t2
Biogeographyof the Late Paleocene
Benthic ForaminiferalExtinction
ELLENTH OIYAS
A B S T R A CT
During the Late Paleocene
ThermalMaximum (LPTM)
benthic fbraminif'era at rniddle bathl'al and greater
depthssufferedextinctionof 30-507cof speciesduling a
lew thousandycars.Extinctionwas lesssevcrcat neritic
to upper bathyal depths, where temporary changc-sin
launal compositionprevailed.Preextinctiondcep-sea
taunaswere cosmopolitanand diverse.and contained
heavilycalcifiedspecies.Imrnediatepostextinction
faunasweremorevariablegeographically,
exhibitedlow dil'ersity.and were donrinatedby thin-walledcalcareous
or agslutinatedtaxa, possibll,becauseCaCO, dissolution increasedglobally liom neritic to abyssaldcpths
just bcfbrethe extinction.Thcseassernbla-ges
weredontinatcd either by long-lived taxa sLrchas Nuttttllide.r
truerrtpt'iorby bulirninidtaxa.the latteraccompanicd
by
a-uglutinants
in somcareas.
Faunasdonrinatedby N. tmentpri rverecontntonin the
Sor.rthAtlantic and at lower bathl'al through upper
abyssaldepth in the lndian Ocean.and might inclicate
oligotrophicconditionsas well as incrcasedcorrr)si\L-ness.Buliminid-dominated
launasmight indicatehigh
ratesof depositionof organicmatteror lor.l-oxygen
conditions.Such llunas werc commonglobllly alongcontinental rnargins.and locallv co-occun'edu'ith sedimcntologic or planktonictaunalindicatorsof high procluctivitr.
In the bathyalcentralPacific.however.buliminid-dominated faunasco-occun'edwith planktonicfaunassuggestinsoli-sotl'ophi'.
and thev could reflectlow-oxygen
conditiorrs
lesultin-u
from sluggishoceancirculation.oxidationof clissociated
methanchydrates.or rlarming ol'
bathl'al irbyssalwaterscausedby a chan-ge
in deep-sea
circr.rlation.
Alternatively,they could indicatethat thc
tiaction ol'organic matter rcaching the seafl(x)rincreasedas a rcsult of decreasedoccanicoxygenf,tion.
214
The latestPaleocenebenthicextinctionthus was cornplex.and factorssuch:ischangesin deep-sea
circulation,
increasedCaCO. corrosiveness.
increasedlemperatures,
decreasedoxygenation.and changesin the patternsof
high productivitymay havecontributcdto its sevcrity.
E nvi ronmental
S etti ng
The late Paleoceneand early Eocenewere the warmest
epochsof the CenozoicEra: polar ice capswere almost
certainlyabsent.and shallowseascoveredlargepartsof
all continents.During theseepochs,major changesoccured in
climate and in the carbon cycle on
-elobal
scales of rnillions to thousandsof years. Long-term
warming of the deep oceansstartedin the mid-Paleocene E poch (Mi l l er et al . 1987a;S hackl et on1987;
Kennettand Stott 1990;Zachoset al. 1992. 19931Seto
199-5).Warm-water pelagic marine organismspenetrated to polar latitudesby the end of the Paleocene
Epoch (Stott and Kennett 1990; Premoli Silva and
B oersma 1984: B oersma et al . 1987: A ubr y 1992;
Berg-{ren1992).Thermophilicvertebratesoccurredin
the A rcti c (E stesand H utchi son1980:McK enna 1980t
Markr.r'ick1994).with vegetationand soil types indicating warrn clirnates(Kemp 1978: Nilsen and Kerr
1978:W ol f' e1994:B asi ngeret al . 1994:Manu m 199, 1) .
Clay mineral associations
in oceanicsedimentsat high
to midlatitudes indicate high humidity and intenscchemical weathering(Antarctic. Robert and Kennett
1992; N ew Jerseymargi n. Gi bson et al . 1993; New
Zealand.Kaiho et al. 1996).Oxygen isotopeclatasuggest that latitudinal temperaturegradientswere ver)/
shal l ow (S hackl etonand B oersrna198l : S t ot t et al.
1990: B arreraand H uber 199l : Zachos et al. 199, 1:
B ral ow eret al . 1995a,1995b;Lu and K el l er 1 995b) .
The Late PaleoceneBenthic ForaminiferalExtinction
Ratesof speciesoriginationand diversitywere high fbr
te n 'e s t r ialv er t ebr at e as n d fl o ra (H o o k e r 1 9 9 1 ;C o l l i nso n 1 983: Reaet al. 1 9 9 0 :W i n ge t a l . 1 9 9 1 ,1 9 9 5 ;Maas
e t a l . 1995) .as well a s fb r p e l a g i co c e a n i co rg a ni sms
(Ro m ein 1979:B oer s mae t a l . 1 9 8 7 :B o e rs maa n d Prern o l i S ilv a 199l: Co rfi e l d a n d S h a c k l e to n1 9 8 8 : M cGo w r an 199l; A ubr 1 , 1 9 9 2 1B e rg g re n 1 9 9 2 : C o rfi el d
1 9 9 3 :K elly et al. 19 9 6 ).T h e l a teP a l e o c e naen d e a r l i est
Eoceneepochsthus constitutea time of major innovati o n of t he bios pher e(Bri g g s 1 9 9 5).
The unusualwarmth of theseepochshas been conlrnonly explainedby high atmosphericpCO, concentrations. probablyciiusedby'platetectonicprocessessuch
as massivevolcanism in the North Atlantic Volcanic
Provinceduring initial opening of the North Atlantic.
of limestoneor oxidationof organic-rich
decarbonation
sedirnents
during the beginningof the India-Asia contincntal collision. and high hydrothermalactivity along
nridoceanicridges (see Thomas and Shackleton1996
tbr a review). Proxy data for atmospheric7rCO. levels
tenttrtivelysupport higher values than todav (Cerling
1 9 9l : F r eer nanand H a y e s 1 9 9 2 ).a l th o u g hth i s re m atns
debatable(Stott 1992).Clirnatemodelin-eindicatesthat
would be much
at suchCO. lcvelstropicaltemperatures
isotope
data. and
oxygen
higher than deduced fiorn
fiorn
heat
transport
incrensed
mechanismsfor highly
temperature
low
latitudinal
krr,i,to high latitudesat the
(Sloanet al. 1995:Sloan
gradientsremain r.rnexplained
a n d Rea 199-)5.
Epochthe world warmedeven
In the latestPaleocene
(Kennett
and Stott 199l: Pak and
more fbr a shorttime
1996).which has
and
Shackleton
Miller 1992:Thornas
Maximum
Thermal
Late
Paleocene
been named the
rvithin
occurred
This
event
TLPTM:Zachoset al. 1993).
Biochron
in
nannofbssil
C2:1r.
paleomagneticChron
NP9 (= Biochron CPt3),and befbrethe last appearance
datur-n (LAD) of the planktonic fbraminif-er Mr.,rrr(Aubry et al. 19961Berggrenand
.ot'ella veluscoen.si.s
Aubry 1996).Oxygen isotopedata indicatethat intermediateto deepoceanwatersglobally warmed by'1 to
6" C over lessthan a f'ew thousandyears(Kennettand
Stott 199l: Thomasand Shackleton1996).but tropical
remained
to subtropicalsurfaceto neritic temperatures
(
S
t
ot
t
1
9 9 5 b1Lu
a
l
.
1
9
9
5
a
.
Bra
l
o
w
e
r
e
t
1992
;
co n s t ant
1
9
9
6
).
T
h
i s rapi d
a
l
.
1995b
:
e
t
K
eller
S
c
h
mi
tz
and
bi' a
explained
has
been
ocean
of
the
deep
warming
durcirculation.
intermediate-water
in
deepto
change
interrnedideep
and
sources
of
the
dominant
ing which
ate waters shifted from high to subtropicallatitudes
(Ke nnet tand S t ot t 1 9 9l : P a k a n d Mi l l e r 1 9 9 2 :T h omas
a n d S hac k let on199 6 ).
Coeval with this short-termwarming was a shifi in
carbon isotope values by about -2'/rr, in surfaceand
deepwatersof all oceans(Kennettand Stott 199l: Pak
215
and Mi l l er 1992 Lu and K el l er 1993. 1995a,199 5b;
Canudo et al. 1995: Aubry et al. 1996: Thomas and
Shackleton1996: Schmitz eL al. 1996). superimposed
on a long-term decreasein values that startedin the
middle PaleoceneEpoch (Shackletonand Hall 1984;
S hackl eton1986. 1987:C orfi el d et ai . l 99l ; C orfield
and Cartlidge 1992 Corfield 1995).An isotopeexcursion similar in magnitudeand estimatedduration occurredin the tooth enamelof herbivoresand in carbonate concretionson land (Koch et al. 1992.1995:Stott et
organicmatter in a
al. 1996).and in terrestrial-derived
New Zealand marine section(Kaiho et al. 1996).The
a
reservoirthus undervu'ent
'uvholeatmospheric-oceanic
rapid.negativeshift in carbonisotopevaluesduring the
LPTM. the durationof which has beenestimatedat between 50 kyr (Thornasand Shackleton1996)and 200
kyr (K ennettand S tott l 99l ).
Massbalanceequationsshow that this carbonisotope
ercursion was so large that it probably could not have
been causedby transf'erof terrestrialbionlassinto the
system or by eruption of volcanoocean-atmosphere
genic CO.: it was so rapid that it probably could not
have beencausedby a changein depositionor eroslon
ratesof carbon in carbonateas comparedto carbon in
organicmatter(seeThomasand Shackleton1996.fbr a
review). This leaves f'ew explanationsopen. and recentlirit has beenspeculatedthat this unusual,negative
carbon isotope excursioncould have been causedby
massi vedi ssoci ati onof (i sotopi cal l yextremel yl i ght t
oceanicmethanehydratesas a resultof the deep-ocean
w armi ng(D i ckenset al . 1995:K ai hoet al . 1996).
B enthi c Forami ni fera
The latest Paleocenebenthic fbraminif'eralextinction
has been describedfrom land sectionsand at oceantc
drill sites. at depths ranging from outer neritic to
abyssal(appendi ces12.1 and 12.2:fi gure l 2.l ). This
extinction was rapid: it afl-ectedbenthic fbranlinif'eral
worldwide.and it occurreddurfaunascatastrophically
ing or close to the beginning of the LPTM and the
ne-eatir,'e
excursionin carbon isotopes.just before the
Epoch boundaryas most commonly
Paleocene/Eocene
defined(Thomas 1989. 1990b:Stott and Kennett 1990;
Nomura 199I in combinationwith Setoet al. 1991: Pak
and Mi l l er 1992;B ral ow eret al . 1995a,1995b;C anudo
et al. 1995in combinationwith Ortiz 1995:Aubry et al.
1996:Thomasand S hackl eton1996:K ai ho et al . 1996) .
This major break in deep-seabenthic foraminiferal
lbunas occuned between the Paleocene and Eocene
epochsif Viewedat coarsetimescales.whereasplanktonic foraminifera i.rnderu'enta major extinction at the
Period(Cushman1946;Beckmann
end of the Cretaceous
1960: von Hillebrandt 1962).After the late Paleocene
215
T HCI ' 1AS
described
inappendices
r2r andr22:nun,bers
(driilsites)and
rerters
[:1::':ii;i,,T::il]li:,:llljH;'il...,1jp,;T,i:::"""'as
Thc p a l e o g o
e g r . p h l r f t h e co nr in enr : in
r h e ,a tr 'p u r e ueen eEp o chi s afi cr Zach.s
benthic extinction faunas were predominantly
of lclw
diversitythroughoutthe bathyaland abyssal
regronsof
the oceitns. whereas coeval planktonic
organtsms
showed high rates of speciesturnover.
hi-eh rates of
specresori-eination.
and hi-ghdiversity(Shaciletonet al.
198-5;Boersma et al. 19137:Corfieia
ana Shackleron
1 9 8 8;B oer s m aanc lp re n to l iSi l v a l 9 9 l :H a l l o c k
et al .
l 9 9 l : A ubr y 1992;C o rfi e l da n c lC a rtl i d g e
l 9 9 l ; Cor_
l i e l d 1993;Lu and K e i l e r 1 9 9 3 .tS g Sa .tb q -rU :
K e l l y et
a l . 1996) .M ajor ex t i n c ri o n so f p l a n k to n i c
a n d b e n thi c
organismsin the oceansthus appearto
have been de_
co u p led( e. g. ,T homa s1 9 9 0 b :Ka i h o l 9 9 z l b ).
The rapidity of the extincrion makes it
difflcult to
compare rmmecriatepostextinctionf'aunas
at dif'fbrent
locations.especiallybecausemany sectlons
contalnun_
confbrrnitiesin the interval stracldling
the extinction
(Aubry et al. 1996).Only fbr
high_resolution
data sers
can_weknow that postextinctionfaunaswere
indeedco_
eval, becausewe can compareI'aunasthat
co_occumed
with the short-termnegativecarbonisotope
event.Fau_
nal comparisonsmay be more difficult because
of dif,f-erences
in taxonomicconceptsof differentauthors
and
becauseof the differencein size fiaction
being studied,
varryingbetween63 and 150pm in most
."r.i, (opp.n_
d i ce s l2. I anc l12. \ .
High-resolution faunal data and carbon
rsotope
analysesare available fiom Ocean Drilling progritm
et al . ( I 99.r).
(ODPI Sites 689 and 690 (Wed<tell
Sea: Thomas and
Shackleron1996).SouthAtlantic (Walvis
Ridge) Deep
S ea D ri l l i ng proj ect (D S D p) S i tes
525 and 5 27
(Thomasand Shackleton1996).
and equatorialpacific
ODP Site 86-5(Bralower er al. 1995a.199-5b:
Thomas
unpublisheddata).Detailed data are also
availablefbr
the Zurnaya and Caravaca land sections
rn Spain
(Canudoet al. 199-5;Ortiz 1995)
and fbr rhe .Iawanui
sectionin New Zealand(Kaiho et al. 1993,
1996).The
short-termcarbonisotopeanomalyhasbeen
recognizecl
in neritic Egyptian secions (Schinitzet
al. 1996.),but
benthicfbraminif'eraldata have not yet been
published
resolution(Speijer 199,1;Sperjerer al. 1996a,
ij.llgh
Isoroperecordsfor ODp Site Z:g re-ristered
1?.96b).
the
6rrC excursion(Lu anclKeller 1993).but
o-ntytimitea
dataron the benthic f'aunasare available (Nomura
and
Kennett. personalcornmunication1995:
Thornas.un_
published data). Faunal data fbr equatorial paciflc
DSDP Sire -577and Bay of Biscay DSDP
Sire,l0l (pak
and Miller 1992)probablydo not cover
the time inter_
val directly after the extinctionbecausethe
rnagnrtude
of the 6lrC anomalyis much smallerat these
sitesthan
at others,suggestingthat its minimum value (and
thus
the detailsof the benthic fbraminif-eralextincrron)
oc_
cured betweensamples.
Most authorsconcludedthat a changein
cleep_water
circulation played a major role in the extinctron.
that
The Late PaleoceneBenthicForaminiferalExtinction
't o1'the deep to intermediatewaters of all oceans
'::rcciat least fbr some time at subtropicallatitudes.
-l that decreasedlevels of deep-seaoxygenationre.rnc fiom the increasedtemperatureof thesewaters
::.J rl c aus olf ac t or ( e .g ..Mi l l e r e t a l . 1 9 8 7 b :T h o m as
-.9 1990b, 1992' .K a tz a n d Mi l l e r l 9 9 l ; Pa k a nd
'.l l cr 1992, \ 995 K a i h o 1 9 9 1 .l 9 9 z l b :N o mu ra l 9 9 l :
,i ro r r i et al. 1991,O rti z 1 9 9 5 ;Sp e ji e r e t a l . 1 9 9 6b).
r-.1-r\.
however.suggestedthat the patternsof f'aunal
, .nctionvary geographicallyand that their complex'.
only
-'.lnnotbe explainedby decreasedoxy_senation
:,:.. N om ur a 199l: C o c c i o n ie t a l . 1 9 9 4 :T h o rn a sa nd
. ,,.'k let on1996) .
r:r this chapter.I first erplore postextinctionbenthic
:',:rninif-eral
fhunal patternsfbllowing the extinetion
.:rg h igh- r es olut io nd a ta l i o m th e s i te s m e n ti o ned
- re . T hen I us e lit e ra tu red a ta (fi g u re 1 2 .1 .a p p e n. -:. ll. l and 12. 21t o try to e v a l u a teo c e a n i ce n v l ron.:rt;rl changes between late Paleoceneand early
-.ne times.to deterrninethe biogeographicpatternof
: :rtinction. and to speculateon its possiblecauses.
Me th o ds
Srn p l e P r epar at iona n d Stra ti g ra p h i cF ra me w o rk
':. H i g h- Res olut ionS tu d y
. . iirs chapternew data are presentedfrom ODP Site
'-5. and a f-ewadditionalsanrplesfrom ODP Sites689
.i 690 and DSDP Site -527lfigures 12.2 and 12.3),as
, :rp a r ed t o dat a in Th o ma s a n d S h a c k l e to n(1 9 9 6).
\.,rrples fbr benthic fbraminif'eralfaunal and isotope
. ..:lrsis were taken fl'om DSDP Site 527 (Walvis
:,.Jge. SouthAtlantic), ODP Sites689 and 690 (Maud
< .c. Weddell Sea).and ODP Site 86-5(equatorialPa, 1r t. They were dried overnightat 50" C and weighed.
:.:n soakedovernightin distilled water.Most samples
, 'rrggregatedreardilyand could be washedover a 63,:: screen.Benthic fbraminifera fbr fhunal analysis
:r'e picked fiom the >63 prrnsize fiaction. fbllowing
lh,rmas ( 1990a). All specirnenswere picked and
': ,rLrnted
in cardboardslides.All samplescontainedsuf:
s
pec
im ensf or a n a l y s i s(> 2 5 0 ).T a x o n o m yi s a s
-.i e n t
:: Thorras( 1990a)and Thon.ras
and Shackleton( 1996).
,,:rdlargely fbllows van Morkhoven et al. (1986). The
richnesswas usedto calculatespeciesrichness
-,rr'Cies
r a sample of 100 specimens usin-s rarefaction
Sa n d er s1968) .
I use the integratedmagnetobiochronologic
fiame'.r.rrrkof Berggrenet al. (1995), which is basedon the
:eomagnetic polarity tirnescaleof Cande and Kent
1 9 9 2)and m odif ied b y C a n d ea n d K e n t (1 9 9 5 ).For
ODP Site 690, I use the biostratigraphicinterpretction
,i Ar-rbryet al. ( l996). givin-ean age of -55.-5
Ma to the
2|7
benthic extinction event and 55 Ma to the NP9/NPlt)
biochronalboundary.I derive numericalagesfbr ODP
Site 690 assuming constant sedimentationrates between the benthic extinction and the NP9NP10 zonal
boundary.and betwcenthe benthic ertinction and the
top of Chron C25. For samplesfiom DSDP Sites ,525
and 527 and OD P S i tes689 and 865 I assi gnnumeri cal
agesaccordingto Ber-ugren
et al. ( 199,5)to daturr levels
i n Thornaset al . (1990).S hackl etonet al . (198.1).and
B ral ow eret al . (1995b).Fi nal l y.I revi sethe numeri cal
ages at thesefbur sitesuntil the benthic fbraminifcral
carbonisotopecurvesagreewith that fbr ODP Site 690
(ThornasanclShackleton1996)in order to correctoptimally for .",arying
sedirr.rentation
rates.especiallyfbr the
dissolutionzone at Sites525 and 527 and fbr the interratesjust abovethe exval with very low sedin.rentation
ti ncti onat S i te 86-5.
Selection of Literature Data Sources
A ppendi ces12.1(D S D Pand OD P si tes)and 12.2(l an d
sections)list sites fbr which data are available that
straddlethe extinction level: hence only a lew sitr's
fiom T.jalsrna
and Lohmann( 1983)are included(figure
l 2.l ). D ata on secti onsi n the N orw egi anGreenl and
Sea and Labrador Sezrwere nclt included becausein
n.ranyof the sectionssedirnentationprobably started
afterthe exti ncti onevent(FIul sboset al . 1989:K arri nski et i.rl. 1989). Data on agglutinateclfaunas were
mostly not included. e\ren thou-ehthey suggestthat
changesclccun'edin agglutinatedfaunasat the end of
the PaleoceneEpoch (e.-s..Geroch and Novak 1984:
K uhnt and K arni nski1990:K ami nskiet al . 1996).be causeexactdataand precisecorrelationsare difficult in
the absenceo1'carbclnate
and thus of isotopedata.Data
fiom the North Searegionare includedbecauscdetailed
stratigraphicdataare available(Gradsteinet al. 199:l).
Ferv paperson neritic faunas are included because
benthicfaunasat thesedepthsare more geographically
variableand planktoniczonationis con.rmonlydifficult:
therefbre.it is not possibleto detenxine whether observedtaunal changecan indeed be correlatedto the
late Paleoceneextinctionin the deep sea.For instance.
paperson neritic to brackish water Paleogenefaunas
fiom the northern North Atlantic were not included
(Berggren1974a:Murray 1989).Dataon neriticsections
in E-eypt(Sperjer199,1;Speiieret al. 1996a.b)were included becausedetailedcarbon isotopedata are availabl e (C hari siand S chmi tz 1995:S chmi tzet al . 1996 ) .
Data trorr.rSpain (Molina et al. 1992 Ortiz and McDougall I 991) were tentativelyincludedbecauseplanktonic fbrarniniferaldatasuggestthat benthicextinctions
were at leastclose in time to the extinctionin the deep
sea. Data fiom coastal New Jersey were tentatively
2I 8
T HOIY AS
N. truempyi, "/.
10
20
30
bi/triserial species, 7o
(infaunal morPhotYPe)
80
40
60
20
o
o
-
G aveli n ell a beccariif orm i s, 7"
5
numberof speci es(100)
40
50
30
20
55.35
55.40
55.45
o
55.50
55.55
55.60
Figure 12.2 Contparisonof the spcciesrichnessand relati"'eabundanceof significanttaxa at the time ol'the late Paleocenebenthic lbranriniferalextinction.Seetext firr clerivationol'thc'age moclel.
anclShacklcton
DatafirrDSDPSites515and517andODPSites6lJ9and690allcrThotttrs
689.and690:datafbr ODPSite865lra\enot1ctbcenpublished.
inc luded( O lss o na n d Wi s e 1 9 8 7 ;G i b s o n et al . 1993:
Gibson and Bybell 1994) becausethe clay mineral
record at thesesitesmay be a meansof correlationto
ODP Site 690 (Robertand Kennett 1994).
