Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319

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High-resolution planktonic foraminiferal analysis from the

Cretaceous^Tertiary boundary at Ain Settara (Tunisia):
evidence of an extended mass extinction
Valeria Luciani 
Dipartimento di Scienze della Terra dell’Universita' degli Studi di Ferrara, Corso Ercole I d’Este 32, 4410 Ferrara, Italy
Received 25 July 1999; accepted 9 August 2001

Abstract

The Ain Settara section, located in the Kalaat-Senan area of north^central Tunisia, spans the Cretaceous^
Tertiary (K^T) boundary which is characterized by a red layer and a thin non-bioturbated boundary clay. Sediment
accumulation across the K^T boundary at Ain Settara was probably interrupted by three short hiatuses and/or
condensed sedimentation. The first hiatus occurs at the top of the CF1 Zone; the second hiatus/condensation
occurred just below the boundary clay and the third hiatus at the P0/P1a boundary, in the earliest Danian. These
hiatuses are marked by weak unconformities, bioturbation and sudden disappearances/appearances of species which
are known to disappear/evolve sequentially in continuous sections. Quantitative high-resolution planktonic
foraminiferal analysis across zones CF1(upper), P0 and P1a(1) reveals an extended and selective mass extinction.
All 41% of the species which disappeared at or below the K^T boundary are rare to very rare and primarily
ecologically specialized keeled deeper-dwelling tropical^subtropical forms (Globotruncana, Globotruncanita, Gublerina,
Planoglobulina, Rosita (Contusotruncana), Racemiguembelina). Their combined relative abundance varies between 10%
and 15% of the total population at the end of the Maastrichtian. The K^T crisis thus appears more catastrophic when
viewed in tropical^subtropical assemblages and based on analysis of larger species ( s 200 Wm) which preferentially
includes the more specialized forms, though, in fact, the K^T mass extinction actually involved a relatively small part
of the foraminiferal population in terms of relative abundance. The pattern of extinction and changes in dominant
population at Ain Settara appear to be very similar to the planktonic foraminiferal turnover of the other north^
central Tunisian sections (El Kef, Elles). The selective mass extinction pattern suggests that the catastrophic effects of
the bolide impact superimposed those related to long-term environmental changes, such as variations in temperature,
sea-level and associated water-mass changes. ß 2002 Elsevier Science B.V. All rights reserved.

Keywords: Cretaceous^Tertiary boundary; hiatus; mass extinction; biostratigraphy; planktonic foraminifera; central^northern
Tunisia

1. Introduction

The Cretaceous^Tertiary (K^T) boundary re-

* Fax: +39-0532-206468. cords a catastrophic crisis in the history of life
E-mail address: lcv@unife.it (V. Luciani). on Earth and is one of the most investigated

0031-0182 / 02 / $ ^ see front matter ß 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 1 - 0 1 8 2 ( 0 1 ) 0 0 4 0 0 - X

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300 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319
stratigraphic intervals. It is now commonly ac-
cepted that the impact of a large meteor caused
a sudden mass extinction (Alvarez et al., 1980).
The circular gravity anomaly structure near
Chicxculub in northern Yucatan (Mexico) has
been identi¢ed as the impact crater (e.g. Pope et
al., 1991; Hildebrand et al., 1991; Swisher et al.,
1992; Sharpton et al., 1992, 1993, 1996). Though
it has yet to be demonstrated whether this crater
is of K^T boundary age (Keller et al., 1993b;
Ward et al., 1995), and whether a meteoric impact
was the sole cause of the biotic crisis.
Among marine organisms, planktonic forami-
nifera which are generally abundant in relatively
shallow to deep marine environments across lat-
itudes are one of the most useful tools for inves-
tigating the nature of this crisis. Although it is
evident that this group su¡ered a mass extinction,
the detailed studies carried out in recent years
demonstrate that the boundary turnover is quite
intricate and composite. A large amount of data
from di¡erent latitudes, depositional environ-
ments and paleodepths document complex extinc-
tion patterns, with deeper-dwelling, complex trop-
ical^subtropical forms more a¡ected by the crisis,
including some forms disappearing before the
K^T boundary, and other forms, among the less
specialized species, constantly found into the
Danian and mainly considered as survivors (e.g.
Keller, 1988a, 1989a,b, 1993, 1996; Keller et al.,
1993a, 1995; MacLeod and Keller, 1994; Pardo et
al., 1996, 1998; Arz et al., 1999).
Although the end of the Maastrichtian was a
time of stressful environmental conditions charac-
terized by rapid climatic changes documented in
marine realm (e.g. Barrera, 1994; Li and Keller,
1998c), relatively few studies have investigated
the biotic response to these long-term changes
(e.g. Abramovich et al., 1998; Li and Keller,
1998a,b,c). The related variations in water-mass
strati¢cation may have pre-disposed populations
to the mass extinction or may have been contrib-
uting causes, albeit probably not the main perpe-
trator of the crisis. To date, neither impact, vol-
canism, nor any other long-term factors has
de¢nitively been linked to the extinction crisis as
testi¢ed by the continuing intense scienti¢c de-
bate. Correspondingly, the controversy regarding
PALAEO 2759 1-5-02
gradual versus simultaneous disappearances of
planktonic foraminiferal species across the K^T
boundary, and hence whether a single impact or
multiple events caused the mass extinction, is still
ongoing.
Tunisian K^T boundary sections with their
high sediment accumulation rates and relatively
continuous sedimentary records have contributed
signi¢cantly to our understanding of this mass
extinction in planktonic foraminifera. The El
Kef section was formally designated the K^T
boundary Global Stratotype Section and Point
(GSSP) (XXVIIIth International Geological Con-
gress, Washington, DC, USA, 1989). As a result,
the section has been extensively studied based on
closely spaced sampling of various microfossil
groups, geochemistry, and stable isotopes (Jiang
and Gartner, 1986; Posphical, 1994; Perch-Niel-
sen, 1981a,b; Perch-Nielsen et al., 1982; Smit,
1982; Donce et al., 1982, 1985; Peypouquet et
al., 1986; Brinkhuis and Zachariasse, 1988;
Brinkhuis and Leereveld, 1988; Me¤on, 1990;
Smit and Tenkate, 1982; Kuslys and Kraehen-
buehl, 1983; Keller and Lindinger, 1989; Robin
et al., 1991; Keller, 1988a,b, 1992; Ben Abdel-
kader, 1992; Keller et al., 1995).
In order to increase the low latitude planktonic
foraminifera dataset and better document the
catastrophic e¡ects of the K^T mass extinction,
other Tunisian K^T transitions were closely
sampled during the Workshop on the K^T tran-
sition, held in May 1998 and made available to
interested parties. This study evaluates one of the
sections, Ain Settara, based on high-resolution
planktonic foraminiferal analysis, similar to that
previously published for El Kef (Keller et al.,
1995) and Elles (Karoui-Yakoub et al., 2002; Kel-
ler et al., 2002) and hence allows for close corre-
lation and comparison with these sections (Fig. 1).
The speci¢c aim of this study is to evaluate the
nature and extent of the mass extinction and the
completeness of the sedimentary record.
2. Location and lithology
The Ain Settara section is located in the Atlas
mountains of north^central Tunisia (Kalaat-Se-
 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319 301

