2 ( corttirtuetl )

Section/ LPT\I Country Formation Ref'erences Depth Resolution bl'rc Size Lithologl

J.2 Italy Possagno Bragact al. 197-5 500-1500 Lor',' No >l-50 Marls

K Austria Reichenhall von Hillebrandt1962 200-800 Lo',v No >l-50 Silts/marls SalzburgBasin

Pat'i.fic

L New Tawanui Hornibrooket al. 1989: -500-1500 High No >63 Siltstone.siltr Zealand Kaihoet al. 1993.in press limestone

M Japan Hokkaido.Tokachi Kriiho 1988 500 1000 Low No >63 Silty marls district The Late PaleoceneBenthic Foraminiferal Extinction 243

\ppendix 12.2 (c ont inue d )

Low O,/ Postextinction/ Postextinction/ High Pr.'-ExtinctionTaxa Short-Term Long-Term Productivity Dissolution

'i bc

\uglutinants.buliminids. nodosariids. Agelutinants.Anomalinoides ? ? -'nticulinids rubiginosus

,i. betcuriifbrnrl.s.agglutinants. First: rg.tu.rptunen.sis. agglutinants Cibicidoide,sspp.. O. umltona- Yes Yes t--.tnidwoyensis, A. ruhigirtosus Second:a-ggllrtinants only lrr.r,agglutinants

\ velleri, agglutinants.B. Agglutinants.buliminids Yes Yes

'l tt/r|dt'elists