Results
H igh- Res olu ti o nS tu d i e s
A t O DP S it e s6 8 9 (p a l e o d e p thl l 0 0 rn ) a n d 690 (1900
m), DSDP Sites -525(1600 rn) and 521 (3400 m). and
O DP S it e 865 (1 3 0 0 -1 5 0 0m ) b e n th i cfo ra n i ni l -eral di versity dropped precipitouslyat the extinction level.
of speciesbecoming extinct (figures
with 35 to 5OC/c
12.2 and 12.3).This percentageis a minimutn estimirte
becauseseveral nodosariidspeciesare rare, so their
rangescannotbe determinedwith accuracy.Some nodosariidsare absentin the intervalconespondingto the
LPTM, but reappearhigher in the section at varyinq
levelsthroughoutthe lower Eocenesection.
At both Walvis Ridge sitesthe extinctionoccursjust
above the base of a level with severedissolution.although benthic fbraminit-erawere well preserved:
planktonic tbraminifera show severe dissolution and
fiagmentation(Thomasand Shackleton1996).No lowof the sedimentoccuned
ering of the CaCO, percentage
at Sites 689 and 865. but faunas at these sites show
more etchingand are lesswell preservedjust abovethe
The Late PaleoceneBenthicForaminiferalExtinction
BFAR (nr/cmZky)
num berof s pec i e s(1 0 0 )
5oo 1000 1500
20 30 40 50 60 o
35
Cibicidoides, %
buliminidspecies,7o
30
20
10
75 0
50
25
0
219
N. truempyi,%
10 20 3 0
40
45
CJ
c)
55
3a
JC
60
Figure 12.3 (a) Speciesrichnessat 100 specimens,(b) benthicforaminiferalaccumulationrates(BFAR). and relativeabundances
spp..and (e\ Nuttallidesrruentpt'iat equatorialPacificSite 865.
Lrf(c) buliminid taxa.(cl)Cibititloicle.s
to C--ande
andKcnt( 1995): agernodelatierBralouer et al. ( I 995a.I 995b)
\urnericalages accurding
extinction level. At Site 690 similar slight dissolution
occurs in the faunas,and CaCO. valuesdecreasejust
below the extinction level (O'Connell 1990; Thomas
t992).
At all sites.Gavelinellabeccariifonnlshas its uppermost occurrenceat the extinctionlevel of severalother
taxa (seebelow). Our data confinn the observationby
Ka tz and M iller ( 19 9 1 )th a t th i s s p e c i e si s mo s t c ommon at higher-latrtudesites: it never reachesabundanceshigher than about 5% at equatorialPacific Site
865. Other speciesthat have their highestoccurrenceat
this level includeseverallargeagglutinatedspecieswith
calcareouscement (Dorothia ox\:('otto.Tritario lturartensis. and Trita.ria pttleocenicci) and calcareous
Bolit'inoide.s
speciessuch as Arugortictt,elasr:oen.sis.
thatrcteBulimina
Bulimina
mitlvruven.si:;.
tlelit'rttulu.s.
hillebrandti.
Neoeponitles
reticulctttt.
Neoflabellina
nis,
Neoeponicleslmntu, and Pullenia c'on'elli.
Immediately above the extinction level, speclnlens
are smaller and have thinner walls than below. even
specimensof speciesthat straddlethe extinctioninterval (e.g., Oridorsali.s umbonatus and Nuttallitles
tuempti). At ODP Sites 689 and 865 the change to
smaller and thinner-walledtaxa occursa f-ewcenttmeters below the highest occurrenceof G. bet'ctrriifitrni's'
possiblybecauseof bioturbation. Tappanirnselmertsis
andAragonieraragonensisare at all sitesmore common
abovethe extinctionlevel than below (as also observed
by Boersma 198,1b:Speijer 1994), but their relative
abundancesvary strongly between sites.Tappcutinasclrnensisreachesup to 40% at the Maud Rise sites.up to
l5c/cat the Walvis Ridge sites.and up to 25Vcat Pacific
Site 865. At all sites.peak occurrencesof T. selmensis
occur directly abovethe extinctionlevel, whereasthose
occur at the top of the LPTM interval.
of A. ctrttgorten.rl.s
Despite these similarities,postextinctionfaunasare
most notable for the major difl'erencesbetween sites.
especially in the abundance patterns of buliminid
species,Ntrttallides truempti, Oridorsali.s ttntbottatus,
Cibit'idoidesspp., and abyssaminidspecies(Thomas
and S hackl eton1996;appendi x12.1).The postext inction faunas diftbr much more from site to site than the
preextinctionfaunas,which are comparableto Velascotype f'aunas at all five sites (see also Berggren and
Aubert 1975;Kaiho 1988).Thesepostextinctionf'aunal
differencescannot be completely attributed to the presenceof stratigraphicgapsand comparisonof noncoeval
f'aunas.becausefaunasat all five siteswere coevalwith
the short-term carbonateisotope excursion. Data were
so there is homogeneityin
collectedby one researcher.
the databasewith regardto taxonomicconceptsand size
fiaction. Postextinction f'aunas differ rnore by geographic region than by depth; f'aunas fiom the two
Walvis Ridge sites and the two Maud Rise sites are
in depth,
quite similar despitelargedifl'erences
At the Walvis Ridge sites.postextinctionfaunasare
donrinatedby lV.truempt'i, O. urnbonatu's,and abyssaminid species:the latter are especiallyabundantat the
22O
I HO I T A S
deepestsite (figure 12.21.Abyssaminids(.Abt"tstttrtitttt
pottgi. Abt'ssrttrtitttr
clutrdrorrl.and Clinttpertinupluni.qrriz) increasein relative abundancepostextinetionat
MaLrd Rise. bLlt much less than at the Walvis Ridge
sites.At Pacific Site 86,5these specieshave very low
throughout,with a slight furtherdecreaseat
abundances
tlre extinction level. Nuttallitles tntemPti declines
stronglyin relativeabundanceat andjust abovethe extinction level at the Maud Rise sitcs as rvell as at Site
865. in stron-qcontrastwith the situationin the South
Atlantic.At the Walvis Ridge sitesthe postextinctionly'.
tnternpt'ispecirnensare smallerand have thinner '"r'alls
than preextinctionspecirnens.but are much more numerousthan in preextittctionfhunas.
of the
At the extinctionlevel the relativeabur.rdance
not
does
at
Site
527.
decreases
buliminid species
-eroup
Maud
at
both
strcxgly
increascs
but
changeat Site 525.
Ris e s it esand S i te 8 6 -5(fi g u re 1 2 .2 ).Ab o v e t he exti t.tction level this group is dorrinatedby snlirllspeciessuch
sp. cf . B. decoralc, and
as T. seltnensi.s,Bolivinrtrrft'^s
Rise sitesSiphttgeneriAt
the
Maud
sirnple.r.
Bulinina
(up to 40%) acrossthe
is
contmon
noicla.s
ltrevi.sltittosu
W
a
l
v
i
s
R
i
c
l g es i te si t i s nl uchl ess
th
e
lev
e
l
,
a
t
ex t inc t ion
rs
(
up
S
i
te
8 6 5 i ts a b u ndance
l
-5
%
).
a
n
d
a
t
to
c onr non
its
has
species
br-rt
the
Walvis
Rid-ee.
at
similar to that
lrighest occun'enceat the extinction level. Bulintirttt
is abundantat Pacitic Site 865 iust above
,semitostcrtcr
but rein abr-rndance
the extinctionlevel.then decreases
Eocene
middle
and
the
lou'er
present
throughout
mains
Series.At the Maud Rise sitesthis speciesappearscl.rly
in Z one NP l2 (5 1 -5 2 M a . B e rg g re ne t a l . 1 9 95).In the
lcrwermostpostextinctionspecirnensof B. setrtit'rtsttrtrt
at site u65 the costaeon the lower part of the test.typical lbr this species.are unusuallvthin and the speciat this
rleus are srnall;possiblythe speciesori-einated
trihedralBulitninuspecles.
time from a noncostate.
o f b u l i m i ni d speci es
A t S it e U65 th e p re d o mi n a n c e
appearsto have lasted fbr a shortertin.rethan at the
M aud Ris e s ite s .At th i s s i te th e i n te rv a lo f bul i rni ni d
about l 0
is o n l y l 5 c n t th i c k . re p re s e n ti ng
c lom inanc e
kyr in our age rnodel.whereas it is about-500cnl thick.
representingabout 100 kyr. at the Maud Rise sites.
Above the buliminid-dominatedintervalat this site fallspp. up to a level in
nas are dominatedby Cibit-idoirft'.s
Z one NP I I ( z o n a la s s i -s n m e innt B ra l o w e re t al . 1995b;
52. U5 53. 61 Ma i n B e rg -e reent a l . 1 9 9 5 ;fi - eure12.3)'
increases
where the relative abundancet'tfN. trLtemp.r'r
to l2-25c/c.At noneof the othcr sitesis therean interval
dorrrinatedby Cibicidoirle.ispecies(up to 307c). Ihe
taxonomyof theseCibicitloitlesspeciesis still underinvestigation:they resemblelar-ee.flat shallow-waterEuropeanspeciessuchas C. pntprius (Brotzen 1948).
A detailedrecordof planktonicfbraminif'eraand isotonesis availablefrom southernIndianOceanODP Site
738 (Lu and Keller 1993),but the benthicfbrarninif'eral
record is limited. lmmediatepostextinctionfaunasare
very poorly preservedbecausethe extinction occurs
r,r,'ithin
a layer rvith very strongdissolution.I observed
very commonsmal l .bul i rni ni dspeci es(i ncl ud ingI . ie1just abovethe extinctionlevel.
ntensi.s
and ,8. .simple.r)
suggestingthat the faunal patternsat this site resen-rble
thoseat Maud Rise.
Detailed clata on faunarsu'ith age control by thc
LPTM carbon isotope excursionare availablefirr the
et al.
Zur-nayaand Caravacasectionsin Spain (Canr-rdo
1995: Orti z 199-5;appendi x 12.2). In both seet iot r s
stron-gdissolutionoccllrsat the extinctionlevel.as indicated by thc presenceof dark gral'. larninatedshales'
Faunasjust above the extinction level of G. beccatiifttnnis are dominatedby the agglutinatedspeciesHaTrlophnrgrrtoitlesretrosept(tAt Zunaya. above the dissc'lcommot.t
lution interval there is a thin interr"alr,'u'ith
Bulinrirtcrtu.rpumettsi.;(Bulitnirttt spp. Assernblage):
higher N. tntempt'i is dorlinant. At Caravaca ,V.
truemltvi is not as comrnonas at Zumirya,and it is acand Oritlorsuli's
cctpitcrttt.s
corrpanied by Anornttlinrtitle.s
The Bulinina spp. Assenrblagen.rayhave
ttmbortttttts.
persistedsontewhatlonger at the someu'hatshallower
but fbr this part of the
sectioltthan at Zut.naya.
Carar.'aca
section age control is not as precise,and short-terrn
ratesnlay obscurethe pattern.
changesin sedimentation
In the Tawanui Section(Neu' Zealand;Kaiho et al.
1993, 1996;appendi x12.2)the benthi cexti n ct ionoccurs in larninated.dark gray.organiccarbon-richshales.
Age control is provided bv recogniticlnof the LPTM
6lrC excursionin terrestrialor-uanicmatter.Within a
.l-cm intern'aldirectly above the hi-ghestoccurrenceof
taunas are dominatedlry Btrlintitttt
G. bet'cctriift,nrlr.i
pupoides. Above
ttr,tltcntensisand Prtteglrtbobulimirtcr
the Buliminu spp.Assernblagethereis an -S0-cm-thick
interval with r,'erystrong dissolution.in which faunas
spp..
taxa (Rhubdutntrtirr.r
are dominatedby a-eglutinant
depres.ra.and Reoplta.rregularis\.
Cottotntchtunntirtu
Higher in the section faunas arc lnore diverse. with
spp.. Arnntttlinoides spp.. and
conlmon Cibic'idoitle.E
B. Iurpanren.si.s.
studies
We can concludefiom thesehi-sh-resolution
by low dithat postextinctionf'aunasare characterized
t,ersity.and that geographicditterencescannot be attributedto insufficientage control.difl'eringtaxonornic
With this in
concepts,or differing sizefracticltlsstr.rdied.
mind. we can try to evaluatethe lower-resolutiondata
availablein the literature.
Literature Data
Beckmann(1960) was the first authorto recogni/e the
imoortanceof the benthicfbraminit'eralertinctionat the
The Late PaleoceneBenrhicForaminrferalExtinction
end of the PaleoceneEpoch. basedon his study of the
hathyalLizarclSpringsFormation(Trinidad).His clara
ta s giv en in B olli e t a l . 1 9 9 ,1 s) h o w a v e ry s tr.o n-{
i n_
creasein highestoccurrencesas well as a decreasein
lowestoccurrences
of benthicforan-rinif-era
in the Lrpper
PaleoceneMoro:.ot,ellavelct,st'oensis
Zone. close to the
Paleocene/Eocene
Seriesboundary(Bolli et al. 199.1).
Benthicfbraminif'eralfaunasin the overlyin-s,:l
zonehat'e
,rnruchlower dil'ersity.Thereare more lastappearances
,)t agglutinated.
noclosariid.
and rotarliidtaxathan of bul rrn inidt ax a ( f igur e 1 2 .4 ).A n o th e re a rl yre c o g n i ti onof
ihe irnportanceof a late Paleocene
benthicfbrantinif-eral
3 \ti nc t ion ev ent wa s b y v o n H i l l e b ra n d t(1 9 6 2 ).w ho
lr)ticed a strongdrop in diversity of benthictbraminiierirl faunastoward the top of his M. t'elct.scoen.sis
Zone
Zone F) in the Reichenhall-SalzburgBasin (figure
i l.-5).In thesesections.however.rotaliid taxa appearecl
', h e t he nt or eeot nn to n\u rv i \ r)rs .
The irnportanceof the benthic fbraminif-eralextinci.n was recognizedby many later authors(appendices
1 .1 and 12. 2)in s t u d i e so f l a n d s e c ti o n sa s w e l l a s of
DSD P and O DP s it e s i n a l l o c e a n s .H i g h -re s o l uti on
studiesdemonstrate
that the extinctionwas a very shortterm event, so contparisonof faunal events between
sites is problernaticfbr data that are so disparrte in
chronologicresolution,a-qecontrol.taxonomicconcepr.
and size fiaction studied,as presentedin the appendices.However.someoverallconclusionscan be drawrr
fion.r these diita. especiallv in contbinaticlnwith the
high-resolutiondata.
In the first place.the extinction\\'asse\ereat midclle
bathyat through abyssal depth. with extinction of
speciesran-eingbetween30 and -507c.
At upperbathyal
through neritic depthsthe ertinctions were much less
severe.but faunalcompositionchangesr.verehighly significant.Thesecorrpositionalchangeswere temporiiry
and many speciesreturnedto the sites (Speijer 199:t;
Gi bsonand B ybel l 199,1).
Second. preextinction Paleocenefaunas were remarkably cosmopolitan,extendingover a wide depth
range(Brotzen 1948:Berg-uren1911a.lgllb: Ber-ugren
and Aubert 1975). The larter authors distinguishecla
continentalshelf fauna(Midway-type)and a lou,ercontinental slope and abyssalplain fauna (Velasco-tvpe).
firstappearances,
% of totalspecies
last appearances,
% of total species
20
=
20
P7
P7
P6b
P6b
P6a
P6a
P5
P5
P4
P4
P3b
P3b
c0
o P3a
c
o
221
benthicextinctiori
P3a
N
P2
P2
P1c
P1c
P1b
P1b
F-igure12.1 (a) Lorvestand (b) highestoccurrences
of speciesbelongingto the four most important
of taxa as a percenta-ce
-sroups
of t he tota l nu nrb ero f spec ies int heLiz ar dSpr ingFor r n a t i o n . T r i n i d a d ( B e c k u r a n1n9 6 0 : B o l l i e t a
l . l g g . l t . T h e h o r i z o n t a ll i n c
showsthe level of the bentliicextinction.
222
T HO IY AS
a^
oo
+
+
aggl utrnants
nodosari i na
+
+
bul i mi ni dea
r otal i i dea
:O
o
!^
5N
C
(9
L
to
<
rogozl
,on"luonnirroeor"not
Figure 12,5 Numbersof speciesof the four mostinrporlant
groupsof speciesin the Reichenhall-SalzbLrr-e
Basin(von
Hillebrandt1962).The verticalline showsthe levelof the
benthicextinction.
although several species(e.g., Bulimina ntitlv'atensi.s
and Con'phostonn midv'avensr.r)occur in both faunas
( e. g. .K aiho 1 9 8 8 , l 9 9 l : N o m u ra 1 9 9 1 ; K ai ho et al .
1993;Sperjer199.1:Coccioniet al. 1994).The Midway
Ihuna typically contains more Cibic'idoirle.iand espectally Anonuilinoide.s species, whereas the Velasco
fiiuna has A. velu.sc'oensis,
Gyroidinoide,sspp.. N.
truempti. and agglutinanttaxa such as Dorothia o.tt'contt, Guudn'ina lnev'igata, Trita,rio lnvanen.si.s.and
Tritcr-riupuleocenico.The common speciesN. truenrytti
generally is consideredtypical fbr the Velascof'auna
(below about -500n). Tapparina selmensisoccurs fiom
the shelf (about 100 m) down to abyssaldepths.but is
rarein bathyal-abyssal
faunasbelow the level ofthe extinction. Both Midway and Velascofaunasare characterizedby extremelyhigh diversities.the occurrenceof
many rare taxa (especially nodosariids).and thickwalled calcareoustaxa.
Third. there were geographicdifferencesin faunal
assemblages
in spiteof the cosmopolitanoccurrenceof
many species.Gavelinellubecc'uriifrtrntis,
fbr instence,
appearsto be more common at high-latitr.rdethan lowlat it udes it es(Ka tz a n d M i l l e r 1 9 9l ). a l th oughi t w as
alsocommonin the Tethys(Speijer1994\.Arugonia yelascoen.sis
is extremely rare at both Maud Rise sites.
whereasS. breyi.spinosais abundantat these sites. but
showslargefluctuationsin relativeabundance(Thomas
1990a. 1990b).The geographicdifferenceshave not
been well evaluated;the availability of more oceanrc
drill sites in depth transectswould help in separating
depth frorn geographiceffects. and in distinguishing
Paleocenebiogeographicprovinces,possiblysimilar to
thosein the latestCretaceousPeriod(Widmark 1995).
Fourth. this high degreeof faunal cosrnopolitanism
endedwith the late Paleoceneextinction(Kaiho 1988;
Thomas 1992:Thomasand Shackleton1996).This ap-
parent difference in provinciality between late Paleoceneand early Eocenefaunasmight be thoughtto result at least partly frorn the lack of a monographon the
early Eocenebenthic fbraminiferal taxonomy comparable to that of Berggrenand Aubert (1975) for the late
PaleoceneEpoch. so that there are greaterdifferencesin
taxonomicconceptsbetweenauthors.This cannotbe the
full explanation,however.becausegreaterdifTerences
between early Eocene faunas fiorn diffbrent locations
have beenobservedin faunasstudiedby one investigator (Tjalsmaand Lohmann 1983:Kaiho 1988;Katz and
Mi l l er l 99l ).
Finally, postextinctionfaunaswere of low diversity
and high dominance,in starkcontrastwith the highly diverse.low-dominancepreextinctionfaunas.The postextinction community thus hiis erstructuretypical for faunas afTectedby a major environmentalperturbance(e.g.,
Pearsonand Rosenberg1978).In theseperturbedcommunities the speciesT. selntensisandA. (rragorren.ri.s
are
comrnon to abundanteverywherebelow shelf depths
(appendi ces
12.I and 12.21B oersma1984a).
Otherwise. f'ew thunal observations are valid on a
worldwide basis.The postextinctionpatternas seenby
Tj al sma and Lohmann (1983) and K atz and M iller
(199l). i.e., dourinanceby 1/. tntempt'i with survivor
speciessuchas O. umhrnatu.r.appearsto be typical for
the South Atlantic and possibly Gulf of Mexico;
abyssaminidtaxa are common below about 2-500m in
this type of assemblage.
Thomas (1990a. 1990b) suggestedthat this associationmight be seenonly in studies
usingthe largesizetiaction (>150 pm) becausemany of
the buliminid taxa are small. High-resolutiondata in
Thomasand Shackleton(1996) and this chapter(figure
12.2) show that this is incon'ect.and that faunas are
dominated by N. truenrpli and abyssaminids even
though the >63-pm fiaction is studied.as was also observedby Mtiller-Merzand Oberhiinsli( 1991).
Patternsin the North Atlantic are more variable. Most
North Atlantic data are from the Goban Spur-Bay of
Biscay Sites,and postextinctionfaunasvary from buliminid dorninatedto N. truentpti dominated.Possibly,
the two types of faunascan be differentiatedby depth:
Buliminid-dominatedfaunasappearto occur at rniddle
bathyaland shallowerbathyaldepths(DSDP Sites5218,
549; Reynolds 1992; Boltovskoy et al. 1992),whereas
N. truentpt'i-dominatedfaunas occur at lower bathyal
throu-ehabyssal sites (Schnitker 1979 Pak and Miller
1992;appendixI 2. I ). All thesesites.however,show increaseddissolutionand occumenceof unconformitiesin
the lower Eocene stratigraphicrecord. which makes
faunalcomparisonsmore diflicult (Aubry et al. 1996).
Bathyal land sectionsclose to the Bay of Biscay
(C occi oni et al . 1995; Orti z 1995) show b ulim iniddominatedpostextinctionf'aunas,although the pattern is
The Late PaleoceneBenthic ForaminiferalExtinction
corlplicatedby the occurrenceof agglutinatedtaunasin
an intervalof severedissolution.In New Jerseymargin
DSDP Site 605 N. trtrentpti-dominatedfaunas occur
above the extinctionlevel (Hulsbos 1987;Thomas unpublisheddata).Shelf faunas(Olssonand Wise 1987:
Gibson et al. 1993) in the region show common Epistominello,Pulsiphonina.and Tunilina, and cannoteasily be comparedto the deeperfaunastcoastalsections
generally have unconfbrmitiesat the extinction level
(Gibson and Bybell 1994).The postextinctionfaunas
fiom Trinidad are buliminid dominated (Beckmann
1 9 6 0 :B olliet al. 199 4 r.