nan area) about 50 km southeast of the El Kef

Fig. 1. Simpli¢ed paleogeographic map of central^northern
Tunisia during the Late Maastrichtian^Early Danian showing
the depositional setting of the Ain Settara section (modi¢ed
after Burrolet, 1967). Isolines indicate depth in meters.
section and west of Elles (Fig. 1). The paleogeo-
graphic setting of these three sections is similar;
all are located on the continental shelf, but Ain
Settara (and Elles) was deposited in a slightly
more proximal position to the Kasserine Island
and at a shallower depth (middle to outer neritic)
with respect to El Kef (Figs. 1 and 2).
An interval of 2.2 m spanning the K^T bound-
ary was sampled at 5^10 cm intervals, except for
the boundary clay where samples were taken at
0.5^2 cm intervals. Sediments of the uppermost
Maastrichtian consist of gray silty marls. The
K^T transition is marked by a lithological break
from silty marl to a thin dark clay layer with a 0.5
cm thick red oxidized layer at its base. A similar
thin red layer is present at the base of the bound-
ary clay at El Kef, as well as at Elles, El Melah
and in all relatively complete K^T boundary sec-
tions, and contains the maximum Ir concentra-
tions and Ni-rich spinels (Rocchia et al., 1995;
Robin et al., 1995).
Details of the lithological K^T transition at Ain
Settara are schematically shown in Fig. 3. Only
the top 60 cm of the Upper Maastrichtian gray
silty marls was sampled due to the steep slope and
debris cover (Fig. 2). At the top of this interval,

Fig. 2. The K^T transition at Ain Settara is located on a steep slope consisting of soft marls and marly shales. The K^T bound-
ary itself is not marked by any major lithological changes, but from a distance is evident by a darker gray clayey shale layer. Up
close, the K^T boundary is marked by bioturbated gray marls underlying a bioturbated dark silty shale (2 cm thick), followed
by a 0.5 cm thin red layer containing the maximum Ir anomaly, a 2 cm thick dark clay (not bioturbated) and a thin (1.5 cm)
‘tempestite’ layer. Upsection, sediments consist of about 1 m of dark gray shaley clays followed by gray shales.

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302 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319

Fig. 3. Stratigraphic column of the Ain Settara section across the K^T transition.

and about 2 cm below the K^T red layer and dark
clay is a weak disconformity marked by truncated
chondrites burrows. A 2 cm thick dark silty shale
overlies the disconformity. The top of this silty
shale is also marked by chondrites burrows
some of which are vertical, truncated and in¢lled
with the overlying dark clay. This suggests anoth-
er disconformity is centered at the K^T boundary.
No burrowing is observed across the red layer
and in the dark 2 cm thick boundary clay. Above
the boundary clay layer is a 1.5 cm thick silty
layer containing diagenetic calcite; this silt layer
may be interpreted as a small storm deposit (tem-
pestite?) (see also Dupuis et al., in press). The
presence of reworked transported species in the
sample from this layer con¢rms the resedimented
nature of this bed. The erosional event associated
to this bed could be responsible, at least in part,
for the unusually thin boundary clay in this sec-
tion as compared with Elles and El Kef. Above
the probable tempestite, the 70 cm sediments an-
alyzed grade from dark gray ¢ssile shaley clay to
gray (sometimes silty) shale (Fig. 3).
The K^T succession from Ain Settara has also
been studied by Molina et al. (1998), Arenillas et
al. (2000) and Dupuis et al. (in press). However,
their study was based on a composite of litholog-
ical features observed at several outcrops along
strike, rather than a single transect as in this
study. Since there are signi¢cant lithological var-
iations along strike at Ain Settara, this may
largely explain the di¡erences in the stratigraphic

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 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319 303