In conclusion,we can speculatethat the northern
Atlantic abyssalto lower bathyal faunas were similar
to South Atlantic faunas (N. truempv-dominated).
whereasthey were buliminid dominatedin the upper-to
mid-bathyalreaches.We cannot be certain. however.
that we are indeed comparing coeval faunas because
tl i sso lut ionand unc o n fo rm i ti easrec o mmo n .
The bathyal faunas in the North Sea (depths
750-1000 m) were dominatedby noncalcareous
agglutinated taxa through the late Paleocene and early
Eoceneepochs(Charnockand Jones1990;Gradsteinet
al. 1994).These agglutinatedtaxa underwenta period
of severe ertinction close to the Paleocene/Eocene
Epoch boundary.after which extremelyimpoverished.
low-diversity. ag-tlutinant faunas remained (e.9..
C h a r noc kand J ones1 9 9 0 ;K i n g 1 9 8 9 :G ra d s te i ne t al .
1994).Direct postextinctionsedimentsindicate stron-q
CaCO. corosivenessand dysoxic to anoxic conditions
(Charnockand Jones 1990;Schroeder1992).At somewhat shallowerlevels(100-500 m) calcareoustaxa (including G. beccuriifonzl.i) were present durin-e the
PaleoceneEpoch (King l9u9). These faunaswere replaced by extremelylow-diversityagglutinatedf'aunas
similar to thosefrorn deeperregions.The timing of the
disappearance
of the calcareoustaxa is difficult to establishbecauseof the biostratigraphrc
complexityof the
North Sea region (Gradsteinet al. 1994;Berggrenand
Aubry 1996).The local North Sea disappearance
of G.
probably
predate
global
not
exbecc'uriiJonms
did
the
tinction ol C. beccariijormis at the LPTM by very much
time becauseit can be correlatedto the lower part of dinoflagellateZone D5 (King 1989).which has beencorrelated to just below the middle part of nannofbssil
Zone NP9. Palynologicalstudies indicate the occurrenceof a peak warm period somewherein dinoflagellate Zone D-5(thus Zone NP9; Schroeder1992).which
might possibly be correlatedto the LPTM in midBiochron NP9 (Aubry et al. 1996).
Tethyan faunas are known fiom land sections only
(appendix12.2).Sectionsin the easternTethys(Speijer
l99ul)from a depthrangeof 500-1000 m show postextinction faunas with common N. truemot'i.Anomali-
223
noides species, and Cibicidolrle.s.Shallower faunas
(100-300 m) containmore buliminids,but on the shelf
Cibic'idoide.sspp. and especiallyAnomalinoitle.saegt'1tllacr.s dominate. Sectionsin Spain (Ortiz 1995) and
Ital y (B raga et al . 1975; D i N apol i et al . 1910:
-500-1500m depth) show stronglv increasedpercentagesof buliminids abovethe extinctionlevel, including
Bulirninct semicostattt.Faunas from the ReichenhallSalzburg Basin (von Hillebrandt 1962; 200-800 m)
show increasedabundanceof rotaliid taxa.but detailed
stratigraphyis not availablefiom thesesections.We can
conclude tentativelythat westernTethys faunas were
similar to North Atlantic fhunasfiom the samedepth.
In the Indian Ocean,no simple faunal patterncan be
extracted.particularlybecausemany DSDP and ODP
siteshave recordswith poor recoveryand unconfbrmities. Faunasat some lower bathyal-abyssalsites (appendix 12.1) seem to resembleSouth Atlantic faunas
(N. truempt'i-dominated),but abyssarninidsare much
less common or absent.possibly becauseno lower
abyssalsitesare available.Other taxa in the postextinction f'aunasare mostly lon-e-termsurvivors such as O.
Ltmbonatu.s,
although B. sentic'o.statomay be present
abovethe extinctionlevel. Faunasat upper and middle
bathyal sites (appendix l2.l) have very different postextinction faunas. dominated by Lenticulinn species
(Mackensenand Berggren 1992)or by Anonutlirnides
species(similar to fhunasin the easternTethys:Speijer
1991). Such f aunas dominated by Artomalinoides
specieshave beeninterpretedas indicatinglow-oxygen
conditions or high productivity (Speijer et al. 1996b;
Nomura and Kennett. personalcommunication1995).
At all thesesites.however.core recoverywas poor and
the existenceof the LPTM 6lrC excursionhas not been
fully documented.High-latitudeODP Site 738 (1350
m), however.wherethe carbonisotopeexcursionis present(Lu and Keller 1993).has the buliminid-dominated
patternof the Maud R i sesi tes.
Data on the Pacific Ocean are available from few
sitesonly at bathyaldepths.NorthernPacificODP Site
883 (Pak and Miller 1995; 1000-2000m) has a poor
upper Paleocenerecord and an unconformity, but its
postextinctionfaunas appearto be of the N. truentpt,idominatedtype. Faunasat DSDP Site 577 (Pak and
Mi l l er 1992;K ai ho i n press;1800-2100m) havecom mon Bttlintirtusemic:ostulrr
above the extinction level,
are much lower than at
althoughbulirninid percentages
Site 865. At this site. however.a 0.-5-rnsectionin the
criticalintervalwas not recovered(betweenCores9 and
l0 in Hole 577), and the sectionprobablycontainsunconfbrmities (Aubry, personal communication 199-5).
Sites 577 and 865 (describedabove) thus have bulimLand sectionsin Japanand New
inid-dominatedf'aunas.
Zealand(500-1500 m: Kaiho 1988: Kaiho et al. 1993.
224
T HC I' 1 A S
1996) show buliminid-dominated f'aunas. although
cornplicatedby dissolutionintervalswith ag-ulutinated
taxa.
Data on the neritic thunasdo not indicatea relationship between the benthic extinction (thus the LPTM)
and changesin sealevel.The cxtinctioneventoccumed
during the secondorder supercycleTA2 of Haq et al.
(1987),and might have been cclevalwith part of shortterm third-ordercycle 2.3. in mid-nannofbssilBiochron
) n d S p e i j e re t a l . (1 9 96b)di d not
NP 9. S pei. je(1
r 9 9 /+ a
observesignificantsea-levelefl'ectsin neritic seetious
in Egypt. Europeanand North Anrericansectionshave
in their upper Paleocene-lower
many Llnconfbrrnities
Eocene stratigraphicrecord (e.-e.,Olsson and Wise
1992; Gibson and Bybell 1994; Berggren and Aubry
1996).su-e-eesting
that sea level was relatively low at
the tirne. In the northernSpanishsectionsthe benthic
extinction event has been correlated to a lowstand
(Pujalteet al. 1995).lt lrasbeenproposed.however.that
sea-leveltrends in the North Atlantic reflectedthe developmentand collapseof the IcelandPlume underthe
North Atlantic Volcanic Province, and thus were regional ratherthan global (Nadin and Kusznir 1995).
Dis c us s ion a n d Sp e c u l a ti o n o n C a C O,
Dissolution, Productivity, and O, Levels
CaCO , Dis s o l u ti o n
The global data set on the benthic fbrantinif'eralextinction event is incompleteand requireshigh-resolution
studiesto describein detail the eventsthat occurreddurin-e the LPTM. Data are neededfiom sites located on
depth transectsin order to discrin-rinatebetween depth
and geographicefl'ects.Most infbrmation is fiorn sitesat
lower bathyaldepths(figure 12.6a).with f-ewdataliom
abyssaldepths.Geographically.most data are fiont the
North and SouthAtlanticoceans.Coveragein the Pacific
Oceanis extremely poor. data being atvirilableonly tion.t
middle to lower bathyaldepths(fi-eure12.6b).Coverage
is slightly betterfor the Indian Ocean,but recovery is
poor at rnany sites.In addition.taxonomicconeeptsft-rr
specificallyfor Bulinrittrrspp..
the postextinctionf-aunas.
spp.,needsto be
Anonralinoicle.t
Cibitidoidesspp..and
global patterns
researchers
befbre
unifbrrnizedbetween
confidence.
degree
of
with any
can be established
Combining lithological and paleontologicaldatir in
that a ren.rarkable
this incompleteset.however.sug-qests
globalepisocleof carbonatedissolutionoccurredat least
partially coeval with the LPTM (figure 12.7). In the
North Sea region calcareousspeciesoccuned only in
the shallowestneritic areas of the time (Kin-e 1989:
Gradsteinet al. l99zl).This severedissolutionhascom-
rronly been attributedto lctcalf'actorssuch as basinr..striction.which certainlyrnay have contributed.but ti:severedissolutionoccurredsynchronouslywith worlcwide dissolution.In the easternTethys(Egypt). Speile
(1994) observeddissolutionlayers over a wide dep'l:.
range.Benjamini ( 1992)reportedsr.rchlayersfiont tit.
north central Negev (lsrael).and Berg-erenand Aubr.
(1996) ti om the K hi eu R i ver secti oni n Geor gia( N\ \
C aucasus).
S i mi l ar di ssol uti oni ntervi tl soccur in lan. sectionsin Spain(TethysandAtlantic sicles:Coecionie:
al . 1994:C anudoet al . 199-5).
Dissolution is more cLfficultto trace in DSDP an.:
ODP sitesthan in land sectionsbecausepoor recover\
is a problem.A thin laver(<,50cnt) of sedinrentmav no:
be recovered.and thereforemany sitesare listed as "n,
data" in fi-ture 12.7 and appendix 12.1.We catlnotb.'
sure of the tirll geographicand bathyrretricextent trl
carbonate dissolution. but it was extensive in antl
aroundthe Atlantic Ocean.as well as at leastin partst,l
the Indian Ocean,along the Tethys margin. and alon-c
the Pacificmargins.We haveno dataon the Pacifrcdeep
basinfloors.which were probablybelow the CCD clLrring most of the Paleoceneand Eoceneepochs(e.g..a:
l.
shown by red clays in Cole LL,I,I-GPC 3. Rea 199-1
We cannotbe sure about the exact tirr.ringof the initirtion of this increaseddissolution. but it may have
started slightly befbre the LPTM (Thomas l99l:
Thomasand S hackl eton1996:K ai ho et al . 19 96) .
The occurence of such widespreadCaCO.,dissoltttion during a period of warming is unexpectedbecause
solalonewould resultin ciecreased
rising ternperatures
ubilit5rof calcite(e.g..Broeckerand Peng 1984).Therefbre. the occurrenceof widespreaddissolutionstrongh
indicatesmassiveaddition of carbon to the ocean-atmospherereservoir.and the patternof dissolutiotr(rnore
severearoundthe Atlantic.lessin the Pacific)resembles
that predicted to result fiom n.rassiveburnin-eof fbssil
fuel and defbrestation(e.g..Walker and Kastin-s1992).
The occurence of the extremelynegativecarbon isotope anomaly in this interval of dissolutionsuggests
that the addedmassof carbon must have been isotopically unusuallylight. The amountof biosphere-derived
organic carbon neededto causesuch a large rsotopie
anomaly would have to be reflectedin widespreaddestructionof land biomass.which does not agreewith
paleontolo-ticalevidence (Thornas and Shackleton
1996).Therefbre.widespreadcarbonatedissolutionassociatedwith the 6lrC excursionslrpportsthe speculation that dissociationof methanehydrates(andtheir oxidation of methaneto CO.) may have been involved
(D i ckenset al . 1995;K ai ho et al . 1996).This speculation implies that the temperatureincreaseof the deep
watersprecededthe carbon isotopeexcursionslightly.
with Thomasand Shackleton( 1996).
in agreenrent
(-)(-)AeC
FF>
66- ^9E
.9r otur gc aG
P€ * € s s
u\v' L:
-o_o_oti6
ZA
0Jlyo);;i
oc:+;
o-c
uso
-t
F i g u r e 12 . 6 D i s t rib u tio n o f site s liste d in a p p e n clices I l . I ancl 12.2 accordi ng to (a) cl cpth and (b) geographi c l ocati on.
: . t u t h o r so l t h e o l i g i n a l pa p e r su n d g r le n r n a p p e n d ice sI l. I a n d I l .l .
n
il
Z
Z
no data
no dissolution
s l i g hdt i s s o l u t i on
d i s s o l u ti o n
O O A( J E
!t>
:::!O;
za
=>
?
-: * >
c(u(c
--o.5dftfi
ar l goal ;i
er 3a- ;
5 =r ? 3 0
<=>
t
Figure 12.7 Occurrcnceof'clissolution
acrossthe intcrr,alof the benthicfbranrinif-eral
extinctionby (a) depth and (b) geographic
location :sitesliste din a ppendic es12. 1and 12. 2.dept hc a t e g o r i e a
s s i n l i - e u r c1 2 . 6 .
ercascd
dissolution
ilndetching
iclencc-.
of nriclotaunas.
butnoclcarscclinrcntar-r'er
226
THOIYAS
carbonreserSuchan increaseofthe ocean-atrnosphere
(1992)
voir is in contradictionwith Stott'sinterpretation
of data on the carbon isotopic compositionof organic
matter in foraminif'eraltests.although such data cannot
be easily interpreted(Goerickeand Fry 1994:Hinga et
al. 1994) .S t o tt' s d a ta (1 9 9 2 ) m i g h t b e e x p l ai nedby
many factors. such as species-specificeff'ects.diageor salinity.
netic efl'ects.effectsof varying temperature.
More problematicfor the methanehydratehypothesis
might be the precisetiming and sequenceof eventsduring the LPTM: dissolutionappearsto haveprecededthe
61rCanornaly.
Dissolved Oxygen Levelsand Productivity
What caused the benthic fbraminiferal extinction. and
what can explain the biogeographicpatternsobserved?
This is a difficult questionto answerbecausethereis no
agreementon the interpretationof Recentbenthic faunas (see e.g., Gooday 1994, Schnitker 1994. fbr reviews). There is no modern analog for faunas dominated by N. truem1tt'i because this species became
extinct in the late EoceneEpoch. Its modern relative,
Nuttallitles wnbonif'era, is common to abundant in
waters, and specificallyin Antarctic
CaCO.,-corrosive
bottom water (e.g., Bremer and Lohmann 1982:
Schrniedl 1995). Others. however. have describedN.
untboniferaas a typically oligotrophicspecies(Loubere
Po s s i b l y,the w aters
1991, 1994;Go o d a y 1 9 9 3 , 1 9 9 .1 ).
in which N. urnboniferais common are CaCO. corrosive becauseof the presenceof abundantdissolvedCO.
derived fiom oxidation of or-eanicmatter, leaving little
food fbr benthic faunas.thus cornbining oligotrophy
and corosiveness. Some support for the view that .ly'.
truentpt'irnay have been an oligotrophicindicatormay
be seenin its Eocenedistributionpattern:This species
had its highest abundanceduring the middle Eocene
Epoch (e.g.,figure 12.3;Kaminskiet al. 1989;Berggren
and Miller 1989;Miller et al. 1992).It then declinedin
abundanceand becameextinct during the late Eocene
Epoch,at a time when global productivityrnay have increasedconsiderably(e.g.,Hallock 1987;Hallock et al.
1991; Aubry 1992;Brasier 1995;Thornasand Gooday
1996) .
In the Recentoceans,low-diversityfaunasdominated
by small, thin-walledspecimensbelongingto the buliminid group commonly occur in low-oxygen envlronr nent s( e. g. . B o l to v s k o ye t a l . 1 9 9l : S e n G upta and
M ac hain- Ca s ti l l o1 9 9 4 : Ka i h o 1 9 9 4 a ).Su ch envi ronments usually occur below a well-developedoxygen
minimum zone,whereoxygenlevelsare low becauseof
enhancedproductivityand oxidationof organicmatter,
and whereCaCO. dissolutionmay be severebecauseof
of dissolvedCO. resultingfiom
the high concentrations
thi s oxi dati on(e.g.,Mul l i ns et al . 19851H erm elinand
S hi mmi el d 1990: Levi n et al . 1991; S chmiedl 199- 5:
Wishner et al. 1995).There is considerablediscussion
whethersuchbenthicfaunasreflectthe low oxygenler'els or the high food supplies(e.g.,Corliss et al. l9U6r
Linke and Lutze 1993: Miao and Thunell 1993; Loubere 1994: Rathburnand Corliss 19941Jorissenet al.
1995).Recentspeciesof Bulimina are particularlycommon in areaswith high rates of depositionof nondegradedorganicmatter (Corlissand Chen 1988; Caralp
1989;C orl i ss1991; Mackensenet al . 1993).Addit ional
factors of importance might be the lack of metazoan
predatorsunder low-oxygen conditions(Wishneret al.
1995).and some thin-walledtaxa have been described
as being opportunistictaxaratherthan directlylinked to
high productivity or low oxygen (Alve 1994; Goodal
I 994).
We thus cannot determine fiom the benthic f-aunas
alone whether the buliminid-dominatedpostextinctiort
faunas primarily reflect low-oxygen conditions.hishproductivity conditions, stressedconditions. or hi-ch
Similarly. we cannot determine
CaCO.,corrosiveness.
whether the N. truempll-dominated fauna reflects the
occurrenceof CaCO,-corrosivewatersor low productivity. We needto considerevidencefrom co-occuring
planktonicorganismsand sedimentaryand iscltopicevidence in order to evaluatethe most probablecausesfor
the extinction. and compare data on dissolutionwith
thoseon benthicfaunas(figures12.7and 12.8).
It appearsprobable that the occurrenceof small and
thin-walled benthicforaminif'erafollowing the benthic
extinction primarily reflects increasedCaCO., corrosivenessof deep waters becauseboth dissolutionand
thin-walledtestsoccurredat almostall locations(figure
12.7).and the associationof thin-walledtestswith corrosive waters is well documented(Boltovskoy et al.
1991).Proxiesfor high productivityor lower dissolved
oxygenlevelsarenot as common as for dissolution.and
thus neither can be seenas the dominant f'actorcontrolling the occurrenceof the thin-walledtaxa (hgure 12.8).
which are complex metazoans,
In addition.ostracodes.
the LPTM at ODP Site 689
during
becamethin-walled
foraminifera (protistans)
the
benthic
same
time
as
ertthe
shown
to form such "dehave
been
did. Ostracodes
graded" tests in hydrochemicalregimes unfavorable
to biomineralization.i.e., in CaCO.,-corrosivewaters
(Steineckand Thomas 1996).
thus probably played a role
Increasedcorrosiveness
in the extinction.It has beenhypothesizedthat the benthic foraminif'eralextinction was largely causedby decreasedlevelsofdissolved oxygen(Thomas 1989),and
a parallel has been establishedwith the Cretaceous
anoxic oceanic events. although of lesser intensity
(Kaiho 1994b).The LPTM benthicextinctionoccurred
The Late PaleoceneBenthic ForaminiferalExtinction
227
Bb
no data
high O2now prod
low O2ihigh prod
l3
10
=>
o!f,= >
=Fd ( g ( !( g
= 33-oi 6i 6
bobbai
3€=3:
5E :=l /
conditionsor increasedproductivityby (a) depthand (b)
Figure 12.8 Occunenceofbenthic l'aunassuggestingeitherlovn-oxy-een
::rrgraphiclocationlsiteslistedin appendicesI 2. I and I 2.2. depthcategoriesas in figure I 2.6.
-iurin-erapid warming of the deep waters, possibly
by a switchto dominanttbrrnationat subtropical
- rlLlsed
..Ltitudes
of warm and salinedeep to intermediatewa:L 'rs( e. 9. .T hom as 1 9 8 9 :Ke n n e tta n d S to tt 1 9 9l : Pak
.,nd Miller 1992. 199,5).Warmer deep waters can be
:redictedto havea lower oxygencontentbecauseof the
.trong decreasein solubility of oxygen at higher tem( T hom as 1 9 8 9 , 1 9 9 0 a :K a i h o l 9 9 l ). If the
:re ra t ur es
.peculationabout dissociationof methanehydratesis
(Dickenset al. 1995:Kaiho et al. 1996).oxida-'rrrrect
:ron of these compounds would have used large
,rrrountsof oxygenand thus havecontributedto the cre.rtionof low-oxygen environments.There is no agreenrentbetweenthe distributionof dissoluticlnand that of
1ou'-oxygenconditions and/or high prodr.rctivitywith
(figures12.7band
rcqardto depth and paleogeography
[.t3 b ) .
The buliminid-dominatedfaunas.however.did not
rrccur
Faunasindicatingeitherhigh productiv-rlobally.
it1'or low oxygen contentswere most cornmon in the
\orth Atlantic over a wide ran-qeof depths(with possible exceptionof abyssalsites).along Tethys.alon-{the
\ntarctic continent,in the centralPacific.and at upper
to rniddle bathyal depths in the Indian Ocean (figure
12.8).Faunasdorninatedby N. tntemp,r'lwere common
rn the SouthAtlantic. and occurredin the northernmost
Paciflcand at middle to lower bathyaldepthsin the Indian Ocean.It is ditficult to interpretsuch patternsbecauseof the small numberof localitiesto compare:The
North Atlantic DSDP sites.fbr instance.are all closeto
the continents(Bay of Biscay. New Jersey ntargin),
whereas the South Atlantic sites are mainly along
oceanicridgesand rise (figures12.1and 12.6;appendix
12.l). We thus cannot determine whether the differencesin distributionpatternsbetweenthe North and the
South Atlantic are true geographicdifferencesbetween
deep waters in thesebasins.or the difl'erencesbetween
open ocean and margin. The lack of indicationsthat
South Atlantic waters were especiallylow in oxygen
that warm and saline
content.however, may sug-qest
waters did not dorninantly flow fiom subtropical regionsof Tethysinto theAtlantic. If Tethysfunctionedas
a sourcearea.the more probableregion of outflow was
the easternTethys, in agreementwith carbon isotope
data (Seto 1995).On the other hand. agglutinatedbenthic fbrarniniferal assemblagesdo support a TethysAtlantic connection(Kaminski et al. 1996).
It remainsthat faunal pattemswere not globally the
oxygencontentas a
same.and.consequently,
decreased
resultof global deep-waterrvarmingdoesnot fully explain the faunal changesand the benthic extinction.
Thomas and Shackleton(1996) used carbon isotope
data to interpret the postextinction.buliminid-dominated f'aunasat the Maud Rise sites as indicativeof a
high rate of supply of organic matter to the seafloor.
This hypothesisappearsto be in contradictionwith the
vier.l' that the Paleocene long-terrn carbon isotope
recordreflectsa decreasein productivity(e.g..Shackleton et al. 1985:Shackleton19t16;Corfieldand Shackleton 1988:Corfield and Cartlidge 1992:Corfield 1995),
I HO IY AS
a view that is in agreementwith bio-qeographical
patterns of planktonic microfbssils (e.g.. Hallock 1987:
B oer s m aet a l . l 9 u 7 : H a l l o c k e t a l . 1 9 9l : O ttensand
Nederbragt 1992; Brasier 199,5).Theoretically,one
would expect extrernely low productivity during the
LPTM. when temperaturegradientswere shallow, so
that wind velocitiesover the equatorialregions were
very low and upwelling and nutrientsupply to the surf'acewaters were almost null (chapter'9). In addition,
benthic fbraminifera have higher metabolic rates at
higher temperaturesand increasedtemperaturesthus
leadto oligotrophy.
Thomasand Shackleton( 1996),however,arguedthat
global productivity might have declined.but that the
carbonisotoperecordsat the Maud Rise sitessuggested
increasedlocal upwellins. and thus possiblyenhanced
local productivity.Along othercontinentalmarginsproductivity might have also increasedduring the LPTM.
possiblycausedby changedpatternsof Lrpwellin-e
as a
water
result of changingpatternsof deep-intermediate
plankcirculation.High abundances
of chiloguembelinid
in
tonic fbraminif'erain lowermost Eocene sedin.rents
the Bay of Biscay and along the New Jerseyntargin
have beeninterpretedas indicatingincreasedlocal productivity and an expanded oxygen minimum zone
( S aint - M ar cl 9 9 l a , l 9 9 l b : P a rd oe t a l . 1 9 9 7).P roductivity may also have increasedlocally in the eastern
Tethys,wheretypically a "flood of radiolarians"occurs
in the dissolutionlayer (Benjamini 1992:Berggrenand
A ubr y 1996) .At In d i a nO c e a nSi te s1 3 8 .1 5 2, and162
the abundanceof chiloguernbelinidplanktonic fbraminif'eraalso increasesabovethe extinctionlevel (Nornura and Kennett.persclnalcommunication1995). In
other easternTethys sections,however.benthic faunas
suggest low-oxygen conditions as well as increased
pr oduc t iv it y( Sp e i j e re t a l . 1 9 9 6 b ).
Local increasedproductivity during the LPTM in
spiteof the decreased
temperaturegradientscould have
resultedfrom an increasedinflux of nutrientsfiom the
continents.An increasedabundanceof kaolinite in the
sedimentsdepositedduring the LPTM indicatesenhancedprecipitationand chernicalweatheringaround
the world (e.g..Robertand Kennett 199,1;Gibson et al.
t i th cl i mate
1993: K aiho e t a l . 1 9 9 6 ).i n a g re e me n w
(chapter
haveled
raintall
could
higher
9). The
rnodeling
wato
coastal
supply
to increasedrunofTand nutrient
have
also
could
fiesh
water
ters:the increasedinflux of
increasedwater stratificationand causedlocal shelf or
s lopeanox ia( Ma l o n e 1 9 9l ; v a n d e r Z w a a n and Jori ss en 1991) .
Increasedproductivitycannotbe seenas the causeof
all occurencesof buliminid-dominatedfaunas:at equatorial Pacitic ODP Site 865 planktonic fbraminif'eral
f'aunasand coccolithophoridsindicative of extreme
oligotrophy occur in the same samplesas buliminiddorninatedbenthic faunas(Kelly et al. 1996).At suclr
sitesthe faunasprobablyreflectlow-oxygenconditions
in the watercolumn.Suchlow-oxygenwatersin the bathyal openPacificrnayhaveresultedfiom the high temperatures(e.g..Thomas I 990a:Kennettand Stott 199I t.
their fbrmationpossiblyexacerbated
by sluggishcirculation dr-re
to the ertremelylou'temperaturegradients.It
may also be possible.however.that the buliminid-dominated faunas reflect the input of a larger proportion of
organic matter on the seafloorunder lorver oxygen conditions, as supportedby the strong increasein benthic
fbraminif-eral
accumulationratesat the level of extinction at Site 865 (figure 12.3: Herguera and Berger
r 99r ) .
The peakabundancein buliminid speciesat ODP Site
u6-5 is just below a peak abundanceof CiLticitloitles
speciesthat was not observedat other sites.The Cibicidctide.sspecirnensresemble the Recent species C.
v"'ueller.storJi.
which lives on elevatedsupportsabove
the sediment-waterinterfaceand is most comntonin regions with active bottom cuffents (Linke erndLutze
had a similar
1993).If the earliestEoceneCibicicloide.s
habitat. its high abundancemight inclicateincreased
botton.rcurrent activity and winnowing, as also indicated by increased percentagesof material in the
>63-um size fiaction. lor'vaccumulationrates,and intensivereworking of planktonicfbraminif'era(Kelly et
al .1996).
The widespread occumenceof the //. trtretnlttiabyssaminidfaunascan be tentativelyexplainedas reflecting increasedcomosiveness
of botton waters. as
shownby the occurrenceof widespreadCaCO.,dissolution (figure 12.6).Thesefaunas.however.might alsoreflect an overall declinein open-oceanproductivityduring the LPTM. The patternobservedby Tjalsma and
faunas
Lohmann (1983), that l/. tmempt'i-don.rinated
migratedto shallowerdepthsin the oceansduring the
late PaleoceneEpoch. could be explainedby the hypothesis that organic matter derived fiom the surf-ace
waters becameexhaustedat shallower and shallower
levelsin the oceansbecauseof the declinin-eproductivity. as also suggestedby agglutinatedmicrofaunas
(Karninskiet al. 1996).This patternwas not global' and
was not observed,tbr instance.at FalklandPlateauSites
698-102 (Katz and Miller 1991.;.
The question remains open concernin-ethe srgnificanceof faunascontainingcommon to abundantZlpa speciesthat occursin postextinction
lttrttino.selmensis,
f'aunas worldwide over a large depth range but was
common only at neritic to upper bathyal depths befbre
the extinction.Although it might be seenas a part of the
buliminid-dominatedfaunas.this is not a satisf'actory
explanationbecausethe speciesoccursin N. tntetnpti-
The Late PaleoceneBenthic ForaminiferalExtinction
clominatedassemblages
as well. At ODP Site 689 it is
nrostabundanttogetherwith an ostracodefauna dominated by taxa related to the present-daypropontocyprids (Steineckand Thomas 1996).Propontocyprids
are opportunisticforms that live dominantly in neritic
environments.They are confined in the deep sea to
$'ood-basedand hydrothernal vent communitiesthat
fbrm in transient,food-rich environments.I therefore
suggestthat Z selmensis,originally describedfiorn
shallow-waterregions, was likewise an opportunistic
taxon rather than a low-oxygen indicator, similar to
nrodern taxa such as Stoinfctrthiu ffusiforml.s(Alve
1 9 9, 1) .
In conclusion.I suggestthat the latestPaleocenebenthic fbraminif'eralextinctionwas a complex phenomenon.Factorsthat contributedto the extinctionmay have
been the global increasein CaCO-,corrosivenessand
the global decreasein oxygenationof neritic through
abl,ssalwaters as a result of increasedtemperatures.
The averageoceanicproductivity may have declined.
br-rtslu_ugishcirculation and low oxygenation could
have led to the sequesteringof or-eanicmatter in the
cleepsea.The upwelling of suchnutrient-enriched
deep
\\'aters.possiblyin combinationwith an increasein nutrient-carryingrunoff along continentalmargins. mty
havecausedincreasedlocal productivity.leadingto the
erpansionof the trophic resourcecontinuum.i.e.. the
occurrenceof highly oligotrophicas well as highly eutro p h icc ondit ions( H a l l o c k 1 9 8 7 ;H a l l o c k e t a l . 1 9 9I ).
Benthic faunas can then be seen as reflecting both
exh'emes,with the buliminid-dominatedfaunasat the
eutrophic end. the |,,tr.truempti-dominatedfaunas at
the oligotrophicend. with patternscomplicatedby the
occurrenceof opportunistictaxa such as Tltpptutintt
.seltnen,sis.
During the LPTM, benthic foraminif'eralfaunasat ntiddle bathyal and greaterdepthsexhibitedhigh rates of
extinction(30 to -50%of species).whereasat upperbath1,althrough neritic depthsextinctionwas less severe
and faunasshowedsignificantbut temporarychangesin
sp e c iesc om pos it ion .
After the extinction deep-seabenthic foraminiferal
tliunaswere typical for perturbedcommunities(low-dir,ersity,high-dorninance).
StrongCaCO, dissolutionoccurred worldwide. resultingin thin-walledfaunas;this
dissolutionmight have resultedfrom the addition of
carbonto the atmosphere-ocean
systemthroughdissociationof methanehydrates.
Two widely occurring postextinctionassentblages
at bathyal through abyssal depths are M trtretnpt,idominatedand buliminid-don-rinated.
In regionswhere
both assemblagesoccurred,the 1/. truempt'i-dorninated
assemblage
lived deeper(lower bathyal-abyssal.).
229
Productivitymay have decreased
globally during the
LPTM. but this efTectcould havebeencounteracted
locally by lower oxygen levels in the water column. resultingin delivery of a largerfractionof clrganicmatter
to the seafloor.Productivitvmay have increasedalong
continentalmarginsas a result of increasedcontinental
runoff and upwellingof nutrient-enriched
deepwaters.
Speculatively,
N. truentpt,i-dominated
faunasmay be
seenas indicatorsof highly oligotrophicconditionson
the seafloor.and bulirninid-dontinated
faunasas indicators of eutrophic conditions.The occurrenceof both
f-aunas
after the extinctionsuggestsan expansionof the
trophic resourcecontinuum.
A CK NO WL E DG ME NT S
This chapteris a contributionto IGCP Project308. The
research was partially funded by NSF grant EAR
9.+0-5784
to Jim Zachos and Ellen Thornas. I thank
Marie-PieneAr"rbry,Murlene Clark. Rodollb Coccioni.
Andy Gooday.and Mimi Katz for their helpful and constructivereviews. and Robert Speijer.Birger Schmitz,
and Kunio Kaiho for sharingprcprintsand reprintsof
their work.
RE F E RE NCE S
Alve. E.. 199.1.Opportunisticfearuresof thc firrarniniler Stainlorthia
Iu.siJrtrntis(Williarnson):Et idence liom Friertjord. Noru ar,.Joir.
ttttl ol Micropulaeonktlogt,r. I 3. p. 2J.
A ubr1".N I.-P . 1992.P al cogcnecal careous
nannol bssi l sl i orn the K c rguel enP l atcau.Leg 120.In Wi se.S . W.. Jr..S chl i ch.R .. er al .. P roceedi ngsof the OceanD ri l l i ng P rogram.S ci enti fi cR csul ts.r'. 1 20:
C ol l egeS tati on.TX (OceanD ri l l i ng P rogri i rl .lp.
. '171,491.
A ubr1.N I.-P .B crggren.W. A .. S tdt. L. and S i nha.A .. 1996.The up per
Paleocene-loucr Eocene strati-uraphic
record and the PaleoceneEocene boundarl, carbon isotope cxcursion: lntplications fbr
In K nox. R . W O'B .. C orU cl d.R . M. and D un a_v -.
-eeochronol ory.
R. E.. (eds.).Cttrrelutiottrtl tlte eurlt Pttleogent itt Nortllrc.stEurr4rc,Geol ogi calS oci etl ,S pcci al
P ubl i cari onN o l 0l . p. -153380.
B arrera.E . and H uber. B . T.. 1991. P al eogeneand earl v N cogene
occanographv
ot the southernIncl i anOcean:Leg 119 fbrani n i l er
stabl ei sotopcresul ts.[n B anon.J..Larsen.8..et al ..P roceedi n gs
of
the OceanD ri l l i ng P rogrant.S ci entrfi cR esul ts.r. I l 9: C ol l egeS tati on.TX (OceanD ri i l i ng P rogram).p. 693-717.
B asi nger.J. R .. Greenu'ood.
D . R .. and S uedi r.T.. |994. E arl y Terti arl ,
regetati onofA rcti c C anadaand i ts rel el anceto pal eocl i ntatiicn tc rpretati on.In B oul ter. !1. C . and Fi sher.H . C .. (eds.t.C enr;oi c
Plcrntstrttd Clitnate.stl the Arctic. NATO ASI Series. 127. Dordrecht:S pri nger-V erl ag.
p. l 7-5 198.
B ecknunn.J. P . 1960.D i stri buri onof benthi cttrrami ni ttraat the Cre
taceousTerti arvboundarl ,of Tri ni dad (\\'est Indi es).R eport 2 l s t
sessi on.Intcrnati onalGcol ogi calC ongress.N orden.C openhagen.
r'. -5.p. 57-69.
B cnj ami ni .C .. I 992. The P al eoceneE ocencboundarvi n l srael :a candidate fbr the boundarv stratot\pc. Neue.sJahrbtrch liir Gcologrt
mttl PuliiotttologieAhhatdlun,qen.r'. lE6. p. :19-61.
B erggren.W A .. 197.1a.Late P al eoceneE arl v E occnc benthoni c
fbrarriniieral biostratigraphvand paleoecolo-lyof Rockall bank.
Micntpuleontologr;r'. 20. p. .+26-.118.
230
I HO I Y AS
B e r g g r e nW
. . A . . 19 7 4 b .Pa lco ce n eb e n th o n ictir r a m in ile ralbi ostrati graphy'.biogeography.and paleoecologl'of Libya and Nlali. Mlcrolrulertntutlttgt',r'. 20. p. 'l'+9-'+65
e la n kto n iclo r a m in ife rmagnetobi osB e r g g r e nW
. A . . 19 9 2 . Pa le o g e n p
tratigraph)'of the southernKerguelenPlate-au(Sites 717 119\. ln
W i s c . S . W . . Jr .. Sch iich . R.. ct a l.. Pr o ce e d in g so f the Ocean
D r i l l i n g P r o g r a n t.Scie n tilicRe su lts.r . 1 2 0 : Co lle g e S tati on.TX
t O c e a nD r i l l i n g Pr o g r a m )p. . 5 5 l- 5 6 8 .
B c r g g r e n . W . A . a n d Au b cr t. J.. 1 9 7 - 5 . Pa le o ce nc benthoni c
and paleoccologl"
paleobiogc-ographl'
fbranriniltral biostratigrirphy.
ol'Atlantic Tethf irn regrons:Midrvav tvpe fauna.Puku'ogetnra1tltt'
r'. 18. p.73 192
Prlcteoecokryt,Pulueotlinntttlo,qr',
Berggren.W. A. and Aubry. M.-P. 1996.A late Paleocene-carlvEocene
correlatton
NW Europeanand North Sea tnagnetobiochronological
n e t u o r k . I n K no x. R. W. O' 8 .. Co r lie ld .R. N4 .a n d D una1.R . F-..
(ctls.).Corrclttirnt ol tlrc Ettrlt Pulcogent'in NttrlhuestEuropt Ge'
l b lica tio nNo . l0 l. p .,3 0 9 - 351.
o l o g i c a lS o c i e tySp e cia Pu
. C.. III a n dAu b r y-M. P - 1995.
B e r g g r e nW . A . . K en t.D. V. Swish e rC.
A rclised Cenozoic geochronologl and cllronostratigraphl'.In
. (eds.).
B c r g g r e nW
. . A.. Ke n t.D. V..Au b r v,I{ .- P a n d Ha r d e nbolJ..
Clcot'hnnoktgt finrc Sculesaul Gltthrtl Strutig,ruplticCttrrtlttt ittrt.
a n d Nlin e r a logi stsS pecral
S o c i c t l ' o f E co n o m ic Pa le o n lo lo g ists
P u b l i c a t i o nN o . 5 ' 1 .p . 1 2 9 - 2 1 2 .
Berg,urcn.W. A. and Miller. K. G.. 1989.Cenozoicbathyal and ab-'"ssal
calcareousbenthic tbrarniniferal zonation. Microltaleontrtlttgt,v.
3 5 . p . 3 t ) t 33 2 0.
Cretaceous-Tertiaryplanktonicibrarriniters tt'otll
Boersma.A.. 198-1a.
n n tic. \[h lvis Rid g e a r e ir .Dce p S ea D ri l l i ng
t h c s o u t h e a s tcrAtla
P r o j e c tL c g 7 , 1 .In M o o r e .T . C.. .1 r ..Ra b in o wltz.P D . ct al .. P roceedingsof the Deep Sca Drilling Project.v. 7'1:\\'ashington{ LI.S.
C o v c r n n r e nPt r in tin gOtticcl. p . 5 0 1 - 5 2 3 .
o ce n ea n d o L h e rT cr tia r ' 1b'cn th ictorarl rni l era
.
B o e l s m a .A . . 1 9 8 .1 bOlig
fiom a depth traversedou'n Walvis Ridge. Dcep Sea Drrlhng Proof the
j c c t L e g 7 5 . l n Ha 1 ' W
. W.. Sib u e t.J. C . e t a l . Pr o ceedi ngs
D e e p S c a D r i llin g Pr cle ct.v. 7 5 : Wa sh in g to n( U.S Governmel tt
P r i n t i n gO f f i c e ) .p . 1 2 7 3 1 3 0 0 .
B o e r s n r aA
. . a n d Pr e m o li Silva . I.. 1 9 9 1 . Distn b u tio nof P al eogene
planktrmicfbrantinitera:analogiesu'ith the Recentl Pltltaogt'rt,qtrtphr. Pulutoclinrutologt.Pakreoetologr: v. 83. p. 29Jlt
Boersnra.A.. Prenroli Silva. I. and Shackleton.N. J.. 1987. Atlantic
Eocene planktonic firrarninif'eralpaleohydrographicindicltors and
Puleotlutogrtryrlnlr'. 2. p 287 331.
stableisotopepaleoceanogtaphy.
B o l l i . H . M . . B eckm a n n .J.- P a n d Sa u n d e r s.J. 8 .. 1991. B erttl ti c
.fon.tntiniferulhiostrtttigrapltt'of the sttuth Curibbean tzgirll. Cantpp.
bndge: CarnbridgeUniversity Press.'1011
benBoltovskol'E
. . a nd Bo lto r sko y.D.. 1 9 8 9 .Pa le o ce n ePlei stocene
i tl
e
t h i c f b r a m i n ile r a le vid cn ceo f ln a .jo rp a lco ce a n o g raphicrettts
. lvis Ridge).Marl rra
t h e c a s t c r nS ou thAtla n tic ( DSDP Sitc 5 2 - 5Wa
Mit'ntpuleonktlogr',v. l'1. p. 283 316.
B o l t o v s k o l ,E
. . . Sco tt.D. B. a n d M e d io li. F . S. 1 9 9 1 .N4orphol ogi cal
Variationsof benthic tirrarninif'eraltestsin responseto changesln
A revierv.Jutrnal ol Palermtoktgt l 65. p
ecologicalparat'netcrs:
175 Iu5.
Boltovskoy.E.. Watanabe.S.. Totah. V. I. and Vera Ocanlpo.1.. 1992.
Cenozoicbenthic bathyaltbrarninif'ersof DSDP Site 5'18(North At
lrntic ). M i t ropaIeonktlt t s t. i. 3t3.I 83-207.
Boltovskoy. E.. Watanabe,S.. Totah. V. and Vcra Ocampo. J.. 199-5.
B e n t h i c t o r a min ifb r stio m DSDP Site 5 1 6 ( u p p e r Maestri chi tan
Quatemary.SouthAtlantic). RetistuEspcriuthdc Micntpulettttlttkt
g i t t , v . 2 7 . p . lll l- 1 9 .
B r a g a . G . . D e Bia se . R.. Gr u n ig . A. a n d Pr o to De cinl a. F. 1975
Foran'riniferibentonici del Paleoceneed Eocene della Sezronedi
Possirgno.In Bolli. H. M.. (ed.), Mttno,qrophiuMicropult'orttttlogittr
Provitttitt di Trt'risirt Itulitt,
.sttlPtilt'ocenee l'Eocene tli Po.tsugnrt.
e h a n d lr L n g evn .9 7. p. U 5-19l .i .
S c h u ' e i z e r i schPa
e ltio n to lo g ischAb
B ral ou,cr.T. J.. P arrou,.M..-fhomas.E . anclZachos.J. (-.. 1 995b.D ata
Report: Stirbleisotopic stratigraphlof the Palcogenepelagtccap at
S i te865.A l l i sonGu1ot.l n Wi ntcrer.E . L.. S ager.W. W.. Fi rth..l .\.
of the C )ccanD r i l l i ng P roand S i nton.J. \{.. (eds.).P roccedi ngs
gram. S ci cnti fi c R esul ts. r'. l '13: C ol l egc S tati on. T X (Oc ean
p. 581-586.
D ri l l i ng P rograrn).
B ral ou'cr.T. J. . Zachos.J. C .. Thomas.E .. P anou. N I.. P aul l .C . K .
K. C .
K el l ey'.D . C .. P remol iS i l va. I.. S l i ter.W. V . and L-ohrnann.
of the equatortl l
to E ocenepal eocel nographv
I 995. Ll te P al eocene
Pacific Ocean: Stableisotopcsrecordetlat C)ccurDrilling Pro-eranl
r" 10.p. 8'tl 865.
Sjte 865. Allison Cuyot. Pulertttttrut,qra1tllr'.
B rasi er.M. D .. l 9c)5.Fo.tsi li ndi tutor.s(t ttul ri (nt l erel s.2 . E tol uti ott
utttl e.rtinr'tiottin rt'ltilittttto oligotroltlt\'.Geolo-sicalSocietySpecial
P ubl i cati onN o t33.p. l l 3 150.
B remer.N {. l -. and Lohtnann.C i .P . | 981. Fr i dcncefbr pri m arl 'c ontrc l
of certai nA tl anti cOceanbenthoni cl b rami ni l -era
of the cl i stri buti on
by degree ol crrbonate saturation.Deep-Stu Reseunlt' r'. 29. p.
9117998.
A msterdi rnlE
: I s c v i er.7' 19
B ri g,es.J. C ., 1995.Gl obul B i ogtttgrzTrhr"
pp.
T.-H .. l 9E '1.The cl i rnatecheni s trvc ol l l l c c W. S . and P en,e.
B roccker'.
l r4ttrttg ntphs ,t29.
C topl tt.si tttlLi ni l rnC i ct4tl tr.si L'ttl
ti crrr.rl rti crl trur
p. 327 336.