Table 1
Relative percent abundances across the K^T boundary at the Ain Settara section, calculated for the s 38 Wm fraction on counts
of 300^500 specimens in each sample (x = rare species, 6 1%)

columns and species ranges between this study
and theirs. Other di¡erences may derive from
the di¡erent sizes of the populations analyzed:
larger than 38 Wm in this study (see below) and
larger than 63 Wm from the other authors.
3. Methods and materials
Samples were disaggregated in dilute H2 O2
(20% vol.) and washed through a 38 Wm screen
in order to avoid the loss of very small species

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304 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319

Fig. 4. Commonly used planktonic foraminiferal zonal schemes across the K^T transition. The datum event sequence is referred
to the data from the most complete K^T boundary sections (MacLeod and Keller, 1991). Zonal indices are in bold type. The
zonal scheme of Keller (1993) and Keller et al. (1995) is followed for the Ain Settara section.

earliest Danian planktonic foraminifera. Keller 4. Biostratigraphy

(1993) and Keller et al. (1995) have shown that
the earliest Danian species are only present in the
smaller (38^63 Wm) size fraction. In addition, rel-
ative species abundances may di¡er widely be-
tween the s 63 Wm and the s 38 Wm size fraction.
The washed residues were dried in an oven at
50‡C. Ultrasonic treatment was not necessary be-
cause the microfossil assemblages proved to be
clean and well preserved. Only in the upper part
of the section are planktonic foraminiferal tests
partly recrystallized.
Population counts are based on representative
splits of 300^500 specimens of the s 38 Wm size
fraction. The remainder of the residue was care-
fully searched for rare species. Results of the
quantitative analyses are listed (in percent) in
Table 1.
The great interest in the K^T boundary in re-
cent years has provided a large database from
di¡erent latitudes and geological settings. These
high-resolution studies have also enhanced the bi-
ozonations across this interval, particularly for
the Lower Danian interval (Smit, 1982; Ben Ab-
delkaber, 1992; Keller, 1988a, 1993; Keller et al.,
1995; Pardo et al., 1996; Berggren et al, 1995;
Olsson et al., 1999). Although high-resolution
studies for the Late Cretaceous are at present
less numerous, the standard zonal schemes (e.g.
Caron, 1985; Robaszynski and Caron, 1995;
Berggren et al., 1995) have recently been im-
proved for the Upper Maastrichtian (from about
70 to 65 Myr; Li and Keller, 1998a).
Fig. 4 shows a comparison between the most

Fig. 5. Planktonic foraminifera species ranges during the uppermost Maastrichtian and earliest Paleocene at Ain Settara (Tunisia).
Note the gradual and extended disappearances of species across the K^T boundary, and the great number of species still present
in the earliest Tertiary. The percentage of disappearing species with respect to the total number is also indicated.

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 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319 307

commonly used biozonations across the K^T pearance of Parasubbotina pseudobulloides

boundary and the main planktonic foraminiferal
events. The use of the standard total range Aba-
thomphalus mayaroensis Zone for the uppermost
Maastrichtian presents some problems because
the zonal marker is often rare or absent at the
top of the Maastrichtian, particularly in relatively
shallow-water sections and tropical regions, and
¢rst and last appearance datums are diachronous
across latitudes. This is because A. mayaroensis is
a deep-water species and generally more abundant
in higher latitudes where it appears earlier (Keller,
1988a; Huber, 1991; Masters, 1994; Barrera,
1994; Canudo et al., 1991; Olsson and Liu, 1993).
In the low latitude Tethys, the uppermost
Maastrichtian sediments are characterized by the
range of Plummerita hantkeninoides, or Zone CF1
(Plummerita reicheli of Masters, 1993, 1984; Kel-
ler et al., 1995; Pardo et al., 1996). Though this
marker species may be too rare or absent in some
localities at the top of the Maastrichtian. How-
ever, even in the absence of P. hantkeninoides, the
top of CF1 which marks the K^T boundary can
be easily identi¢ed in low latitudes by the disap-
pearance of the largest and most complex globo-
truncanids. Biozone CF1 spans the upper half of
C29R below the boundary, or the last 200^300
kyr of the Maastrichtian based on the Berggren
et al. (1995) time scale. Keller (1993) and Keller et
al. (1995) distinguish the P0 and P1a zones in the
lowermost Danian (Fig. 4).
Zone P0 generally spans the boundary clay,
from the LAD of Plummerita hantkeninoides
(and the larger, more ornate globotruncanids) to
the FAD of Parvularugoglobigerina eugubina and/
or Parvularugoglobigerina longiapertura. Zone P1a
corresponds to the total range of P. eugubina. The
zone has been subdivided into two subzones,
P1a(1) and P1a(2), on the basis of the ¢rst ap-
(Fig. 4). At Ain Settara the CF1, P0 and P1a(1)
zones have been recognized (Fig. 4). Taxonomic
criteria are mainly based on Robaszynski et al.
(1983^1984), Caron (1985), Nederbragt (1991)
and Keller et al. (1995).
4.1. Uppermost Maastrichtian Plummerita
hantkeninoides (CF1) Zone
The Plummerita hantkeninoides (CF1) Zone was
recognized at Ain Settara below the red layer and
boundary clay. The 60 cm analyzed below the
boundary clay does not encompass the base of
the zone, because the marker already occurs in
the basal sample. Abathomphalus mayaroensis is
absent in this interval, possibly because of the
relatively shallow-water environment at Ain Set-
tara during the latest Maastrichtian. A total of
57 species are recognized at the top of the Maas-
trichtian, in the Plummerita hantkeninoides (CF1)
Zone (Fig. 5). (P. hantkeninoides was distin-
guished from Plummerita reicheli based on the
criteria given in Pardo et al. (1996).
Relative species abundances indicate that the
bulk of the population consists of small relatively
unornamented heterohelicids (particularly Hetero-
helix navarroensis, Heterohelix globulosa, Hetero-
helix dentata and Pseudoguembelina costulata,
Fig. 5). Hedbergella monmouthensis and Globige-
rinelloides aspera are also quite common. Large
keeled forms and complex heterohelicids represent
a minor part of the population. Guembelitrids,
including Gu«mbelitria cretacea, are not abundant
in the uppermost Maastrichtian (less than 10% of
the total population).
Six species (Globotruncana insignis, Globotrun-
cana duwi, Rosita (Contusotruncana) contusa,
Pseudoguembelina hariaensis, Heterohelix morema-

Fig. 6. Relative abundance changes of planktonic foraminifera species across the K^T boundary in the s 38 Wm size fraction at
Ain Settara. Note that the species disappearing at or below the K^T boundary are relatively rare, their total relative abundance
being less than 15% of the total fauna. The disappearing species mainly belong to the larger, more ornate, tropical^subtropical
and deeper-dwelling group. The biserial heterohelicids which represent the latest Maastrichtian dominant population (in particular
Heterohelix navarroensis, Heterohelix dentata, Heterohelix globulosa, Pseudoguembelina costulata) decline above the K^T boundary,
and are substituted by the triserial low-oxygen-tolerant guembelitrids. The relative abundances of the new Tertiary species are low
at ¢rst; their percentages increase when guembelitrids decline, suggesting the progressive recovery after stressful environmental
conditions.

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308 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319
ni, Globotruncanella citae) disappear simultane-
ously below the weak unconformity above sample
4 (Fig. 5). In the 35 cm below the unconformity,
another six late Maastrichtian species are rare and
sequentially disappear (Globotruncanita angulata,
Globotruncana falsostuarti, Heterohelix planata,
Plummerita reicheli, Gublerina robusta, Gublerina
cuvilleri).
The K^T boundary is placed at the thin red
layer at the base of the boundary clay, which is
a characteristic lithostratigraphic marker that gen-
erally contains maximum Ir- and Ni-rich spinel
concentrations (Robin et al., 1991). Twelve spe-
cies disappear at this interval and another seven
disappear in the overlying clay and silt layer (tem-
pestite, Fig. 5).
4.2. Lowermost Danian: P0 Zone and P1a(1)
subzone
Zone P0 at Ain Settara coincides with the thin
(2 cm) boundary clay. The ¢rst Tertiary species
are present in this clay. They are represented by
very small, rare Globoconusa conusa, Woodringina
hornestownensis and Globanomalina archaeocom-
pressa (Fig. 5). In most expanded boundary clays
these species have been recorded as appearing in
sequence (e.g. El Kef stratotype, Keller et al.,
1995). 56% of the latest Maastrichtian species
are still present in P0 Zone (Fig. 5).
The P0^P1a zonal boundary is marked by the
¢rst appearance of Parvularugoglobigerina eugubi-
na and/or P. longiapertura. These species ¢rst ap-
pear just above the boundary clay, together with
Eoglobigerina eobulloides, Eoglobigerina simplicis-
sima, Eoglobigerina edita, Eoglobigerina fringa. In
other K^T sections these species ¢rst appear se-
quentially (Olsson et al., 1999) (Fig. 4). At the
studied interval only the lower part of P1a(1) sub-
zone is represented as evident by the fact that the
FAD of P. pseudobulloies is not encountered, and
only those events generally occurring at the base
of the subzone have been identi¢ed, such as the
¢rst occurrences of Chiloguembelina midwayensis
and Globanomalina taurica (Fig. 4) (Olsson et al.,
1999). Relative abundance of eoglobigerinids and
parvularugoglobigerinids is low at the base of
Pla(l) and increases upsection where Cretaceous
PALAEO 2759 1-5-02
species, in particular guembelitrids, decline. A
marked decrease in abundance of Globoconusa
conusa also coincides with this level (Fig. 6).
5. How complete is the Ain Settara K^T
transition?
The interval across the K^T boundary is mag-
ni¢ed in Figs. 7 and 8 in order to highlight the
relationship between lithology, species distribu-
tion and population abundances at Ain Settara.
In the latest Maastrichtian CF1 Zone, closely
spaced samples emphasize the relationship be-
tween truncated species ranges in sample 4 and
the disconformity at the top of the bioturbated
interval (Fig. 7). These features suggest that the
simultaneous species disappearances (7% of spe-
cies) are due to a hiatus and/or condensed sedi-
mentation related to the disconformity, rather
than biological factors. Since the six species are
relatively rare, deeper-dwelling tropical^subtropi-
cal taxa, their disappearance has little e¡ect on
the overall foraminiferal population.
Burrows in¢lled with dark clay and truncated
at the top just below the K^T boundary (red
layer) are also evidence of hiatus and/or non-de-
position. The species that simultaneously disap-
pear at the K^T boundary may here be also, in
part, artifacts of the incomplete or condensed
record making di⁄cult to identify the authentic
biotic response to the K^T event.
Hiatuses at or near the K^T boundary have
been widely documented and related to sea-level
changes. On the basis of a sequence stratigraphic
model, together with faunal evidence, several
studies have documented a sea-level lowstand pre-
ceding the K^T transition, followed by a general
rise in sea-level in many sections of continental
shelf to basin margin settings (e.g. Donovan et
al., 1988; Baum and Vail, 1988; Mancini et al.,
1989; Keller, 1989a; Savrda, 1991, 1993; Habib et
al., 1992; Moshkovitz and Habib, 1993; Stinnes-
beck et al., 1993; Keller et al., 1993a, 1994; Pardo
et al., 1996; Keller and Stinnesbeck, 1996). Some
of these authors have interpreted clastic deposits
at or near the K^T boundary as sea-level low-
stand deposits, and others interpreted the same
 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319 309