Brotzen. F.. 19.+lt.The SwedishPuleocerrcuntl it.s.fbruntiniletal
.frutnt
seri esC . no '191 (i i rs bok -i .
S l eri ges Gcol ogi skaU ntl ersokel se.
19.18).1.10pp.
C ande.S . C . anclK cnt. D . V .. 1991.A neu'gcomagncti cpol ari t) tl l l c
andC enozoi c.Jttuntulrtl G ertl thts i Lul
scal el br the LateC rctaceous
r" 97rB l 0t. p. 13.i )17 l 3.t)-5J.
R e.scunh,
antl
K cnt. D . V .. 1995.R eri setlcal i brati onof th e geoma-gS
.
C
.
C ancl e.
netic polaritl titt.tc scale fbr the l-ate Cretaceousantl Ccnozoic.
v. 100(B'l).p. 6093 6095
.lourna! oJ Cit'oplttsitul Re.;eurch.
C anudo.J. L. K el l er. C i .. N Iol i na.E . and Orti z. N . l 99 5 P l ank ti c
turnover tnd 6l rC i sotopcsacrossthe P al eoc c nefbranri nrl 'eral
Eocene transitittnat Cara\lca and Zunlava. Spttitt,.Puluertgertgntr'. I ll. p. 75 100
Pulucot'r'rrlogr',
phr. Pulttcot'litnutrtlttqt.
of B ul i tni ttuari l l r and Mc l otti sburC aral p.M. H .. 19E 9.A buncl ance
Relationshipto the qualit) tlf marineorganicmatter.Groleecutttnt.
Marine Letters,r'. 9. P. .j7J3.
E vi denc etrom
C crl i ng.T. E .. 199 1. C arbondi oxi cl ei n the atl nosphere:
ttf S c i etttt"r"
C enozoi canclMcsozoi cpal eosol s.A rttt'ri ttttt.l rtunutl
29l . p. -r77J00.
C hari si .S . D . and S chnri tz.B .. 1995.S tabl c(6rrC .61sO)ands tronti un.)
(ErSr/865r)
isotopesthrough the Paleoceneat Gebel Au'eina.eastern
Tethl's region. Ptrlueogcogrttpltv.Pttlaeoclinutolttgt, Ptlueot'tolo,gr:v. l l 6. p. l 0-l -129.
Charnock. N'[.A. ancl Joncs. R. W.. 1990. A-uglutinatedfbrarninitcra
from the Palaeogencof the North Sea. In Hernlclren.C , Karrinski.
B i os truti :4N {.A .. K uhnt.W. and S cott.D . B .. (eds.).P al eoat'ol o.qt'
rtl' Agglutittttttd frtru
uul'littttrnntt
rttpln, Pult'oL'erutrtQtttpht
and Physical
ninileru, NATO ASI Scrics. Series C: N{alherr.ratical
S ci cnces.r'. 327. D ordrecht:K l uu'er A cademi c. p. 129 263.
Cllark. M. W. and Wright. R. Cl.. l9ti'1. Paleo,ueneabyssal benthic
fbrarni ni l ersfi onr the C ape l nd A ngol a B asi ns.S o uth A tl anti c
Ocean.In H si i . K . J.. LaB recque.J. L.. et al . Ini ti al R c portsof the
(U .S G o\c rnment
D eep S ea D ri l i i n-eP roj ect.r'. 73: \l 'ashi n-eton
P ri nti ngOffi ce).p.'159 '11i 0.
. . 199' +l.ntegratc c l
C occi oni .R .. D i Leo. R .. Gal eotti .S . andN {oncchiS
biostratigraphyand benthictirranlinifcralfaunal turnoveracrossthe
boundarl at Trabuaka Pass section. Northern
Paleocene-Eocene
r'.'+. p. 87 100.
Spttttt.Puluet4rcirl,qo.r,
l l tnrsof tl teLrntl ottC l rl r:P al aeontol ogC ol l i nson.N {. E .. i 9U 3. I-ossiP
als Fi el clGui desto Fossi l s.r" I (P al aeontol ogi cA
i cal A ssoci ati on.
soci ati on.London).l 2l pP .
The Late PaleoceneBenthicForaminiferalExtinction
('or1le l d R
. . M . . 1 9 9 3 .D e p th h a b ita tsa n d th e Pa la e o ce nrea d ia ti onof
t he p l a n k t o n i cl b l a m i n ile r an r o n ito r e du sin g o xlg e n a n d carbon
iscrtopes.
In Lees.D. and Edu'ards.D. (cds.).Etolutitmurv Pottent.t
at t l P t n t ' e . s . s e
L .i n
s ,n c a nSo cictvo f l,n n d o n .p . 5 9 - 7 0 .
Ct r1f lg l 6R. . N , { . 1
. 9 9 5 .P a la e o ce noece a n sa n d clim a te :a n is( ) tr r liepcrspective.Earth Scierre Rzlfu,lt r. 37. p. 225 352.
C, rrf ie l d .R . N { . . a n d C a r t lid g c'J.
. E,.. 1 9 9 1 .lso to p ice vid e n cetb r the
dep t hs t r a t r f i c a t i oonf l ir ssila n dr e ce n tGlo b ig e r in in aa: r cvics. H i .rLrt r i u l B i o l o g v r ' . 5 . p . 3 7 - 6 3 .
i Lrriie l dR. . M . a n c lC a r t l i clg eJ.
. E.. 1 9 9 2 .Oce a n o g r a p h racn d clinrati c
implications of the Palaeocenecarbon isotopc maxiinrtnt. Terra
rVala, v..1. p. -1.13J55.
, , rrlie l d .R . M . a n d S h a c kle to nN.
. J.. 1 9 8 8 .Pr o d u ctivr tych a n g cas a
control on planktonic lbraminiteral evolution after the Cretir
ceous/Tertiaryboundarv.Hi.sntritul Blolo,qr:r. I . p. -123 3'1-1.
r'' rrf ie l c lR
. . M . . C r i r t l i d g e.J. E.. Pr e m o liSilva I. a n d Ho u slcy.R. A ..
199 1 .O r y g c n a n d c a r b o n iso to p cstr a tig r a p h yo l th e Pa la e o-rene
and C r e t a c e o u lsi m e s to n e sin th e Bo tta ccio n eGo r g e a n d th c ContessaHighu av sectiol.ls.
Urnbria.ltaly. Terru Nolr, r,.3. p. .11"1J22.
i, rrlis s . B . H . . 1 9 9 l . M o r ph o lo g y'a n d m icr o h a b r tapt r e t' e r e n ceosf benthic firraminif'erafronr the nortlr$'estAtlantic Ocean. trlurine illitrr4tuleorttolo.qt.
r'. | 7. p. lt).5 236.
. -, , rlis sB
. . H . a n d C h e n .C .. l9 ll8 . M o r p h o typ ep a tte r n so l' No r we-ei an
Scad c e p - s e ab c n t h i cf or a r r in iti' r aa n d e co lo g icaim
l p lica tio n s.Geo1o , q rt .: 1 6 .p . 7 1 6 7 1 9 .
. orliss .B . H . . M a r t i n s o n .D. G. a n d Ke ft' e rT. .. 1 9 8 6 .L a te Qu a tc rnarv
. lee p - o c e acni r c u l a t i o n (. ito lr r yita l Stxie tt r tf An te r ict Bu lle t i n,t.
97.p . l l 0 6 - l l 2 l .
(t
-u\irnrarr. J. A.. 19.+6.Ultper Crt'taceous.fttrarninift'ro the Gttl.l
Coa.stulRegiorrof the United SIute.suul tdjucertl arerr.i.U.S. Geological Survey ProfessionalPaper206. 2.li pp.
Lrilev . D . H . . 1 9 8 3 . L a tc Cr e ta ce o u sa n d Pa le o ce n eb cnthi c
ibr a r n i n i t e r sf r o m D c cp Sca Dr illin g Pr q e ct Site 5 1 6 . Rio Gr ande
Rise .n e s t e r nS o u t hA t la n ticOce a n .ln Bir r ke r P
. F ' ..Ca r lso n .R. l -..
J oh n s o n .D . A . . e t a l .. ln itr a l Re p o r tso f th c Dccp Sca Dr i l l i ng
Pro j e c t . v . 7 2 : W a s h in g to n( U.S. Go ve r n m e n tPr in tin g Ofl l ce).
p. 7 5 7 - 7 8 2 .
)rek en sG
. . R . . O ' N e i l . J . R .. Re a .D. K.. Ou cn . R. M .. 1 9 9 5 .Dissoci at ion o f o c e a n i cm e t h a neh vclr a te
ls a ca u seo f th c ca r b o niso lo peercursion at the end of the Paleocene.Pulttxeunttgrultll.r;r'. lt). p.
965-97 L
-r:\ apo l i A l l i a t a . E . . P r o kr - De cim aF. . a n d Pe le g r in i.G.8 .. 1 9 7 0.S tu
. t r a t i g r a licoe. M icr o p a le o n to lo g ico
d e i Din to rntdi
dio G e o l o g i c o S
Bclluno. Mentoric Sot ieta Gertlo,git'uIIuliunu. r. 9. p. I 2ll
r. t . 's . R . a n d l ] u t c h i s o n .J. F t.. 1 9 8 0 .Eo ce n elo u ' cr \cr te b r a te sIrorn
. l a n a d ia Ar
. lu ttr ye o entpl tt,
Ell e s r n e r eI s l a r r d C
n ctic Ar ch ip e la g o Pu
v. -10.p. -315 3.17.
Pulut'oclinntologt.Pulueoet'olrry1;
Freen r a nK. . H . a n d H a 1 ' e s.J. \4 .. 1 9 9 1 .F r a ctio n a tio no fca r b o n i sole ve ls.C1obo1
t op e sb y p h 1 ' t o p l a n k toann d e stir r tltcso l a n cie n CO.
t
Bio g e o lt t t , t n i tu l C r r ' / r ,r r. ' . 6 . p . I 8 5 l9 tt.
I rc r0c h .S . l n d N o l a k . W.. 19 8 - 1 .Pr o p o sa lo f zo n a tio nlir r th e l rte
Tithonian lrte Eocene.based Lrponarenaceoustirraminil'erafionr
t hc o u t e r f - ' a r p a t h i a n s.
Po la n d .BENT HOS ' 8 3 . Se co n dIn tcrnl i t ion r l S l , r r r p o s i u m
o n Be n th ic F o r a n r in ifcr a{.Pa u . F r a n ce .A pri l
I 98 3) . p . 2 2 5 - 2 - r 9
t iibs on .T . G . a n d B 1 ' b e l l L
. . \{ .. 1 9 9 .1Se
. d in r cn ta rpva tte r n sa cr o ssthe
Pll e o c e n eE o c e n eb o un d a r yin tl.r cAtla n tic a n d Gu ll' co iistr lp l ai n\
ol'the UniteclStates.Brrlfu,trrr
dL' lu Srttiet! Belge tle Glolo.qit, t.
I 03 . p 2 3 7 - 1 6 5 .
(iif rron .T . G . . B y b e l l . L - .M . a n d O*e n s. J. P. 1 9 9 3 .L a te stPa le occne
Nr-u
lit h o l o g i ca n d b i o t i c e r,e n tsin n cr iticd cp o sitso l so u th we ste rn
.lersev.Rrlt,rrrrrno.qrultln.t'. 8. p. -195 5l.l.
(ioeric k e .R . a n d F r y . 8 . . 1 9 9 .1\'. a r ia tr o n so f m a r in ep la n klo n6 1 r Cni th
lat i t u d e t. e n r p e r a t u.r e
d
in th e u o r ld clcca nGl
. rbrrl
a n d d isso lVe CO.
Bio q t ' o c h e n i tu l ( \ ' r ' 1 r ,r8, . lt5 9 0
23 I
speci esw hi ch ex Goodal '.A . J.. 1993.D ccp-scabenthi cforami ni t'eral
pl oi t phytodetri tus:
and control son di strihuC haracteri stifeatures
c
rion. Ilaritrc Mitropuleonlolo,qr;v. 22. p. ltiT-20-5.
Gooday.A. J.. 199.1.The biolo-eyof dcep-sealirraminifera:A ro'icu, of
Paluio.s,
someadvancesancltheir irpplicationsin paleoceanognrphy.
r'. 9. p. l :1-31.
C radsLei n.
F. M.. K ani i nski .M. A .. B erggren.W A .. K ri sti ansen.I. L.
and D 'Ioro. N {.A .. 199.+.
o1'theN orth Sc a
C enozoi cbi ostrati graphy
anclLabrador She)f. Micropaleontolo,qt,r'..10(Suppl.). J52 pp.
i n the trophi crcsourceconti nuum:a tac H al l ock.P . 1987.Fl uctuati ons
raph\ t'. 2. p. 157-17 l .
tor in gfobal diversitl cvclesl Paleooceunog
H al l ock. P . P rcrnol iS i l va. L and B oersma.A . 1991.S i mi l ari ttesb c tueen planktonic and larger fbraminiferal evolutionary trends
throtrgh Paleogenepaleoceano,eraphic
changes.Palaeogt'ogropht
Pttlueoclitnatrtlogy.
Pulaeoetologr',r'. 83. p. '{9 64.
. andV ai l . P R .. I 987. C hronol ogyof 1'l uctuatH aq. B . U .. H ardenbolJ.
r'. 235.p. l l -56 1167.
i n-useal cvel ssi ncethe Tri assi c..S ci orcr,,
W. H .. 1991.P al eoproducti vi ty
from benH ergucra.J. C . and B er-eer.
thic firranrinif'eralabunclance:
Glacial to postglacialchangesin the
."r'est-equatorial
Pacific.Geologr',v. 19. p. 1173 1116.
H ermel i n.J. O. R . and S hi mmrel d.G. 8.. 1990.The i mportanceof the
orvsen minimurr zone ilnd sedimentgeochemistryin the distribution of Rccentbenthic fbraminiferain the Nofihs'est Indian Occan.
Mttrine Geologt; r'. 91. p. I 29.
H rnga.K . R .. A rthur. M. A .. P i l son.M. E . Q. and Whi taker.D .. 19 9.1.
Carbon isotope fiactionation bv marine phytoplankton in culture:
the cffccts of CO2 concentration.pH. temperature.and species.
C l ri l nl R i ogeocl tenti cal
C ycl es,v. 8. p.9l -102.
und sei neForanti ni tc renV on H i l l cbrandt.A .. 1962.D as P al aeozaen
fauna i m B ecken von R ei chenhal lund S al zburg. B ayeri sc hc
Akademie der Wissenschaften.Mathematisch-Naturs'issenschaftN eueFol ge.v. 108. I 82 pp.
l i chc K l asse.A bhandl ungen.
i m A l ttert i aer
V on H i l l ebrancl t,
A .. 1965.Forani ni fcrenS trati graphi e
von Zumava (Provinz Cuipozcoa.N\'\LSpanicn)und ein Vergleich
mit irnderen Tethl's-Cebieten.Bayerische Akademic der WisK l asse. A h
senschati en.Mathemati sch
N aturni ssenschal tl i che
hancl l ungen.
N eueFol ge.v. l 2-3.62 pp.
H ookcr.J. J.. 1991.The scqucnceof mammal si n thc Thi rnetrana nd
Y prcsi an of the London and B el gi an B asi ns. [-oci i ti on of the
Palaeocene-Eocene
boundarv. Na+slelterson Stratigntpht. t. 25.
p. 75 90.
H orni brook.N . D e B .. B razi cr.R . C . and S trong.C . P . 1989.Manualof
Neri ZealandPermianto Pleistoceneftrraminif'eralbiostratigraphy.
NevrT,eolund()eological Sunet Pultorttologitul Bulletin Ntt 56.
l 9l 3pp.
bound ary
H ovan. S . A . and R ea. D . K .. 1992. P al eocene/E ocene
changesin atmosphcricand oceaniccircr-llation:A southernhenrisphererecord.Geol oi qt,r. 20. p. I 5 I 8.
H . andWri ght .R .
H si i . K . J.. N IcK enzi e.J. A .. Oberhl i nsl iH. ,. Wei ssert.
In Hsii. K. J..
C.. l9tj.1.SouthAtlantic CenozoicPaleoceano-eraphy.
I-aB recque.
J. L.. et al ..Ini ti al R cportsofthe D eepS eaD nl l i ng I' ]roj ect. r. 73: Washi ngton(U .S .GovernncntOl l l ce).p. 771 785.
fronr the upperc onH ul sbos.R . 8.. 1987.E ocenebenthi cfi rrarni ni tcrs
Deep Sea Drilling Project Site 605. In
tinentai risc ot'l Nerv Jerse1".
ran H i nte.J. E .. Wi se.S . W.. Jr..et al .. Ini ti al R eportsof thc D eep
S eaD ri l l i ng P roj ect,r. 93: \l ashi n,cton(U .S . GovernmentOl 'flc e).
p. -525-5-18.
H ul shos.R . E .. K roon. D .. Jansen.H . S . N '[.and van H i nte.J. E .. 1989.
Lou'er L,ocenehenthic fbraminrferaand paleoenvironmcntof the
outerV ori ns P l ateau.N oru'egi anS eaTD S D PS i te 338).Mi t'ntp ul totttuthtg.t,
v. 35. p. 256-273.
J. G. V .. 1995.A concep
Jori sscn.F. J.. de S trgter.H . C . and Wi cl nrark.
tual nroclelerplaining lrenthic fbranrinif'eralrnicrohabitars.M a r i t te
Mi tntytl tortnthr,qr,r'. 26. p. 3-l -5.
232
TIOIYAS
K a i h o . K . . 1 9 8 8 .Up p e r n u r srCr cta ce o u s p a le ogene
to
bath_yal
benthi c
l o r a n rin ile r abl io str a ttg r a p hovfJa p a na n d
Ne ri Zeal and:Latestpa_
l c u c r ' n er n id d le F .o u cn ch ,.n r h i..r t,r
..
' r r i,,ir t,"-r
r:!:;:'
_.!,,.,," a,r,,u,,t,i,,toi illrr#lHT'l]'J::i]
K a i h o . K .. 1991. Global
paieogene
rurno\
ef.
changes of
aer obi c /anaer obi c ben_
ancl deep-sea circulation. pultreogeo,qraplt.r..
Pt l u erx l i n u k t k t,q pa l trco e t.o l oqrt r,.
_t.,
[j-3,p. 65_116.
K a i h o . K .. l99.la. Benthic fir r ar linif.er al
dr ssolvec l _ox y gen i ndex antl
d i sso l ved oxyuen lcr els in thc ntr ,tler r r
lceiln. C eol ( r ,q\; l . 22.
thic
lbranrintl'era
p . 7 1 9 - 7 2 "2
K a i h o . K . . l9 9 .lb . p la n kto n r ca n d b e n th ic
fb r ami ni f.cralexti ncri on
events during the Iast I00 nt.y. pulueo,gt,o,qrupht.,
paltteoclirttuktl_
ogt. lltlueoeutlo,gr:r.. I I l. p. -+5_71.
K a i h o . K . . in p r e ss.Disso lr .e cl
o xyg e nch a n g e sin thc deep_sea
duri ng
the last r00 miilion years based.n benthic
lirranrntfer^. parueo
gert,qr up h t, puk rerx-I i n u to I ttg t, pttk rco t
et r I o.q.t.
K a i h o . K . . M o r g a n s.H. E. G. a n ti Oka d a .
H.. t9 9 :. Faunalturno.,.erof
lntermediate-waterbenthic fbrarlinif.era
during the palcogcne in
New Zealand.Morine Mit.r.oltulettntoltLgt,,
v. 21. p. 5l_E6.
K a i h o . K . . Ar in o b u . T .. Isliiwa r a .R.. M o r g a n s.g .
E . C .. Okacl a.H ..
'fakeda.
N.. Z h o u . G., Ka jiwa r a .y. M a tsu r n o to.R .,
H trai .A ., N i i t_
s u m a . N. a n d Wa d a . H.. 1 9 9 6 . l_ a r cst pal eocenc
benthi c
t o r a n t i nit' e r ael xtin ctjo na n cl e n vir o n m e n ta l
changcsat l aqanui .
New Zeaiand.puleoceturtt,qrupllr;
r,. ll. p. aa7 A6i.
K a m r n s k i ,\ ,1 .A.. Gr a d ste ir rF. . M . a n clBe r g g r e n .
W A .. t989. pal co_
gene bcnthic tirranriniter biostratigraphv
and paleontologyat Site
6 : 1 7S
. o u th e r nL a b r a d o rSe a .In Sr iva staS.
. p . Arrhur.M.. C l entent.
B . . e t a l .. Pr o ce cd in gosf th c Oce a nDr illin g p r o lram.
S ctenti fi cR e_
s u l t s . r ' . 1 0 5 : Co llcg e Sta r io n .T X tOce a n
Dr il l i n-r programl . p.
70-5 730.
K a m i n s k i . N { . A.. Ku h n r . W. a n d Ra d le y.
J. D.. 1996. pal aeocenc
Eoccne dcep water aggrutinarcdrbraminif'era
riorn the Nurnidian
flysch (Rif. nortltern Morocco): their significance
lirr the palaeo_
ccirnographyol' thc Gibraltar gate!\,ay.Jrturrutl
of.M it.ntlsaleottol
o , q . rr;' . 1 5 .p . J- 1 9 .
K a t z . N { . E . a n d NIille r , K. G.. l9 9 l.
Ea r ly pal eogencbenthi c
t o r a m t n ife r a la sse n r b la g cs
a n d sta b le iso to p e si n the S outhern
O c e a n .l n Cie se lski.p F .. Kr isto ffe r scny.
, e t al .. proceccl i ngs
of
t h e C ) c e an
Dr illin g p r o g r a n t.Scie n tificRe su lts.r ,. I l .l :
C ol l egeS ta_
t r o n .T X (Oce a nDr illin g p r o g r a m ) p
. ..{ g l_ - 5 1 1 .
K e l l v . D . C . . Br a io u e r . T . J.. Z a ch o s.J.
C.. p r e nrol i S i l va. l . and
T h o m a s .E .. 1 9 9 6 .Ra p idd ive r sifica tio o
n f p la n kroni ctbrami ni tera
r n t h e t r o p ica l p a cific ( ODp Site E6 - 5 )clu iin g
the l are pal eocene
Therntal Maximunr. Geoktg.t,v 2a. p .123J26.
K e n p . . E . M . . 1 9 7 u .T e r tia r yclin r a r ice r lo r u ti.n
a n d . ,.ecetati .n
hi stor\,.
in the SourheasrIndian Ocean region.pulueoge,,griqrhr.p,rlur,,,,.li.
rnututlog\Pulutot,t.ologr,r,. l.l. p. 169-20tj.