Fig. 7. Detail of the Ain Settara section, showing species distribution against the lithological changes across the K^T boundary.
The enlarged interval highlights the truncation in species distribution in correspondence with the unconformable surface at the
top of the uppermost Maastrichtian CF1 Zone. These sedimentological and biostratigraphic features are probably the result of a
hiatus and/or condensation. Another hiatus/condensation probably occurred just below the K^T boundary as suggested by bio-
turbation truncated at the top. Note also the simultaneous appearance of six new Tertiary species (which usually appear in se-
quence) in the Lower Danian (sample 8) associated with the deposition of a ‘tempestite’ level. These characteristics are also prob-
ably related to a hiatus. Hiatuses at the P0^P1a boundary are widely documented, and are generally related to a sea-level
lowstand in the Early Danian (e.g. Keller and Stinnesbeck, 1996; Pardo et al., 1996).

deposits as impact-generated megatsunami depos-
its (e.g. Bourgeois et al., 1988; Hildebrand and
Boynton, 1990; Alvarez et al., 1992; Smit et al.,
1992, 1994a,b; Bralower et al., 1998).
There is strong evidence for a sea-level low-
stand below the base of the Plummerita hantkeni-
noides Zone at about 65.5 Ma, coincident with the
late Maastrichtian maximum cooling (Keller and
Stinnesbeck, 1996; Pardo et al., 1996; Li and Kel-
ler, 1998b). After this cool event there is rapid
warming between 200 and 400 kyr before the
K^T boundary, followed by a rapid cooling dur-
ing the last 100 kyr of the Maastrichtian (Li and
Keller, 1998b). It is possible that the disconform-
ity 2 cm below the boundary at Ain Settara is
related to the cooling of the last 100 kyr of the
Maastrichtian, associated increased current activ-
ity and possibly a sea-level lowstand. At Elles II,
a cross-bedded foraminiferal packstone also
marks a disconformity just below the K^T bound-
ary red layer and clay (Keller et al., 2002).
A third disconformity and hiatus at Ain Settara
is present at the P0^P1a boundary, associated
with deposition of the silty layer (sample 8) above
the boundary clay (Figs. 7 and 8). The silty layer
appears to consist largely of reworked sediments,
as indicated by an increase in the abundance of
Maastrichtian species (Fig. 8), and may represent
a storm event (‘tempestite’). Maastrichtian spe-
cies, however, are also present above and below
the silty layer. This suggests either that reworking
is ubiquitous above the K^T boundary, or that
the Maastrichtian species are actually survivors,
but that most of the specimens in the ‘tempestite’

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310 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319

Fig. 8. Enlarged interval across the K^T boundary at Ain Settara showing the relative abundance changes of selected planktonic
foraminiferal species plotted against the lithology. It is clearly visible that the increase of uppermost Cretaceous species in lower-
most Danian within the ‘tempestite’ (sample 8) is related to reworking by the deposition of the ‘tempestite’ level. On the other
hand, the specimens in the non-bioturbated boundary clay and in the shales above may be survivors. This is suggested by their
good preservation, consistent presence at Ain Settara and by the fact that most of these species have been recognized as survivors
in other K^T transitions.

are transported and reworked. There is some sup-
port for a hiatus and the reworking interpretation
in that six new Tertiary species simultaneously
appear within the silty ‘tempestite’ layer, includ-
ing Parvularugoglobigerina eugubina and P. lon-
giapertura. Since these species evolve sequentially
(Fig. 4), such simultaneous ¢rst occurrences gen-
erally indicate a hiatus and/or condensed sedimen-
tation, and at Ain Settara they are also associated
with the disappearance of four globotruncanids.
The relative abundances of the new Tertiary spe-
cies in the ‘tempestite’ layer and in the overlying
5 cm are very low. This may either indicate the
early part of their evolutionary range where most
species populations are small, or winnowing and
poor preservation (dissolution and mechanical
breakage). Since the latter is not observed and
the Danian species are characteristic of the lower
part of the Pla Zone, the hiatus at the Ain Settara
section is probably very short, corresponding to
the lowermost part of Pla(l) and part of P0.
Hiatuses at the P0^P1a boundary are widely
documented in numerous expanded shallow-water
K^T sections (Keller and Stinnesbeck, 1996; Par-
do et al., 1996). As previously pointed out, the
database derived from detailed multidisciplinary
studies aimed at determining the paleodepths
and relative sea-level changes, indicates a gener-
ally rising sea-level after a main lowstand at the
end of the Maastrichtian. The sea-level rise was
interrupted by at least two short-term lowstands
in the earliest Danian, one of them at or near the
P0^Pla boundary. The lowstands are generally
marked by hiatuses involving part or entire bio-
zones and/or by condensed sedimentation related
to the transgression which follows the lowstand.
Thus, the transition between the zones P0 and
P1a is generally characterized by a hiatus, and
marks the ¢rst short-term Paleocene sea-level low-
stand (Keller and Stinnesbeck, 1996; Pardo et al.,
1996). The deposition of the ‘tempestite’ layer at
the Ain Settara section appears to be related to
this lowstand.
6. Extended and selective K^T boundary turnover
6.1. K^T boundary extinctions
Species distribution and relative abundance

PALAEO 2759 1-5-02

 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319
changes in species populations reveal a progres-
sive and selective planktonic foraminiferal turn-
over at Ain Settara (Figs. 5 and 6). 20% of the
Maastrichtian species disappear between 35 and
5 cm below the K^T boundary. Thirteen species,
or 21%, disappear at or below the boundary
and their combined relative abundance is less
than 10% of the total population (Fig. 6). All
41% of the species which disappeared at or below
the K^T boundary are rare to very rare and pri-
marily ecologically specialized keeled deeper-
dwelling tropical^subtropical forms (Globotrun-
cana, Globotruncanita, Gublerina, Planoglobulina,
Rosita (Contusotruncana), Racemiguembelina).
Their combined relative abundance varies be-
tween 10% and 15% of the total population at
the end of the Maastrichtian. The K^T crisis
thus appears more catastrophic when viewed in
tropical^subtropical assemblages and based on
analysis of larger species ( s 200 Wm) which pref-
erentially include the more specialized forms,
though, in fact, the K^T mass extinction actually
involved a numerically small part of the foramini-
feral population.
The extinction pattern at Ain Settara K^T
boundary is very similar to the extinction pattern
observed in other sections from north^central Tu-
nisia (El Kef and Elles; Keller, 1988a; Keller et
al., 1995, 2002; Molina et al., 1998; Karoui-Ya-
koub et al., 2002) and other low latitude K^T
boundary successions, such as those from Italy,
Spain, Mexico, and Texas (Luciani, 1997; Canu-
do et al., 1991; Keller, 1989a,b; Keller and Lo-
pez-Oliva, 1994; Lopez-Oliva et al., 1996). Varia-
tions in the extinction pattern are primarily due to
di¡erent taxonomic concepts (lumpers versus
splitters, see Karoui-Yakoub et al., 2002), di¡er-
ential preservation, analytical methods used (size
fraction analyzed, sample intervals) and local en-
vironmental conditions.
Closely comparable planktonic foraminiferal
turnovers in di¡erent sections increase the reliabil-
ity of data. In the above mentioned studies, spe-
cies disappearing at or below the K^T boundary
mainly consist of large, complex, specialized trop-
ical^subtropical taxa, with a combined relative
abundance of less than 20%. The data from Ain
Settara con¢rm this selective extinction pattern
PALAEO 2759 1-5-02
311
and indicate that the K^T crisis was more severe
in low latitudes where it selectively eliminated the
highly specialized tropical to subtropical species
group. The numerically abundant survivors and
the pre-K^T disappearance of some species sug-
gest that environmental factors other than a me-
teorite impact also contributed to the K^T crisis.
6.2. Reworked or survivors in Danian sediments?
Above the K^T boundary, in the thin non-bio-
turbated boundary clay (sample 7), 33 (or 56%) of
the Maastrichtian species are present and 7% of
these species disappear at the P0^P1a boundary;
34% gradually disappear within the P1a(1) sub-
zone, and 16% are present to the top of the sec-
tion analyzed (Fig. 5). Are any or all of these
species reworked? Or are they survivors? The ex-
tent of the mass extinction largely depends on the
interpretation of these species, whether reworked
or survivors, in Lower Danian sediments.
The Cretaceous species present in the Lower
Danian are mainly small cosmopolitan surface-
water dwellers such as Heterohelix, Pseudoguem-
belina, Globigerinelloides, Hedbergella, and Guem-
belitria (Plate I). All of these taxa are of small
morphology with little or no surface ornamenta-
tion, similar to the new evolving Early Danian
fauna, and they are relatively evenly distributed
in Lower Danian sediments of Ain Settara. There
are also isolated occurrences and a few poorly
preserved specimens that are considered reworked
(e.g. Racemiguembelina powelli in sample 7, Glo-
botruncana aegyptiaca and Rugoglobigerina ma-
crocephala in sample 32 and Plummerita hantke-
ninoides in sample 20, Fig. 5).
The Cretaceous species observed in the P0 and
P1a zones at Ain Settara are largely the same as
in the El Kef and Elles sections (Keller et al.,
1995, 2002; Karoui-Yakoub et al., 2002). The sur-
face dwellers Rugoglobigerina rugosa and Rugo-
globigerina hexacamerata and the keeled deeper
dwellers Globotruncana arca and Globotruncanita
stuarti have also frequently been observed in the
lowermost Danian, but these are not generally
viewed as survivors (MacLeod and Keller, 1994;
Keller et al., 1995).
Although reworking versus survivorship argu-
312 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319