K e n n e t t J. . P an d L . D. Sto r t.1 9 9 0 .p r o te u s
a n d p r o b _Oceanus:
A nces_
tral Paleogeneoceansas revealed1.romAntarctic
stabic rsotopcre_
s u l t s .I n B ar ke r .p F .. Ke n n e tt.J. p . e t a l.. p r o ce e di ngs
o1.theC )cean
D r i l l i n g P r o g r a m .Scr e n tillcRe su lr s.r .. ll3 :
Co llel e S tati on.TX
( O c c a nD r i llin g p r o g r a m ) .p . g 6 - 5 - 8 g 0 .
K e n n c t r J. . P a n d Sto tt.L . D.. 1 9 9 1 .Ab r u p t
d e e p _ seu,arrni
a
ns.pal aeoc e a n o g r a ph icch a n g e sa n d b e n th ic e r tr n ctir ) n s
a t the end of the
Palaeocene.
N(rilre, \,.3-53.p. 22-5_229.
K i n g . C i . .I 9 8 9. Ce n o zo ico f th c No r th Se a .
In D. G.. Jenki nsand J. W.
Murra:-, (eds.). StrutigruplticttlArlu.rof. lo.ssi!
Foruntittilera,(2nc)
e d . ) .B r i t i s h M icr o p a le o n to io g icaSo
l cie tl,Se r ie s.Chi chester:
L.l hs
Horu'ood l_inrired.p. .ll g{g9.
K o c h . P I - . . Z a ch o s.J. C. a n d Gin g cr ich .p
D.. 1 9 9 2.C orrei ati onbe_
t!!ecn isotopc rccords in rnarine and continental
carbon rescrvorrs
r c a r t h c P a la e o ce n e /Eo cebnocu n d a r 1 Nu
,. tu r t,,t. - 15Ep.
. -l I 9 322.
K o c h . P l - . . Z ach o s.J. C. a n clDcr r m a n .D.
t_ .. 1 9 9 -5.S tabl ei sotope
s t r a t i , u r a phat,n tl p a le o clin ta to io gol,f th e p a lco g e ne
B i ghorn B asi n
(Wyonring. USA). Rrlazo,geo,qrupltl.,palaeoc!irnuktloet.
tntd
Pulueoecologt,v. I I 5. p. 6l_g9.
K uhnt. W. and K anti nski .M. A .. 1g90.pal coeco l ogy
ol Late C rc ta_
ceous [o paleocenedeep_wateragglutinatetl
lbramrnif.erafront the
North Atlantic and tr,esternTeth1,s.In
Hemleben. C.. Kamtnski.
M. A.. Kuhnt. W. anclScott. D. 8.. (eds.).Rtlcoecolrtgr..
Birt.rtntti.g_
raphr, Poleoceawtgruphtuttl Tu.ronotnt.
of'i\,q,qlttinttretl
Frtrttntirti_
li,ra. NATO ASI Series.SeriesC: Mathernatilal and phl,sical
Sci_
ences.v. 327.D ordrecht:K l uu,erA cademi cpubl i s hers .
p. a-13-505.
Le C al vez, y. 1970. C ontri butron i I.6tude
cl es Forami ni fi res
pal dogdnes
du B assi nde pans.C ahi ersde pal 6o ntol ogi e.
92 pp.
Ler,i n.L. A .. H uggett.C . L. andWi shner.K .
F.. 199 1.C onrrolof dc ep_
seabenthtccontmunitv structurebv oxvgen
and organiu_mattcr
grF
dients in the easternpacific Occan. Journu!
rf- ilrtrine Reseurtlt.
v. .+9.p. 763_tt00.
Li nke. P and Lurze,G. F.. 199-1.Mi crohabi tat
pref .erenc es
ol .benthi c
forantinii'era_a static
1bodacquisition.,r,,,,'i,i"ii,f,',,ii),j#il,,"1
Loubere.P. I 991. Deep-seabcnthic fbraminif.eral
assemirlage
respouse
to a surtaccoceanproductivity gradrent: Atest. pult,ocettno.gftrlttt\.,
r,.6. p. l9-t_10.1.
Loubere.P . l 9g.l . euanti tati r,eesti mati onof
surtaceoc eanproc l uc ti t,
i ty and bottom w ater
using ben
rninif.era.
o,,,,.
thic fbra-
r,:;:t;:.:l ; ;l;"]li;r
",,,
I-u. G. and K el l er.G..
1993.The pal eocenc_E occne
trans rtl oni n rhe
Antarctic Indian Ocean: Inl'erencettom planktic
ltrrantinif.cra.
,ty'a_
ri tteMi crcpul eontol og\.,
\..21.p. l 0l -112.
Lu. G. and K cl l er,G.. 1995a.pl ankti ctbramrni l .eral
fh unalturnorc rsi n
the subtropicarpacrlic truring the late pareocene
to carrv Eoccne.
Journ aI of Forttrrti n iferu I Rt,.rettrty't,l. 2-5.p. 97I ) 6.
Lu. C . and K ei l cr. G.. 199-5b.
E col ogi calstasi sand sal tati on:S pec i es
richnesschangein planktrc tbranriniferaduring
the latc paleocene
to early Eoccne.DSDP Site 577. pulueogcogrupht,pakrctx.lintutol_
ogt',Palueoacologr:v. I I 7. p. 2) | 227.
Maas.M. C .. A nthony.M. R . L.. Gi ngcri ch.p
D .. Gunnel l .G. F-.and
K rause.D . W.. 1995.Mantntal i angeneri cdi versi t y
and turnov eri n
the late Paleoceneancl earlv Eocene of the
Bighorn and Craz1,
Mountains Birsins.W1,,om
ing antl N{ontana(ll S,i.). prtt crettgat rtr
ry
pht, Polaeot.lintttktlo.qt,
palttetx,rttlo,q.t,
v. I | 5. p. I g l_107.
,
MacJi enscn.A . and 13erggren.W A .. 1992. paleoeenc
benthi c
forarrinil'erstr.ontrhe SourhernInclianOccan (Kereullen plateau):
B i ostratrgraphv
and pal eoecol ogl In
,.. Wi se. S . W. J r.. S c hi i c h.
R.
et al .. P roceedi ngs
of the Ocean D ri l l tng progran t.S c i enti l j cR e_
sul rs, r'. 120: C oJl egeS tari on.TX (Ocean
D ril l i ng program).
p. 603-630.
Mackensen.A ..
Fi i terer.D . K .. Grobe.H . and S chmi edlG..
. 199-1.
B en_
thic lbrantiniferal assenrblagesfiont the eastern
South Atlantic
P ol ar Front regi on betw ,een
35. and 75" S : D i stri butron.ec ol ogv
and fbssifization potential. Murine Mitntpcrlaontolttgt.,
v. 22. p.
33-69.
Malone. T. C.. 1991. Rirar.flow,
ltlntrtphtrtktonprotluttiort trnrl oryqert
depl(tiott irt ChesopeakeBa.r:Geological Society
Spectalpublica_
ti on N o. 58. p. 83-93.
Manum, S . B .. i 99.1.The pal i rer:rgene
l l ora ol .spi tsber gen.
In B oul ter.
M. C. and Ftsher.H. C.. (cds.). Canrt:oit.ptunt.s
lurc! Clinrcte.srtl
rtte Arcti(. NATO-ASI Series. 127. Dordrechr:
Springer_VcrJag.
rl5
rr
)l)
Markw i c.k.P J.. l 9g,+..'E quabi l i tv...
conrrnenral i r),.
an tl Tc rti arv..c l i _
mate": the crococl i l i an
perspecti \c.Gcol tl ql .,v. 21. p . 613 616.
McGouran. B .. 197.1.
Forami ni tera.
In ron cl erB orch.C . C .. S c l ater.J .
C .. et al .. Ini ttal R eportsol the D eep S ea D rrl l i n-gproy ec t.
r.22:
\,\rashrngron
(U.S. Governmenrprinring Oftice.r.
p. 009-OUl.
McGow ran.8.. 1991.E rtrl ul i on antl envi .onnent
i n th e earl v pal eo_
gcne.Menoir ol tlte Geohtgicu! Sot.it,ttrl lndiu.
v. 20. p. 2l--5-j.
The Late PaleoceneBenthic ForaminiferalExtinction
\ lc K e n n a .M . C . . 1 9 8 0 .E oce n ep a le o la titu d eclim
. a te .a n d r n a m n ralol
s'
E ll e s m e r eI s l a n d . P r rlr r e o g e o g r a p ltt.
Pa h te o clitttu to lo gPu
t. lutotcologrt r'. 30. p. 3:19-.362.
\ liao. Q . a n d T h u n e l l . R. C.. 1 9 9 3 . Re ce n t d e e p - sca b enthi c
t br a r r . l n i f e r adlr s t r i b utio n in
s th e So u thCh in a a n d Su lu Se a s.Marin e M i c r r 4 t a l e o n k t l o .gt.2
t, 2 . p . I 3 2 .
\ lillc r. K . G . . K a t z . M . E . a n d Be r g g r e nW.
. A.. 1 9 9 2 .Ccn o zo icdeepsea benthic tbraminifera:A tale of three turno\,ers.Studie.sin Renrltic fitrurniniferu, BENTHOS '90. Tokai Unirersitl' Press.Sendai.
p. 6 7 7 5
\ lillc r. K . G . . F a i r b a n k s.R. G. a n d N' lo u n ta inG.
. S.. 1 9 8 7 a .T e rti ary
l
n
ory g e n i s o t o p cs t n t h csis.se ale ve l h isto r y.a n d co n tin e n tamargi
crosion.Pnltolcano,qrupln,v. 2. p. I 19.
\ liller. K . G . . J a n e c e k T.
. R.. Ka tz. M . E. a n cl Ke il. D. J.. 1 987b.
. \ h y s s a lc i r c u l a t i o na n d b e n th icfb r a m in ite r ach
l a n g e sn e a rth e P ar'. 2. p. 711-16l' .
leocenc/Eoce
ne boundarl'.Puleot'eoutant1tftr,
\ lolina . E . . C r n u d o .J . J . . M a r tin e z- Ru iz.F . a n d Or tiz. N.. 1 9 9 .1.Intcgrateclstratigraphvacrossthe Paleocene/Eucene
boundirrvat Caravrca. southern Spain. Ecloqne gt'ologittt Helvetiue, v. 81.
p. - 1 7 6 1 .
\ lolina . E . . C a n u d o .J . J . . Gu e r n ct.C.. M cDo u g a ll.K.. Or tiz. N.. P as
c u a l .J . O . . P a r e s J. . O .. Sa m so .J. lv{ ..Scr a - Kicl. J. a n d T o squel l a.
J . . 1 9 9 1 .T h e s t r a t o t l p icIle r d ia nr e visjtcd :In te g r a te dstr a tigraph)
boundarv.RevuecleMfuntpaliutlolorcross the Paleocene/Eocene
qlr , ,r . . l - 5 .p . l - 1 3 l - 5 6 .
\ lLille r - M e r z E
. . a n d O b c rh iin sli.H.. 1 9 9 1 .Eo ce n eb a th l' a la n d a byssal
benthic fbraminifera from a South Atlantlc transcctilt 20' -10"S.
l. 8-1. p.
Pukteo,qeoqrupltv.
Puluerxlirttulologt, PtLlaettetzrlo,gr;
1t 7 t 7 l .
\ lullin s . H . T . . T h o m p s o n .1
T. L..
. .B.. M cDo u g a ll.K. a n d Ve r co u te r c.
J985. Oxygen-rninimur.r.r
zone edgc eft'ects:Evidence frorn the
r'. 13.
central Califirrnia coastal upu'elling system. Ge.)1o,q11
p. - 1 9 1 J 9 . 1 .
\ lurrav . J . W . 1 9 8 9 .P a l e og e n a
tront
e n d Ne o g e n eb e n th icfb r a m in if' e rs
. e t a l.. ln itial R cRo c k a l lP l a t c a u I. n R o b e r ts.D. G.. Sch n itke rD..
po r t s o f t h e D e e p S ca Dr illin g Pr o je ct.r ' . lll: Wa sh in g to n(U .S .
G o v e r n m e nP
t r i n t i n gOflice t.p . 5 0 - l 5 3 .+ .
\ , rc lin . P A . a n d K u s z n i r.N. J.. 1 9 9 - 5Pa
. la e o ce nuep lifi a n d Eocene
subsidencein the northernNorth Scabasin liom 2D fbrq'ard and re
Verse stratigraphicrrodcling. Journul ol the Geologit'ul Socien.
Lt n u k t t . v . I 5 2 . p . 8 3 3 - 8 .1 8 .
\ ils en . T . H . a n d K e n . D . R.. 1 9 7 8 .Pa le o clir n a tic
a n d p a le o g e o graphi c
la te r itc( la to so l)o n th e lcel andinr p l i c a t i o n sn f a l o u e r T e r tia r ,"FaeroeRidge. North Atlantic region. Ceolo,giculMuga:.ine,t'. 115.
p. 1 5 3 - 2 3 6 .
\ ornur a . R . . 1 9 9L P a l c o ce a n o u r :r p hoyf Up p e r M a e str ich t i anto
Eo c e n eb c n t h i c t i r r a min ile r a a
l sse m b la g east Site s 7 5 2 . 7 5 1 and
. T :r ylo r .E.. A l t.
75 . 1 e
. a s t e r nI n d i a nO ce a n .In We isse l.J.. Pie r ce J..
J . . c t a l . . P r o c e e c l i ngosf th e Occa n Dr illin g Pr o g r a m .Scic nti fi c
Re s u l t s .r . l 2 l : C o i l e g e Sta tio n .T X ( Oce a n Dr illin - uPr o gratr).
p. 3 - 2 9 .
s n dCe nozoi c
O Con n c l l .S . 8 . . 1 9 9 0 .Va r iltio n sin Up p e r Cr cta ce o u ir
N'laudRise. \\reddell Sea. Antarccalcium carbonatepercenta-qes.
t ic a .I n B a r k e r .P F . . K cn n e tt.J. P. e t a l.. Pr o ce cd in gos1 ' th eOcean
'fX
Dr i l l i n g P l o - u r a n rS. c re n tificRe su lts.r ' . ll3 : Co lle g e Sta tio n.
lO c e a nD n l l i n g P r o g r a m )p. . 9 7 1 - 9 1 3 .1 .
Olsson. R. K. and Wise. S. W.. Jr.. 1987. Upper Puleocettelrt ntiddla
Eot errctlepositiottalsequetue.sarul liutu;e.s itt tlrc Netr .lt'rsa AtluttiL nturgin. Cushr.nanFoundation for Foranrinit'eralResearch.
Sp c c i a lP u b l i c a t i o n2 . 1.p .9 9 | 1 2 .
O rue E r t e b a m i a .X . . A p p e llin iz.E,..Ba ce ta .J. I.. Co ccio n i.R.. N '[ons Pu.i al te.
. 1 ' r oA..
c ch i . S . . N u n e z - B e t e lli.K.. Pa r e s.J. M .. Ja u r n ePa
V.. S a n s o .J . a n c iS e r r a - Kie l.J.. in p r e ss.Pr o sp e ctivePa le ocene
Eocene boundrr') stratot)'pescctions in the decp $ater Basquc
233
basi ns. w estern P 1'renees.S parn: brostrati graph)'and mag netostratigraphyof the TrabakuapassanclErmua sections.Mictrtpuleottlo l ttgt.
Ortiz. N.. 1995. Diftbrential patternsof benthic fbrarninittral extinctions nearthe Piileocene/Eocene
boundaryin thc North Atlantic and
r'. 26. p. 3.+1 359.
the \\'esternTethys.Mrrrlnc Micntpuleorttutlosr;
Orti z. N . and N l cD ougal l .K .. 1991.B i ostrati graphland pal coeco l ogl
ot the srnallcr bcnthic forirminifersfiorn the EuropeanPaleo-eenc
rrari nc Il crdi anS tagc{N E S pai n).Geol ogl tttlS oti en' rtl A nteri t' tt,
Abstnu t.swillt Pnt,qrtuns,r'. 23. p. A37.
di verA . J.. 1992.P l ankti cforami ni f'eral
Ottens..l .J. and N ederbragt.
sity as indicatorof oceanenvironrnents.MzlrlrteMicnpuletnttolo,qr.
r,. 19.p. I 3-28.
P ak. D . K . ancl Mi l l er. K . G.. 1992. P al eoceneto E oceneben thi c
Inrpl i cati ons
tor deepwater
tbrami ni l erali sotopesand assembl agcs:
r'. 7. p..105 -+ll.
circulation.PaIeoct,tutographrt
P ak.D . K . and Mi l l er. K . C .. 1995.Isotopi cancll aunalrccordof P al eogene deep-w atertransi ti onsi n the N orth P aci fi c.In R ea. D . K ..
of the
B asor'.L A .. S chol l .D . W. andA l l an.J. F..(cds.).P roceedi ngs
OceanD ri l l i ng P rogram.S ci enti fl cR csul ts.r'. 1.15:C ol l cgeS tati on.
TX (OccanD ri l l i ng P rogramt. p. 265-281.
P arcl o.A .. K cl l cr. G.. Mol i na. E . and C anudo.J. I.. 1997. P l ank ti c
l brarri ni feraltLrrnovelacrossthc P al eoceneE ocenc trl nri ti ou nt
D S D P S i te .101.B av of B i sca,"-.
N orth A tl anti c.Muri rteMi ctr4n l t'
rutl ohte\,r. 29. p. 129 l 5tl .
PopP arker.W. C .. C l ark.\4. W.. Wri -sht.R . C . and C l ark.R . K .. 198.1.
ul rti on cl vnami cs.P al eogencabyssalbenthi cforami ni fers.castern
J. L.. ct a]..Ini ti alR ep orts
S outhA tl anti c.In H si i . K . J.. [-aB recc;Lre.
of the D eep S eaD ri l l i ng P roj ect.r'. 7-3:Washi ngton(U .S . Gcx'ernnrentPrinting Ofhce). p. .18l -1116.
R .. 1978.Macrobcnthi csl l cce\\i unj rr l eP earson.T. H . and R osenber-u.
l ati on to organi ccnri chnrentand pol l uti onof the mari nc envl ronn1enl.Oae(uto,qrophtuntl Mttrine Biolo.ry rlrrrtitrtlRetlt'tt: r'. 16.
p.129 3l l .
P renrol i S i ha. I. and B ocrsma. 198.1.A tl arrti c E ocene pl anktoni c
tbraminiferal historical biogeography rncl palcohvdrographicinrlices.Pultteo,qt'ogrupltt.
Pulaeot l inutolttgt, Pulueoetrtlttgt.t. 67.
p.315 356.
X .. P a rds .
l . Orue-E xtebarri a.
P uj al te.V .. B aceta.J.-1..D i nards-TurelJ..
J.-\{. and P ayros.A .. 1995. B i ostrati graphi cand magr)et(' \tri tti depos i graphi ci ntcrcaJi brati on
l nd P al eocene
of l atestC l retaceous
ti onal sequcnccs
fronr the deepuatcr B asqueB asi n.\\csternP v rc nees.S pai n.E urtl turul P l urtel (rt S (i etI(( Lett(rs,\. I 36. p. I7 30.
R athburn.A . E . anci C orl rss. B . H .. 199-1.Thc ccol ogy of l i vi ng
(stained)deep sea bentlric tbraminilera fiorr the Sulu Sca. Pnlt'oceutograpl tr.t. 9. p. 87 150.
R ea.D . K .. 199-{.The pal cocl i mati crecordprovi cl edb1'eol i andep os i
of G aoti on i n the deepsea:the geol ogi chi storyof u'i nd.R el l t'rr'.s
7rlrr'.ri.r, 32. p. 1-59 l9-5.
R ea. D . K .. Zachos.J. C .. Ou'cn. R . M. and Gi ngeri ch.P D .. 1990.
Gl obal changc at thc P al eoceneE occnc houncl ar\':cl i rnati cand
evofuticlnart'consequencesOf tectonic erents. Prt/rt?a,gettgntpltt',
Puluatxlitnutologl. Iltlaeoet oltryr: r'. 79. p. I I 7 128.
the P u
L.8.. 1992.Fuurtulcl tartgei rthettl ti c.fbrurni ni Jeruu
R ey'nol ds.
leot-ene/Eot
ene bounlurr. DSDP Site,5J9.Nlasterof ScienceThcsi s.Orono:U ni versrtlof Mai nc. 130pp.
and E ocenckaoli ni te di s R obert.C . and K cnnett.J. P . I 992.P al eoccne
tri buti onrn the S outhA tl anti c and S outhernOcean:A ntarcti cc l i rrati c and pal coceanographc
i mpl i cati ons.Muri ne C eoktgt:r. 1 03.
p.99 I 10.
Robcrt.C. and Kennett.J. P. 199.1.Antarctic subtropicalhurnrdcpisode
at the Palcoccnc-Eoceneboundarv.Clav rlineral evidcnce.Gcalogrl r'. 22. p. 2l l -21.1.
234
TtO\,1AS
nannoe lcareous
R o m e i n .A . J . T . . 1 9 7 9 .L in e a g e sin e a r ll Pa le o g e n ca
Bu
p l a n k t o n .U t r eth t M icr o p u le o n to lo ,q ita
l lle tin ,v. 2 2. 2-31pp.
studvrtl '
a n d p xle o e n vir o n mcntal
Saint-N'larP
c .. 1 9 8 7 .Bio str a tig r a p h ic
Paleocenebenthic and planktonic lbraminit'ers.Sitc 60-5.Decp Sea
D r i l l i n g P r o j e ctL e g 9 3 . In r a n Hin t6 .J. E.. Wise . S. W .. Jr..et al ..
I n i t i a l R e p o r tso f th e Dccp Se aDr illin g Pr o je ct.r ' .9 3: Washi ngton
( U . S .G o v c r n n r e nPr
t in tin gOffice ) .p . - 5 3 9 - 5 .1 7 .
S a i n t - N 4 a r cP.. l 9 9 la . [.e Pa l6 o cd n e t le p a ssa g ePa ldocc:nc-E oci ne
sur la bordureseptentrionalccluGolf'ede Biscayc(AtlantiqueNord.
e t p a l6 ocdanographi c.