PALAEO 2759 1-5-02

 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319 313

ments are still a matter of intense debate, it is now
currently accepted that at least a certain number
of species may have survived the K^T mass ex-
tinction. Criteria for evaluating species survivor-
ship include dwar¢ng, continuity in stratigraphic
range, good preservation, wide geographic distri-
bution and Cretaceous carbon isotope signals in
foraminiferal shells. In addition, the decline of
potential survivors coincides with the prolifera-
tion of new Danian species and suggests ecologi-
cal competition, rather than mechanical rework-
ing (e.g. Keller, 1988, 1989; Barrera and Keller,
1990; MacLeod and Keller, 1994; Keller et al.,
1995). Biogeographic analysis of Cretaceous spe-
cies in Danian sediments reveals higher extinction
rates of the survivor fauna at the P0^P1a transi-
tion in low latitudes than in higher latitudes where
species tend to persist into the overlying P1b and
P1c zones. This complex paleobiogeographic pat-
tern is further support of survivorship (MacLeod
and Keller, 1994).
All the above mentioned criteria are encoun-
tered for the Cretaceous species found in the low-
ermost Danian at Ain Settara. Moreover, the
presence in this section of the resedimented ‘tem-
pestite’ at the P0^P1 boundary provides the op-
portunity to separate the e¡ects of reworking
from normal deposition. For example, the in-
creased percentage of Cretaceous species within
the ‘tempestite’ seems to be related to reworking,
whereas the specimens in the non-bioturbated
boundary clay and in the shales above may be
survivors. This is suggested by the fact that
most of these species have been recognized as
survivors in other K^T transitions (see also Arz
et al., 1999), but also by their good preservation
and consistent presence at Ain Settara.
6.3. Species survivorship patterns
In higher latitudes, the K^T crisis had little
e¡ect on species survival rates, but signi¢cantly
reduced the relative abundances of species in the
earliest Danian (Keller, 1993; Keller et al., 1993c;
Pardo et al., 1998). A major change in the dom-
inant population also coincides with the K^T
boundary at Ain Settara as well as at El Kef
and Elles. Species that dominated during the
Late Maastrichtian generally survived into the
Early Danian, though their relative abundance
drastically declined after the K^T boundary (e.g.
Heterohelix globulosa, Heterohelix navarroensis,
Heterohelix dentata, Fig. 6). The decline of the
biserial group was accompanied by the rising
dominance of the triserial group. Guembelitrids,
in particular Gu«mbelitria irregularis, became dom-
inant in the planktonic foraminiferal assemblages
above the K^T boundary. The same replacement
of biserial by triserial taxa is observed in all Tu-
nisian sections (Keller et al., 2002) and is repre-

Plate I. Scale bar = 50 Wm. Some of the simple ecological generalist species which are possible survivors of the K^T mass extinc-
tion from the Ain Settara section, Tunisia.

1 Heterohelix navarroensis Loeblich. 90^95 cm above the K^T boundary.

2 Heterohelix dentata Stenestad. 20^25 cm above the K^T boundary.
3^6 Globigerinelloides aspera (Bolli), 3: 90^95 cm above the boundary; 4: 3 cm below the boundary; 5: 35^40 cm
above the K^T boundary; 6: 50^55 cm above the K^T boundary.
7^10 Heterohelix globulosa (Ehrenberg). 7 and 9: 3 cm below the K^T boundary; 8: 7 cm above the K^T boundary;
10: 10 cm below the K^T boundary.
11^13 Gu«mbelitria trifolia (Morozova). 11: 2 cm above the K^T boundary, from the boundary clay; 12 and 13: just
below the K^T boundary.
14 and 15 Gu«mbelitria irregularis (Morozova). 12^14 cm above the K^T boundary.
16 and 17 Gu«mbelitria cretacea (Cushman). 16: just below the K^T boundary; 17: 2 cm above the K^T boundary, from the
boundary clay.
18^21 Hedbergella monmouthensis (Olsson). 18: 4 cm below the boundary clay; 19: just below the K^T boundary;
20 and 21: 2 cm above the K^T boundary, from the boundary clay.
22 and 23 Hedbergella holmdelensis Olsson. 22: 2 cm above the K^T boundary, from the boundary clay; 23: 4 cm below the
boundary clay.

PALAEO 2759 1-5-02

314 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319
sentative of the faunal turnover in all low and
middle latitude sections. The decline of common
Cretaceous survivors in the Early Danian is grad-
ual and associated with the increased abundance
of evolving early Tertiary species. This suggests
increasing competition with Danian species better
adapted to the changing environment (Canudo et
al., 1991; Keller, 1989a,b; Keller et al., 1993c,
1995).
The marked increase in guembelitrids above the
boundary to about 70% at Ain Settara is charac-
teristic for most K^T boundary successions. These
very small triserial morphotypes are present in the
latest Maastrichtian, but rare in deep-sea and out-
er continental shelf regions, and more abundant
in shallow epicontinental seas (Leckie, 1987; Kel-
ler, 1989; Liu and Olsson, 1992; Schmitz et al.,
1992; Keller et al., 1993c, 1998). Numerous data
from ancient successions as well as analogies with
the extant triserial morphotypes (Gallitellia vivans)
indicate that guembelitrids were tolerant of unsta-
ble environments, and that they adapted to low-
salinity and low-oxygen conditions and variations
in temperature and nutrients (e.g. Wang et al,
1985; Kroon and Nederbragt, 1990; D’Hont
and Keller, 1991; Keller et al., 1998a,b). Evidence
of an expanded oxygen minimum zone which in-
vaded the shallow continental shelf has been
documented at El Kef on the basis of a detailed
integrated study (Peypouquet et al., 1986; Donce
et al., 1985; Keller, 1988b, 1992; Keller et al.,
1995).
It is possible that the marked increase in guem-
belitrids at Ain Settara corresponds to the expan-
sion of the oxygen minimum zone which may
have invaded the Kalaat-Senan shelf with the ris-
ing sea-level across the boundary. The decrease in
abundance of these forms coincides with a litho-
logical change from dark gray clayey shale to gray
¢ssile shale. This change also coincides with the
consistent increase in abundance of the earliest
Danian species thus suggesting the progressive en-
vironmental recovery after stressed conditions.
7. Summary and conclusions
The Ain Settara succession, as well as those at
PALAEO 2759 1-5-02
El Kef and Elles, were deposited during the K^T
transition in a region of the North African plat-
form which was strongly in£uenced by sea-level
£uctuations and variations in the oxygen mini-
mum zone. These e¡ects may have been ampli¢ed
in this relatively shallow shelf region and com-
bined with the superimposed global changes in
thermal and chemical strati¢cation, as well as
the e¡ects of a meteorite impact, were likely re-
sponsible for the extinction of the tropical^sub-
tropical faunas at the K^T boundary in both shal-
low and deep-sea environments. However, major
climatic changes began well before and continued
well after the K^T boundary. The survivorship of
the cosmopolitan species appears to be directly
linked to their greater tolerance of these environ-
mental £uctuations. Their decline and eventual
demise in the Early Danian seems to be a conse-
quence of direct competition with the evolving
and better adapted Early Danian fauna.
The following conclusions can be reached based
on the Ain Settara section:
(1) The thin red layer that marks the K^T
boundary and contains enhanced Ir and spinel
concentrations is present at Ain Settara. Never-
theless, the sediment record across this section is
not continuous and three disconformities have
been identi¢ed based on lithological changes, bio-
turbation, and the simultaneous appearance and
disappearances of many species which are known
to evolve or disappear sequentially in continuous
sections. The three disconformities occur a few cm
below the K^T boundary, just below the red layer
and at the top of the 2 cm thick boundary clay
and overlying silty ‘tempestite’ layer which marks
the P0^Pla boundary. However, there was rela-
tively little active erosion at these disconformities
as suggested by the presence of a 2 cm thick
boundary clay and Zone Pla (l).
(2) The species recognized, the relative popula-
tion abundance and the extinction pattern appear
very similar, and comparable with those of the El
Kef I and II (Keller et al., 1996) and Elles I and II
sections (Keller et al., 1995, 2002). At Ain Settara
21% of species disappear between 25 and 5 cm
below the K^T boundary; these species (Plum-
merita reicheli, Globotruncanita angulata, Hetero-
helix planata, Gublerina robusta, G. cuvilleri, G.
 V. Luciani / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 299^319
insignis, G. duwi) also gradually disappear below
the boundary at El Kef I, as do the ¢rst four
species at Elles II (Keller et al., 1995; Keller et
al., 2002). The simultaneous disappearance of six
other species below the boundary at Ain Settara is
presumably related to a short hiatus and/or con-
densed sedimentation as marked by a weak un-
conformity. The species becoming extinct at this
level are mainly deeper dwellers, and belong to
the tropical^subtropical group. Their disappear-
ance could be related to the cooling documented
at the very latest Maastrichtian (100 kyr; Li and
Keller, 1988c).
The main extinction event involved 22% of the
total species, which constitute only 10% of the
total population, in terms of relative abundance.
The extinction pattern is gradual and selective:
most of the tropical complex and deeper-dwelling
species disappear below or at the boundary
whereas the majority of the small cosmopolitan
surface dwellers are still present in the Lower
Danian. These forms largely correspond to those
found in the P0 and P1 zones at El Kef and Elles
(Keller et al., 1996, 2002).
A hiatus is implied by the deposition of the
‘tempestite’ level and is marked by the simultane-
ous appearance of six Tertiary species, included
Parvularugoglobigerina eugubina. Hiatuses at the
P0^P1a boundary are widely documented in shal-
low-water sections, and are related to a sea-level
lowstand which re£ects a negative in£ection in the
general rising trend in the earliest Danian (e.g.
Pardo et al., 1996; Keller and Stinnesbeck, 1996).
The opportunist low-oxygen tolerant guembeli-
trids became the dominant population in the ear-
liest Danian, substituting the latest Maastrichtian
dominant groups. The relative abundances of the
emerging Tertiary species are low at ¢rst; their
percentages increase when guembelitrids decline,
suggesting the progressive recovery after stressful
environmental conditions.
In conclusion, the extended pattern of extinc-
tion and changes in dominant populations at Ain
Settara show close analogies with the planktonic
foraminiferal turnover of other sections of cen-
tral^northern Tunisia (Keller et al., 1995, 2002).
This indicates that this area was a¡ected by com-
parable environmental changes at the K^T tran-
PALAEO 2759 1-5-02
315
sition. These changes appear peculiar and largely
related to the shallow continental shelf depth
(Keller et al., 1995). Sea-level £uctuations, expan-
sion of the oxygen minimum zone and related
water-mass changes may have ampli¢ed their ef-
fects in this area with respect to other shallow-
water K^T successions, thus explaining the
marked decline of survivors above the boundary
unrecorded in other K^T transitions. Albeit it is
possible that the disappearance of the Maas-
trichtian species at the boundary clay and above
it is related to the environmental changes resulting
from a bolide impact (Arenillas et al., 2000; Du-
puis et al., in press), it appears more probable that
the e¡ects of a bolide impact may have superim-
posed those related to long-term environmental
changes such as temperature variations, sea-level
£uctuations, major volcanism and possibly multi-
event scenario of impacts (Stinnesbeck et al., in
press) with associated perturbation in the water
column.
Acknowledgements
I am grateful to G. Keller for initially proposing
and encouraging this study, and for providing the
samples and for helpful comments and suggestions
of an early draft of the manuscript. Samples were
collected during the May 1998 Workshop on K^T
transitions and I am grateful to T. Adatte and W.
Stinnesbeck for sampling the section and supply-
ing the lithological description, and to A. Pardo
for kindly providing outcrop photos. I sincerely
thank H.P. Luterbacher and X. Orue-Etxebarria
for the careful review, and the editor J. Remane
for further improving of the final manuscript. This
work is financially supported by the Ministry of
the University, Scientific Research and Technol-
ogy (ex 60% Program, resp. V. Luciani).
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