S i t e D S D P ' 1 0 1 . l- e g .1 8 ) .Br o str a tig r a p h ie
Bulletin de lu Sotidtl Giologitluc de Frunce,v. 163.p. I ll-5 Il3l.
et pal6ocdanographiclues
Saint N'[arc.I 99 I b. Donn6esstratigraphiques
s u r l e s d € p 6 tsd u Pa ld o cb n csu p ir ie u rd e la p a r tic occi dental ede
l ' A t l a n t i q u cN o r d ( fjo ssecle Ba ltim o r e .Site DSDP 605). P ukteogeogrupht, PulaeocIinuilologt, Palaetrccttl
rtgr:l. li5. p. -135-350.
S a n d e r sH
. . L . . 19 6 8 .M a r in e b e n th icd ive r sitl,'A: co n tpal atr\estud\'.
A r n c r i c u t N u tu r a list,v. 1 0 2 .p . 2 .1 3 - 2 8 2 .
S c h r n i c d lG
. . . 1 9 9 5 .L a teQu a te r n a r )b' e n th ictb r a r lin itcr alassernbl ages
t i o n t h e e a s te r nSo u thAtla n tic:Re co n str u ctioonf d e epu rtet ci reuchanges.Reports(nt htlur Re.stunh,Allietj
lation and prcrductrvit-v
v. 160.207 pp.
WegenerInstilutelor Polur utttlMurine Reseur<'h.
S c h m i t z .B . . S p e ije r .R. P a n d Au b r y. M .- P. 1 9 9 6 .L a testP al eoccne
b e n t h i c e r t i n ctio u e \e n t o ll th e so u th cr nT e th ya n shel l '(E -cl -pt):
v. l '1.
F o r a r l i n i f i r a l sta b leiso to p ic( 5 1 ' ' C.6 r sO)r e co r d s.Ct'o1o,qr:
p. 317-35t).
Schnrtker.D.. 1919.Clenozoicdccp water benthic firlarninitcrs.Ba-"of
B i s c a y .I n M o n ta d e r t.L .. Ro b e r ts.D. G.. ct a l.. ln itia l R eportsot the
D e e p S e a D r iL lin - ePr o je ct.r ' . .{ 8 : Wh sh in g to n( U.S. Govcrnl rcnt
P r i n t i n gO f f i ce .)p. . - 1 7 7 - - 1 1 3 .
S c h n i t k e rD
. . . 1 99 1 .De e p - se ab cn th ictb r a n r in ite r s:F o od and bottorl )
w a t c r r r a s s e s.ln Z a h n . R.. Pe d e r se nT. . F .. Ka r n in s ki .N '1.A . and
L a b e . v r i el.- . . ( e d s.) .C' u r b tn tCttlittg in tlp Glu tia l Ott'tttt; C on. u ' rbrk: S pri ngcr
s t r a i n t su t t h e Oce u n ' sRo la itt ( ]lttb u lCiia ilg c'Ne
V e r l a g .p . . 5 1 9 5 5 - + .
Schrijder.'L. 1991.A pall'nological zonation lbr thc Paleoceneof thc
r , Il . p. II3-l 26
N i r r t l rS e aB l t str ,..lo u n n o
l l M itr ttp u ltte o ttto lo gt.
S e n G u p t a . B . K. a n d M a ch a in - Ca stillo .N' I. L .. 199-1.tsenthi c
lirrarniniterain orlgen-poor habitats.ivlurirrcMitrttlttrletntloktgt.r'.
20.p. 183 201.
of the InSekr. l9t).5.Carbon anclorvgen isotopic paleoceano-eraphy
n ra n d p al eo-oeerlei
c l i a na n d S o u thAtla n ticOce a n s:p a le o clim a te
S er.C . t. 10.
c u l r t i o n .. l o u nu l o f St ie n te o l Hir o sltitttttIJttive r .sitt.
p. 393 -1t35.
. 1 9 9 1 .Da ta r e llo rt:Orrgen ancl
S e t o .K . . N o m u r a.R. a n d Niitsu n r a N..
n l ouer E occne
c a r b o ni s o t o per e co r d so f th c L lp p e rN{ a e str ich tiato
n d i an Occan. In
b e n t h i c f b r a r n in itir sa t Site 7 5 1 in th e H,a ste r ln
ol the
. Alt. J.. e t r l.. Pr o c cedi nss
\ ! ' e i s s e l J. . . P ie r cc.J.. T a 1 ' lo rE..
.
n tih cRe su lts.r ' . l2 l: Col l egeS tati on.
O c e a nD l i l l i n g Pr o - e r a mScie
T X ( O c e a nD r illin g Pr o g la r r ) .p . llti5 8 1 t9 .
S h a c k l c t o nN
. . J .. l9 tt6 . Pa lco g e n esta b lciso to p ee r e n ts .R tl rtt'o3ea,qv. 57. p. 9 l | 02.
rupht'.Puluetxlinruttilttgt Puluertecrtlrtgt,
S h a c k l e t o nN
. . J .. 1 9 8 7 .T h e cr r b o n iso to p er e co r do f the C cntl zoi c:
H i s t o r - ,o- l ' o l ga n ic ca r b o nb u r ia l a n d o f o r lg e n in the oceanatrd
J. a n d F le e t.A. J.. ( cd s.) .M u r i nt' P el ntl t'trnt
a t f n o s p h c r eI n
. Br cxr ks.
S o u n e R o c k .s.Ce o lo g ica l So cie ty'Sp e cr a l Pu h licati onN o f6.
p. .123J34.
. 1 9 8 1 .T h e clir n a tcol the E ocene
S h a c k l e t o nN
. . J . a n d Bo cr - sn r aA..
o c e a n . . / o r r nr r rol l th e ( ie r lr tg iL ' u lSo cie tvttl ln n d on, l . 138. p.
| 5l-l 57.
. r h o niso to p estri rti graph)'of
ShacklctorN
l . . J . a n clHa ll. M . A.. 1 9 8 - 1Cla
b u l k s e d i n r c n ts.
ODP Site s6 li9 a n cl6 9 0 . In Ba lke r .P F.. K ennett.
J . P . c t a l . . Pr o ce e d in gos1 ' th eOce a nDr iJlin gPr o g ram.S ci cnti l i c
R e s u l t s .r ' . l l 3 : Co lle g e Sta tio n .T X ( Occa n Dr illing P roql am).
p. 9lt5-t)89.
l 9E ' 1.
N . J. and N '[embers
of thc S hi pboardS ci enti l i cP art,"-.
S hackl eton.
l n l l oore. T. C .. J r..R abi A ccul nul ati onratesi n l eg 7-1sedi tnents.
noui tz. P D .. et al .. l ni ti al R cportsol 'the D eepS eaD ri l l i n,eP ro.j ec t.
v. 7.1:Washi ngton(U .S .GorernnrentP ri nti ngOfhce).p. 621 631.
S hackl eton.
N . J.. C orfl el d.R . N I. and H al l . N {.A .. 1985.S t abl ei s uope
pl anktoni cfbranl i ni tera../orrrdataand the ontogcn)oi P i tl eocenc
R esettnl t.r'. 15.p.321-.136.
nul ol Forunri rti fenLl
S i te 2-15.[n S i rnpson.E . S . W.. S c hl i c h.R .. et
S chl i ch.R .. et al .. 197.1.
al .. tni ti al R eportsofthe D eepS eaD ri l l i ng P roj ect.r.25: W hs hi ngton (Li .S .Go\ernmentP ri nti ngOfti ce).p. 187 2.36.
A ttrospheri ccarbonc l i ox i deand
S l oan.L. C . anclR ea. D . K .. 199-5.
carl )' E ocenecl i rratc: A generalci rcul ati onnl odel i n gs etts i ti ri tr
stutlt. Pttl ueogcog nt plt\ Pul att tt l i nutttt l o gt. Pulaeoecttl ogt. 119' p
275-292.
T. C . Jr.. 1995.P os s i b]erol e
S l oan.L. C .. \['al kcr..l .C . G. and N 'l oore.
of oceani cheattranspotti n eari v E ocenecl i mate.P rl l rac l ul l ogrup/rr',r'. 10. p. -l-17356.
S peqer. R . P . 199.1.E rti ncti on artd recovcry patterns i n benthi c
firraminit'eralpalcocomrttunitiesacross the Crctaceous/Paleogene
buundaries.Geologittt Ultruicctirtu. MedrrnclPalcocene/Eocene
edelin.qen vutt rle Fucull(it Aonl\t el('ttstlutlrytert Ltniversiteit
Utretli. r'. I 2.1.l9l pp,
R . P . r'artdct Zuaan. G. J. arrtlS chnti tz.8.. I996a.The l mpac t
S pei .j er.
of Paleoccnc/f:ocenebourtclart'events on middlc neritic bcnthic
lirranrirriteralassctrblagesfi-cltrEgl'pt. Mtritte trlitroltuletttttttlttllt
v. 28. p. 99 I 32.
S pei j er.R . P . S chni tz. 8.. A ubr1.N {.-P and ('hari si .S . D .. 1996b.The
l atestP al eocenebcnthi c erti ncti on event: P unctuatedttl rnov c r l l l
outcr ncritic torarriniferal taunls frttnt Gebel Au'eina. Egypt ln
B ound. .. (eds.).P al eocenc/E oc ene
A ubry'.M.-P antl B ertj anti niC
r' .
arv E ventsi n S paceand Ti nre.Isnrcl Jouurulrl E urth .S ti rrl tr' .r,
-+1.p. 2t)7 222.
S tei neck.P L. anclThornas.E .. 1996.Thc l atestP al eocen ceri s i si n the
dccp sea:Ostracodcsuccessionat Maud Risc. SouthernOc.^anGeo/o,qr'.r. 2-1.p. 5tl3-5E6.
and l ouer oceani c7 ;C O.:A c l i S ton.L. D .. 1992.H i ghcr tcmperatures
nrateenigtnaat thc end of the PaleoceneEpoch.Rift',t ertnrt,qntpltt.
v. 7. p. 395-.10.1.
S tott. L. D . and K ennett.J. P . 1990.A ntarcti cP al eoge nepl ank toni c
: P Leg l l 3. S i tes689 and 690. l n
fbrami ni l erbi ostrati graphlOD
ol the Oc eanD ri l l i ng
. ..et al .. P rttceedi ngs
B arker.P F.. K ennett..lP
P rosram.S ci enti fi cR esul ts.r'. l l 3: C ol l egc S tati on .TX ((J c earr
D ri l l i ng P rogl anr).p. 5'+9 569.
W. A .
S tott. L. D .. S i nha.A .. Thi ry-.N I.. A ubr1. N 4.-Pand B er-r:gren.
tl te P al eoceneE oceneboundarl ' :
1996.Gl obrl 6l l C changesacr()ss
conel ati ons.Itt K nor. R . W . O' B .
C ri teri a 1or terrestri al -nl ari ne
Corticlcl.R. N'l.antl Durral'.R. E.. (cds.). Corrt'lutiortrt the Earlv
Puleogerrcin Northtest Eltrry;r'.Geological Societ)'Special Publicati r)nN cl . l 0l . p. -r8l 399.
N . J. anclC orti cl d.R . C .. 1990.
S tott.L. D .. K ennctt..l .P . S hackl eton.
In
duri ngthe P al eogene:
The evol uti onot A ntarcti csurl i tcer'r'atets
tlrences front the stable isotopic contposititln of pliinktonic
l brami nrfers.OD P Leg l l 3. In B arker.P F.. K el l nctt.J . P . et al ..
P l oceecl i ngs
of thc OceanD ri l l i ng P rogranr.S ci enti ti cR es ul ts .r' .
I I 3. p. E -19863.
Thurras. tj .. I989. D crel i tprnent of C enozoi c deeps ea benthi c
A.. (ed.t.Orl '
forarl i ni f'erall aunasi n A ntarcti c\\ater\. i l l C l tanl e..J.
gitt.sttntl Ewtluliotr ol tltt t\rttunlit Blata, Geolo-eicnlSocietl' Spcci al P ubl i cati onN o.17.p. l 8.l -196.
benthi c
throuuhN eogencdee p-s ea
Thorl {s. E . 1990a.Lrte C l rctaccous
In B ark er.P F..
fbrami ni l ers(N 'l audR i se.Weddcl lS ea.A r.rtarcti ca).
K ennett.J. P . et al .. P rocccdi ngsof thc OceartD ri l l ing P rogranl .
S ci enti fi cR esul ts..v. I l 3: C ol l egeS tati on.TX (OceanD ri l l i ng P rogranr). P. 57 I -5q'+.
The Late PaleoceneBenthic ForaminiferalExtinction
Eo' ce n em a sscxtin ctlo ns1n
irirrnn sE. . . 1 9 9 0 b .L a t e Cr e ta ce o u s- ca r l1
tlrc deep sea.In Sharpton.V. [-. and Ward. P. (eds.).Cktbul Cutu\tr()plk's irt Eurth Hi.\lor\': Att ittterdistiplittttt-tCttrtlen'rtt't'rtnlnt
ptttts.Vtlcati.sttt.tuul Ma.s.s
Mrtrttilitt, GeologicalSocictv of Alnerrc aS p c c i a lP a p e rN o . 2 .{ 7 .p ..1 8 1 - - 1 9 - 5 .
E.. 1992.Clenozoicdeep seacirculation:Evidencefiorl deep
. irtrr.ncs.
\ er b e n t h i ct i r r a r n i n i f b r aln
. Ke n n e tt.J. P a n d Wa r n ke .D.. ( e ds.).
A Perspectitetttt Gktbul Change.
Tlrc AnturttiL Puleot'rttintturrcnl:
.\merican Geophl'sical Union Antarctic Reselrch Scries No. -56.
p. 1 . 1 1 - 1 6 5 .
: h()m a s .8 . . a n d G o o d a l . A. J.. 1 9 9 6 .Dccp - se ab e n th icfb r a n tin ifera:
tracers firr changes in Cenozoic oceanic productiritl'. Caolo,qr',
p. 3 5 5- 3 5 8 .
. J.. 1 9 9 6 .T h e Pa le o ce n eEo ce n eb e nthi c
f hLrnr a sE.. a n d S h a c k l e t onN.
t irra m i n i t c r a le x t i n c t i o n a n d sta b le iso to p ea n o tr a lie s.In Knor.
. r r t' ktti rtrt
rt.l
R. E.. ( e d s.) Co
R. W . O ' 8 . . C o r f i e l d .R. Nl. a n d DL r n a y.
tlte Earlt Pttlcogenein Nortlnl(sf ElrnTre.GeologicalSociety Spec ial P u b l i c a t i o nN o l 0 | . p . .1 0 1.+ .1I .
inLrm a s E
. . . B a n e r a .E . . H a m ilto n .N.. Hu b e r .B. T .. Ke n n e tt.J. P. O'Con n e l l .S . B , . P o s p i c ha l.J. J.. Sp ie ss.V.. Sto tt.L . D.. Wti. W. and
e a tig la ph)'of
\ \ ris e .S . W . . J r . . l t ) 9 0 .Up p e rCr cta ce o u sPa le o g e n str
. tsa r kcr .P F .. K enSit c s6 8 9 a n d 6 9 0 . M a u d Rise ( An ta r cticir )In
nc tt .J . P . e t a l . . P r o c c cclin gosfth e Oce a nDr illn g Pr o g r a n l.Sci cn. X { Oce a nDr illin g Pr o g ri l m).
t it ic R c s u l t s v. . l l - l : C o l lcg e Sta tio nT
p. 9 0 1 9 1 . 1 .
T honra sE
. , . .S h a c k l e t o nN. . J. a n d Ha ll. \{ . A.. 1 9 9 2 .Da ta Rcp o n :Carlron isotope straiigraphyof Paleogenebulk sedirnents.Hole 76lC
r E\nrouth Platcau.easternIndian(Jccan).In von Rad.U.. Haq. B. Lt..
of the OccanDrillng Program.ScientillcResults.
ct al.. Procecclings
r. 1 2 2 :C o l l e g eS t a t i o n.T X ( Oce a nDr illin - qPr o g r a m )p. . 8 9 7 9 01.
T t rls nr a .R . C l . . 1 9 7 6 .C e n o zo icfb r a m in ile r afio n t th e So u thAtlanttc.
DS D P L c g - 1 6 .l n B a r ke r .P F .. Da lzie l.L \\' . D.. In itia l Re p o rtsof
t he D e e pS c a D r i l l r n gP r o je ct.\' . - 1 6 Wa
: sh in g to n( L L S.Gtxe r n ment
P rin t i n gO l f i c e ) .p . . 1 9 3 - 51 7 .
T . jrls n u . R . ( 1 . . 1 9 8 3 .E o ce n cto N' lio ce n eb cn th ic lb r a n tin ile r sfi orrr
De e p S c a D r i l l i n g P r o.ie ctSit!' 5 1 6 . Rio Gr a n d eRisc. In Barkcr.
P F . . C a r l s o n .R . L . . Jo h n so n .D. A.. e t a l.. In itia l Re p o r tso f the
Dee p S e a D r i l l i n - eP r o ie ct.r ' . 7 2 : Wa sh in g to n( U.S. Go vcr nment
Pri n t i n gO i h c e l . p . 7 3 1 - 7 - 5 5 .
. . C . l n d L o h m a n n .G. P. 1 9 8 3 .Pu lto t' t' tte - Etn t' trbeu tl nul
T jrls rr t a R
tntl ultt'.tsulltt'rtlhit lorunrini.llnt.fhm tlte Atlutttit Ot artrl.\'[icropaleo n t o l o g vS p e c i a lP u blica tio nNo . - 1 .9 0 p p .
A. S.. I 986.
\ rn iV l o r k h o l e nF. . P C . M.. tse r g g r e nW.
. A. a n clEd r va r tls.
Ceno:.oicto.sntopolitutdaclt-ttuter bertltit'lorunittifi'rzl. Birlletin
n Aclu itai ne.
des C c n t r e sc l e R e c h er ch csEr p lo r a tio n - Pr o d u ctioElftr4er.noir
I l. :121pp.
235
V an dcr Zu'aan.G. J. anclJori ssen.F. J.. 1991.B i ofaci al pattcrnsi n
rir er-inducedshelf anoxia.Journal ol the Gettlttg,itulS'oclerrtl'. 58.
p. 65 82.
V i ncent.E . and B run. L.. 1911.P al coceneand earl l E ocenenl i cro facies. benthonic lbraminitera. and paleobathl'metryof Deep Seu
Drilling ProjectSitcs 2-16and 2-17.WesternIndian Ocean.In Fisher.
R . t-.. B uncc.E . T.. et al .. Ini ti al R eportsof the D eep S ea D ri l l ing
P roj ect. r'. 2'+: Washi ngtorl(U .S . Gorernment P ri nti ng Otl i ce).
p. 1359885.
und sei neForami ni f'e renV on H i l l ebrandt.A .. 1962.D as P al aeozaen
tauna i m B ecken l on R ei chenhal lund S al zburg. B al eri sc he
Mathemati sch-N aturni ssensc haft
A kademi e der Wi ssenschafi en.
N euc Fol ge.r'. 108.I 82 pp.
l i che K l asse.A bhandl ungen.
\V al ker.J. C . G. and K asti ng.J. F.. 1992.E ftectsoffuel and l orestco nserration on tuture levels of atmosphericcarbon dioride. PclneoPalaeot'tttlogr;r'. 97. p. l5l 189
Palueotlirnutolrt,e,r.
,q,eogruphr,
Wi dmi i rk. i . G. V . 1995. \4ul ti pl e deep-r'atersourccsand trop hi c
regi mesi n the l atestC retaccousdeep-sea:E l i dence from benthi c
lbraminif'era.Murine Mk ropuleotttologt,v. 26. p. 361 3tt-1.
P l antand mi rmmaldi v erWi ng. S . L.. A l roy. J. and H i ckcl . L. J.. 199-5.
sitl- in the Paleoceneto earlv Eoceneoithc Bighorn Birsin.Pnlrteor'. I l -5.p. l l 7-155
geogrupl n',
P ul ueot'col oqr,
P al aeotl i matol ogt',
Wi ng. S . L.. B ou'n.T. M. andObradovi ch.J. D .. 1991.E arl y E ocenebi '
otic and climatic changc in interior sestern North Arnerica.(ierr1
a.qrtr'. 19.p. l l 89-1192.
Wrshner.K . F..A shj i an.C . J.. C el fhan. C .. C o$i n,s.N {. l \'1..K ann. L..
P cl agi cand
J.. 199-5.
I-evrn.t-. A .. Mul l i neaur.L. S . and S al tzrnann.
bcnthic ccology of the lo$'er interfaccof the EasternTropical Pa
t. '12.p. 93-l l 5
ci fi c oxvgenmi ni mum zote. I)eep-S euR eseurclr.
Wolt'e.J. A.. 199.1.Alaskan PalaeogeneClimates as inf'erredfionr the
In: B oul ter.i !{. C .. and Fi sher.H . C .. eds..C trto
C LA MP database.
:oic Plunt.suntl Clintute tl tlte Arctic, N,{TO ASI Scries. ll7.
Frankfurt:S pri ngerV erl ag.p. 223 138.
Zachos.J. C .. R ca. D . K .. S eto.K .. N omura.R . and N i i tstrma.N ..1991.
Paleogcneand early Ncogenedeep \\'aterpalcoceiinographyof the
Indian Oceanas determineclfiom bcrrthicforatniniler \table cilrb()n
and orygen i sotopcrecords.In D ttncan.R . A .. R ea.D . K .. K i dd. R .
. K .. (eds.).5t;rtl rs.riofr R esul ts
B .. ron R ad. U . and Wei sselJ.
l htnt
Drilling in the huliun Ot rarr. GeophvsicnlMonograph70.
SLienti.fic
U ni on.p.3-51 -185.
Washi ngton.
D . C .: A meri canGeophl -si cal
Zachos.J. C .. Lohmann.K . C .. \l ral ker.J. C . G. anclWi se.S . W.. 19 9]
A brupt cl i matechangernd transi entcl i matescl uri ngthe P al eog enc :
r'. 101.p. l c)l -l l 3
perspecti re.
A r.nari ne
Jtntnrulol 'Geol rtqr',
E vol uti onof ca rl v
Zachos.J. C .. S tott.L. D . and Lohtnann.K . C .. 199-+.
r .9. p. 353-3It7.
Paleot'eurtogrupltt,
Ccncrzoicmarinetemperatures.
236
T HO \1 AS
A p p e n d ix 12.I
Recovert,/
Site
L o ca tio n
Re f- e r ence
D epth
R esol uti on
r-pnr 6rrC
S i ze
H i ch
Y cs
>63
Unc onfbrmi tr
Soutlt Atlutttit'
I .I
oop 689
We d d cll Se a
T h o m a s 1 990a1
I I (X )
Thornasand
Shackleton1996
Medi r.rnr
ul tc ontofl nrt\
1.2
onp 690
WeddellSea
Thomas1990a:
Thomasand
Shac k let o n1 9 9 6
1900
Hi-sh
Yes
>63
Good
2.1
oDp698
Falklancls
Platcar.r
Katz and
Millcr 199I
800 900
Lor.r'
No
> 150
Poor
2.)
onp 702
Falklands
Plateau
Katz and
Miller. lc)9|
I 700- 1800
Low
No
> l-50
Poor
2.3
ot)p7(X)
Falklands
Plat eau
Katz and
M iller 199I
2l-50 2600
Low,
No
> l-50
Poor'
2..1
osor'329
Falklancls
Platciru
Tjalsrra 19771
Tjalsnraand
Lohm ann1 9 8 3
17-10l7.l-5
Low
No
> 150
Mcdiurn
3. I
nsop20C
Mid-Atlantic
Ridge.west
l jalsmaand
Lohrlann 1983:
Miiller-Merzancl
Oberhiinsli199|
ll7-5 3000
Lou
No
> 150.
Poor'
Dailey 1983:
Tjals m a19 8 3 :
Boltovskovet
al. 1995
1000-2000
Lo*
No
>l-50.
> l-50.
>63
Goocl
>63
3.2
osop-516
Rio Grande
Ris c
-+.I
DsDp-52:l
\\alvis Ridge
Clark and Wli-sht
198' 11
Hs u c t a l .
I 98.11
Parkercr
al. I 98-1
3-500-.1000
Lou,
No
> 150
Poor
1.2
osop-525
Walr,isRidge.
Thomasand
Shac k let o n1 9 9 6 :
Boersrra198-l:
Boltol skov and
Bolt ov s k oy1 9 8 9
1600
High
Ycs
>63
Poor'/r.p
lrr
Thomasancl
Shacklcton1996
Boersma198.1
-i.+00
Pa k a n d M ill cr
1000-2000
-+.3
trsop527
Walr'isRidge.
ln core
High
Ycs
>63
Poor/rplrr
l n c ofe
Puci.ftc Ot eart
5
oot'883
No r th Pa cific
Lou
No
>150
199-5
6
osop-577
Subtrt'rpical
North
Millcr ct al. 1987b:
Pa cific
Pa k a n d M ill el l 99l r
P oor/
uncontornritr
I Sm-l100
High/
rneclium
Partim
High
Yes
> 150
Good/gap
unc onl onnl t\
Kaiho in press
7
ttop 865
EquatorialPacific
Braloweret al.
lc)9-5a.b:
Thomas.
unpublis he d a t a .
this chapter
1300 l-500
>63
Good/
unconfbrrnifi'.)
The Late PaleoceneBenthic ForaminiferalExtinction
237
A p p e nd ix 1 2 .I
Postextinction/
Slrort-Ternr
. -t r. t in ctio n
Low Ol/
High
Productivitl'
P()stextinction/
Long-Terrn
r t t t ri i f o r n t i s , I J .r n i d t r ate n .si.i,
B. sin p le . r.
I se lr n e n si.s,
B ul i mi ni ds (8. .setni t'osttrttr).
rt'mpti, B. tlttnelen.si.s,
A. urttgonensis, S. brevisltirtrt.str
S. elegurtta, N. truentpti.
ipet e n s i s N
, . truentpt'i
' . , t rt rii l r t r n i s .
T . se llr cl,sis, B. sir n p le.r,
B ul i rni ni ds (8. .serni L-tL.strttu).
a b yssa m in id s.S. b r e vis l ti rtosti
5. el egurtttt,N . trttetttpt'i
C i bi ci tl oi tl cs spp.N
C i l t i c i d o i d a s sp p ..
-..Lt ttri{forni.s. Gt ntidinoitle.s spp..
N. tnrenpti,
Y es
Y es
truerrt|tti ,
N o'l
Yes
bulinlinids.
No'l
Lenti cul i nu spp . o rurtbottatus
- ir1oi r l e .sr p p .
-. t t t t rii f o r n t i s , P . c o r t e l l i.
A b1'ssani ni ds. C i hi ti tl oi de.s spp..
r , e o a p o n i t l e s sp p .
. l. 1r(.) e r ? r i . N
hul i mi ni ds. N . fruent1tt'i
--. rLt ina n t s .G t n t i t l i r t o i d e . t sp p .
S l i g ht
O. untl xtnrtttts
nc lli, l n l i m i n i t l . t
. , , 1L t t r i 4 f r t r r t r i .P
s ,. c r t r t ' t ' lli,
Y es
O. urrthttnutus
ltrtnitle.sspp.. 5. brzll.r1ruro.r'rr
t c uri i . f b r t r t i . sR, . t u r p e n tie r u e ,
Dissolr.rtion
Ab 1 ' ssa m in id s.N. tr u e mpt'i ,
A. ttr u g ,o r tr llsis
N . l ntenpt'i , O. unbotktl us,
'f. sel tnertsi s,
A . tra,qortert;i s,
N o'l
N o'l
B .;tttti tt'.:!Ltl ,t
. , t c uri 1 f p l r t i 5 , b u l i r n i n i c ls.
' . t t loit l e . ss p p . . a g g l u t i n an ts
Ab - vssa m in id s.N. tr u e mpt'i ,
A byssarrri rri ds.N . truentpti ,
,1 . tr n tg u r cn sis
A . urugonettsi i , bul i rni ni ds
N o'l
l
N. truentpti,O. tortbonutus,
P.proli.ut
,\. unt,qurcn.si.s,
No'l
Yes
'l
N. Iruentpri,abyssan'rinids.
Il. t ri hedru. 7'.seIntettsi.s
Yes
Yes'l
N. tr u e m p ti. a b yssa n ri ni cl s.
N . truentpt'i , bul i mi ni ds.
No
Yes
A. r tr tr g orten.si .s
T ..se h n e n .sis,
C i bi ti tl ol rl e.r spp.
i, t t c or i i f o r n r i . s .A . t ' e l t t s ttte n sis.
N. tr tte r n p ti, a b yssa m i ni ds.
N . trttctnpti . bul i rni ni ds.
No
Y cs
: rt rern 1 t t i ,B . t h u n e t e n . s i.s
T ..se lttte n sis
C i l ti ci dtti des spp
I
N . l ruentpt'i , A . urugonertsi s.
N o'l
'l
N . tntempyi . bul i nl i ni ds
Y es
S l i ght
C i l ti ti tl oi ti e.s spp. N . l rrrerrl Tr,r'i ,
bul i rni ni ds (Il . sotti ttt.sttrttr\
Y es
S l i ght
't ' Lt uri i f b n t t i s . P . t o n a l l i,
. / rrit o a l i s t s ,S . l t r e v i s l t i ttttstt.
: t 'I it t t t u I u s , N . t n t e r n T t t i
,t t t uriilrtrnri.s, Ncoeponitle.s spp..
-: . Lrt ina n t s
: . LLt ar i | l o n n i . s , N . t r u e r np ti.
N e o e p o n i t l c s sp p ..
--: lut ina n t s .
:'r, llolrlr,s spp.
: bat (t rr i i f b r r n i s . B . r n i d t tu tcr t.sis.
l b.vssarni ni ds
-rlut inan t s . I ' . t o r t ' e l l i
: hc t 'c rr r i | f l t r n r i s ,P . c o n e l li,
', t , lrt . tt ' o e n s i . s . B . r n i t l t o ve r t.ti.s
' ! T ..se lr r te n si.s,
A. u r u gttnen.si .i ,
Ncr te p o r r r r lt' .tsp p .
: lrt , c t t tr i i f o r r r u . r , . l . r ' t , 1 a .r tr ;r ,r r .r l.r , T ' .se lm e n .si.t,
B. se n ti( o.\l (tt(t
. \ (, nt i(r i b r ( i u g a g , t l u t i n a n ts
238
THOtaAS
Appendix 12.l ( conrinued)
R ec ov ery /
Site
L o ca tio n
Re f' e r e nce
D epth
R csol uti on
l prl r 6l rC
S i ze
U nc onformi ty
Indiun Oceart
8.1
oopJlJ
Kerguelen
Mackensenand
Berggren1992
2000-3000
Low
No
>l-50
good/
unconformitl
8.2
oop 7,18
Kerguelen
Mackensenand
Berggren1992
600-2000
Low
No
>150
Poor
9
oup 738
Kerguelen
Lu and Keller 1993; 13.50
Nomura and Kennctt
personalcomnrunicationlThomas
unpublished
data
High
Ycs
>150.
>63
poor/
lprlt in
core
l0
osop2:15
SouthMadagascar Sigal 197'1
Basin
600-1000
Low
No
>150
Poor
II
osop237
MascarenPlateau
Vincentand Brtin
191I
600 2500
Low
No
> 150
Poor
12
osop2l-5
NinetyeastRidge
McGowran l97,ll
Hovan and Rea
1992
600-2500
Low
No
>150
Good/lprivt
not tn corel
13
oop 752
Kerguelen
Nornural99l ;
-500-1000
Nonruraand Kennett
personalcommunlcation
High/
medium
Partim
> 150
Poor
l,t
r:tov762
EasternIndian
Ocean
Nonruraand Kcnnett 1000-1500
pcrsonalcomnrunication:Thomasunpublisheddata:
Thomaset al. 1992
High/
medium
Partirn
>150.
Poor/
disturbed
>63
North Atlantic
l-5.1 osop-5:19
Bav of Biscay/
GobanSpur
Reynolds1992:
BerggrenandAubry
l 996
600-1500
Lol'n'
>l-50
Yes/no
benthics
Good/
unconlormtty
HiglV
medium
Partim/
no/
no
yes
>1 . 5 0
Go o d /
r.rnconlormity
15 .2 DSDp :10 1 Bay of Bis c ay
Pak andM i l l e r 1 9 9 2 : 1 8 0 0
Schnitker1979:Saint
M ar c l99la . 1 9 9l b ;
Pardoet al. in press
15.-l rrsr;r''1004' Bay of Biscay
Schnitker1979
3100 .+2(X)
Low'
No
> l -50
Poor
15..1 r;sop5:1[J
GobanSpur/
Bay of Biscav
Boltovskoyet al.
1992
600-2000
Medium
No
>6]
good/post
l6
New Jcrsey
margin
Hulsbos1987:Saint 1800-2300
Marc 19871l99lb:
Thomas,unpublished
data
Low/high
No
>150.
Good/
uncontblrrrrtl'
nsor,605
>6-l
The Late PaleoceneBenthic ForaminiferalExtinction
239
\ppendix l2.l ( tontinuetl )
-Extinction
:':.c
T.rra
Postextinction/
Short-Term
'i. bet'carii. ortni.s, Cilticidoide.r spp..
P c onelli , N . I r u e r r t p t i
'l
Postextinction/
Long-Tcnl
N. truentpti. Cibit'idoitle.s spp..
Low O2l
High
Productir,'ity
Dissolutitttt
No'l
O. Ltmborttttus
N o'l
:
N. truentpti, buliminicls.
Cibicidoide.sspp
Yes
Yes
G . belt or i i l o r m i . s , A . t ' e l u .st' o e n sis.
') T. .sel ntensi s,
A . onL,e.utert.si s,
Y es'l
). weller i , B . t h u t e t e n s i s
N trrterrtl 'ti
j. hettttriifitrmis,
N. truentpti. Attontttlinttitles spp.
No'l
T. selntensi.s,
Arugoria spp.
Yes'l
Atromulirutitles
N. truenrytvi,
durtit'trs,
O. rurtbortatus
Yes'.)
N. Iruentpti, buliminids.
Cibicidrtidesspp.,A. azrgrrtrerisis,
O. ruttbttrtcttus
No
'
j. bec c u r i i l o m r r s , b u l i r n i nid s
?
Stilosttntellu spp.
Lenti<'ulirut spp.
,'. bet'curii.forntis.
N. truentpti.
3. tIuneten.si.s,
agglutinands
7'.selntensis.buliminids
N. truentp.t I
.. lt t't t'u ri i.lbrnt i.s,N. t ruentpti,
. . : ! lut ina n t s
r'. bec t a r i i . f o r n i s , P . t o r y eIIi,
( il, irirl, ti l e . t . p p . . N . I n ! ( t ttlt\ i.
bulirninid s
N. truentpt'i,
G. bet.r.urii.fbnni.t,
spp..B. thanetensis,
Cibicidctitle.s
B. tlelitunilu.s
N. truentpti, B. sentito.slttttt
Cibitidoides spp..
'l'..selntensis
G. becca r i i.fonnrs,a-e-ulutinants
i.s
Bulinrinids.T. seItnert.s
Yesl
Ycs
G. bet'turiiforni.s,C. lrt'plnlu.r.
(1.ntitlxut'ettsis,
B. delitutulus,
Neoeporti c!e.sspp.. G r roi d i ruti de.r
') N. truentpti, ab1'ssaminicls.
buliminids(8. senricostutu)
N. lruempti, abvssaminids.
bulinrinids.Cibicitlttides
spp.
Ycs
Yes
Cibicidoide.sspp..N truenrytt'i,
G: rrti tl i rtoi tles spp.
abyssanrinids.
l
Cibicidoidesspp..Aragonia
a rugortettsi.s.
Ci bit'idoi des spp.
Yes
Yes
No data
u,
T. selnrcrtsi.s,
S. brevi.spirto.s
A. uragonertsis,B.
seilltcostutu
N. truentpt'i,buliminids.
Yes
Yes
'l Probabll'unconformtty
N. trtrenpti. Cibit'idoide.sspp..
Yes
Yes'l
G. beccuriilortni.s.Cibititloide.!spp..
A . t 'eI a s c o en s i . s B
, . n t i t l t t u yen .st s
P . t t t ne I I i , a g g l u t i n a n t s
C i hi t i tl t,i tl t'r :pp.
abyssami ni ds. T. seI tttertsi s,
bul i rni ni ds (B . setni t ostu)
THOf'1AS
240
App e n d ix 12 .2
Country
North Atlarttir'
Spain
A.l
Section/
Formation
LPTM
References
Depth
Resolution
6llc
Size
Lithology
Llmaya(Atlantic)
Ortiz 1995;Canudoet al.
1995:Molina et al'
von Hillebrandt
199'11
1965
600-1500
High
Yes
>63
Marl' calcarenite
turbidites
A.2
Spain
TrabakuaPass.
Ermua.west
Pyrenees
Coccioni et al. 1994:OrueExtebaniaet al in press
800-1000
High
No
>100
Limestones/marls
A.3
Spain
'l'remp/Campo
?50-500
Low/
medium
No
>150
Maris
(Ilerdian stratotype)
Molina et al. 1992:Ortiz
and McDougall l99l ;
von Hillebrandt 1965
B
France
ParisBasin
Le Caivez 1970
10-200
Low
No
>150
ClaYs/silts
c
Trinidad
Lizard Springs
Formation
Beckmann1960;Bolli
et al' 1994
500-t500
Low
No
>150
Marls
D.I
usA
New Jersey;
Vincentown
Manasquan
Formattons
Gibsonet al. 1993:Gibson
andBYbell 1995
50-200
Low
No
>63
Clayeysands/
claYs
D.2
USA
New Jersey;
Vincentown
Manasquan
Formattons
OlssonandWise 1987
50-250
Low
No
>150
Silty sands/clays
E.l
NW
Europe
North Searegion
King 1989
100-500
Low
No
>150
Sand/clays
8.2
NW
Europe
North Searegion
Gradsteinet al. 1994
750-1000
Low
No
>150
Sand/clays
F
Spain
caravaca(Tethys)
ortiz 19951Canudoet
al. 1995
500-1000
High
Yes
>63
Marls' limestone
G
Israel
Nahal Avdat (Negev)
Speijer 199'1
500-700
Low
No
>125
Marls/limestone
H
Egypt
Abu Rudeis- Wadi
Nukhl (Sinai)
Speijer1994
500-700
Low/
medium
No
>125
Marl/limestone
I.l
Egypr
GebelAweina
Speijer1994:Speijeretal.
1996a:Charisi et al'
1995:Schmitzet al' 1996
100-150
Low/
medium
Yes
>125
Marl/limestone
1.2
Egypt
Bir lnglisi = Gebel
Duwi
Speijer1994;Speijeret al.
1996b;
Schmitzet al. 1996
50-200
No/
Low/
rnedium
Yes
>125
Marl/limestone
Tetht's
J .lltalvBellu no DiNapoliet al. lgT0500_1500Low N o >l 5 0 M a r l s / s c a g l i a s
The Late PaleoceneBenthicForaminiferalExtinction
241
A p p e ndix 1 2 .2
.::-ExtinctionTaxa
Postextinction/
Short-Term
Low O./
Postextinction/
Long-Term
High
Productivity
Dissolution
. utccariifonnis, Cibici.doides,
- - rnrinids,P. con'elli
First agglutinants,O. umbonatus,
IhenB. tu.rpanten.si.s
M truempyi,abyssaminids,
O. ttmbonutus,buliminids
Yes
Yes
. ueccrtriiJbrmis,
N. truempl.i.
,::lutinants,buliminids,
-, t nalino ide sv'elle ri
Rare agglutinants.then small
buliminids.N. truempt'i
A. di.ssonata,
N. trttemp.vi,
buliminids.O. umbonattrs
Yes
Yes
. ,:rbigirtosus,Bulimina trigonalis,
: ,,r'telli, agglutinants
Cibicidoidesspp.,Ut,igerina.
spp.,Hanzuv'ttiaspp
?
'!
.. ,telphidiunt,pulsiphottina,
;.t,idoides,buliminids
Buliminids, Cibicidessuc'-.
cedens,Bolitintt spp.
Yes?
?
- . r'lnscoensis,
B. mitlwayensis,
-.,.rnrinids,
Cibicidoidesspp.
N. truempyi,T. seltnensis,
(buliminids)
B. semicostata
Yes?
'!
..:.elinella spp..Cibit:itloidasspp..
- : 'tualinoidesspp.,Ut'igerina
T. selmensis,Epistominellusp.A
A., buliminids, Pseudouvigerina spp.
T. selmensis,Pulsiphonina
prima, Arutmalinoid€.tspp..
EPistominelln
Yes
Yes
:...elinellcrspp.,Anomalinoidesspp.,
- :':cidoidesspp.
Agglutinants,T. selmensis
Epistominellasp.,T. robertsi,
N. truempti
Yes
Yes
;.et.t'ttriiformis,
B. mitlw'atensis,
' ' inincr trigonalis, Cibicidoides
- .htntmina ruthyenmurrd)'i,
Low-diversity agglutinants
T. brevispira,Gttudryinahiltermcrzni,buliminids
Yes
Yes
Low-diversity agglutinants
T. ntbertsi,R. amplectens,
R' intermediaassemblage
(agglutinant)
Yes
Yes
RareHaplophrugmoides(barren),
T. selmensis,buliminids.
A. aragonensts
Buliminids, N. truempti,
Anomalinoidesspp.
Yes
Yes
M truempt'i,Cibicidoides
spp.
Yes?
Yes
paupera,
' - ::, tilrrythragtniunt
,..:nrblage(agglutinant)
:
: ;,t'c't'ttriifonnis,
A. t,elascoensis,
,.t-r.elli,Gt,roidinoidesspp.
: ueccrrriifolnls,agglutinants,
,,nelli, buliminids, Cibicidoides
' ii
:
: ht't.t.uriiJormis,
agglutinants,
: .,,r't'elli,buliminids,
.- "icidoides spp.
(?) Anomolinoidesspp.,vaginulinids,Stainforthiaspp.,
buliminids
T. truempti, Cibicidoidesspp.,
buliminids
T selmensis,
Yes
Yes
.- iit'irloitles spp.,lagenids,C. midA. avnimelechi
,;rgrr.sis,
(?) B. callahani, Cibicidoidesspp..
P. v'ilcoxensis
A. aragctnensis.
Buliminids, Cibicidoidesspp.,
Artornalinoidesspp.
Yes
Yes
:,:rtnalinoitles, spp.,Valvulinerlaspp.,
Anomalinoidesaegrptai(us,
Lenticulina spp.,L. applinae
Vtlt'ttlineria scrobiculata,
Cibicidoidesspp.,Anomalinoide.sspp.,B. .foraJraensi s
Yes
Yes
ctpplinae
-. ,.tttstomoide.s
:.:gltiinants, N. truempti, G. bec-
I B. semicostata,B. trinitatettsis
N. truempti, Anomalinoides
spp.,buliminids
Yes?
?
buliminids,A. velusco.;rii.fctnnis,
,' :ili. P co n-e lLi
I HO XA S
242
Appendix
12.2 ( corttirtuetl )
LP T\I
Se ctio n /
C o u n tr y
F o r m a tio n
R ef'erences
D epth
R esol uti on
bl 'rc
S i ze
Li thol ogl
J.2
Italy
Possagno
Bragact al. 197-5
500-1500
Lor','
No
>l-50
Marls
K
Austria
Reichenhall
SalzburgBasin
von Hillebrandt1962
200-800
Lo',v
No
>l-50
Silts/marls
L
New
Zealand
Tawanui
Hornibrooket al. 1989:
Kaiho et al. 1993.in press
-500-1500
High
No
>63
Siltstone.siltr
limestone
M
Japan
Hokkaido.Tokachi
district
Kriiho 1988
500 1000
Low
No
>63
Silty marls
Pat'i.fic
The Late PaleoceneBenthic ForaminiferalExtinction
243
\ppendix 12.2 (c ont inued )
Postexti
nction/
Pr.'-ExtinctionTaxa
'i bc<curiiforntis,
N. truetrrlttl,
.,_rglutinants.
buliminids,A. yelas,,ertsis,
P. coryelli
Short-Term
M tntetnpti,O. tonbonutus
\uglutinants.buliminids.nodosariids.
-'nticulinids
,i. betcuriifbrnrl.s.agglutinants.
t--.tnidwoyensis,
A. ruhigirtosus
\ velleri, agglutinants.B.
' l t t / r |d t ' e l i sts
First: rg.tu.rptunen.sis.
agglutinants
only
Second:a-ggllrtinants
Low O,/
Postextinction/
Long-Term
High
Productivity
Dissolution
N. truem1tt'i,
B. senicostutu,
buliminids
Yes?
Yes
Agelutinants.
Anomalinoides
rubiginosus
?
?
Cibicidoide,sspp..O. umltonalrr.r,agglutinants
Yes
Yes
Agglutinants.buliminids
Yes
Yes