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Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231

Late Maastrichtian to early Danian calcareous nannofossils at

Elles (Northwest Tunisia). A tale of one million years across
the K^T boundary
Silvia Gardin 
ESA-CNRS 7073, De¤partement de Ge¤ologie Se¤dimentaire, Case 117, Universite¤ Pierre et Marie Curie, 4 Place Jussieu,
75252 Paris Cedex 05, France

Received 29 October 1999; accepted 9 August 2001


A detailed study of calcareous nannofossils of the Elles section (Tunisia) was extended over an interval of about
1 Myr including the K^T boundary. The late Maastrichtian assemblages are rich and well preserved. Pulses of surface
water cooling, probably related to episodic upwelling are revealed by the increase in abundance of cool water taxa
(mainly Kamptnerius magnificus, Nephrolithus frequens, Gartnerago spp.) in certain late Maastrichtian samples,
paralleled by a decrease of the warm water species Watznaueria barnesae. A sensible decrease in abundance of
Arkhangelskiella cymbiformis and its return to smaller specimens was also recorded at Elles, probably as the result of
surface water productivity changes in the latest Maastrichtian. Some Maastrichtian samples at Elles contain
Neobiscutum romeinii and Chiasmolithus? sp. These taxa, formerly thought to be Tertiary coccoliths, are here
considered as truly Maastrichtian occurrences. As at El Kef, the Danian of Elles is characterised by successive blooms
of surviving and newly evolved species. Though the abundance of calcareous nannofossils rapidly reached the same
level as in the Maastrichtian, the lower diversity still points to a mesotrophic, poorly stratified marine environment.
The reworking versus survivorship of Cretaceous species found in Danian samples remains an unresolved question,
but a hypothesis of survivorship is also strengthened by the presence of several coccospheres of W. barnesae and
Bidiscus rotatorius well above the boundary. The effect of the marine environmental change across an expanded K^T
boundary section as Elles appears complex if seen through calcareous nannofossils. From a geological point of view,
the K^T boundary floral turnover at Elles seems dramatically synchronous with the signatures of an extraterrestrial
impact, superimposed on long-term, background environmental and climatic changes. However, a precise succession
of events based on appearance/disappearance of nannofossil species across the Cretaceous/Paleocene transition is far
from being unanambiguous. ß 2002 Elsevier Science B.V. All rights reserved.

Keywords: calcareous nannofossils; K^T boundary; Tunisia

1. Introduction

Over the past few decades, faunal and £oral

* Fax: +33 (1) 442 73 831. analysis across the Cretaceous^Tertiary (K^T)
E-mail address: (S. Gardin). boundary have suggested two di¡erent basic pat-

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 3 9 7 - 2

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terns: gradual, stepwise extinction at, in, or below exterminated by the impact of a large extraterres-
the boundary versus one catastrophic mass extinc- trial body highly fascinates the imagination. The
tion. The greatest part of marine microfossil stud- geological and geochemical signatures of a large
ies have concentrated on the few hundred cm extraterrestrial body impact in the Caribbean re-
above and below the Ir anomaly, in an e¡ort to gion are now irrefutable, but the exact nature,
evaluate the few thousand years before and after rate and tempo of the marine environment reac-
the well-documented impact event. Such a short tions are still di⁄cult to establish. The following
time interval did, however, not yield the necessary major controversies still exist among palaeontol-
environmental information needed to evaluate ogists :
possible long-term trends and palaeoenvironmen- (a) Was the marine biotic diversity on the decline
tal changes preceding the K^T event. Thus, sev- before the K^T boundary event?
eral high-resolution researches on planktonic for- (b) Were the e¡ects of the K^T boundary event
aminifers (Li et al., 1999; Li and Keller, 1998a, globally catastrophic and/or geographically con-
1998b, 1998c) have recently documented the last trolled?
million years of the Maastrichtian marine envi- (c) Is the meteoric impact the only and unique
ronmental changes at di¡erent low-latitudes sites, cause of the biotic crisis?
strengthening the multi-causal scenario for this
best studied mass extinction. 1.2. Calcareous nannofossils and the
The aim of the present work is to document K^T boundary
through a high-resolution study, the stratigraphic
occurrence and the palaeoecology of late Maas- Calcareous nannoplankton provides much infor-
trichtian to early Danian calcareous nannofossils mation about the changing environment across
at Elles. The time slice investigated across the K^ the K^T boundary. The calcareous shields con-
T boundary, about 1.2 Myr, is longer than that tribute to the bulk of marine sediments, providing
usually reported in published literature, being not a record of climatic and productivity changes
just the ‘1 m below and above’ the boundary through the N18 O and N13 C ratio retained in
which is known in detail from di¡erent latitudes. them. Calcareous nannoplankton has developed
Thus, a more detailed and wider dataset of calca- strategies of adaptation to extremely low nutrient
reous nannoplankton palaeoecology during the availability (Kilham and Soltau Kilham, 1980)
last 650 kyr of the Maastrichtian can provide fur- and the maximum diversity observed today is in
ther information on the marine palaeoenviron- the oligotrophic oceanic gyres, where the vertical
ments preceding the K^T boundary event. Un- distribution of species depicts the strati¢cation of
fortunately, besides a limited number of speci¢c the photic zone. It is therefore reasonable to infer
papers on Maastrichtian calcareous nannofossil that past diversity changes in the calcareous nan-
stratigraphy (Burnett et al., 1992) and biogeogra- noplankton community re£ect changes in the
phy (Watkins, 1992; Henriksson and Malmgren, strati¢cation of the photic zone. Also, due to their
1997, among others), no detailed calcareous nan- continuous geological record and rapid species
nofossil quantitative study of the whole Maas- evolution, calcareous nannoplankton have an ex-
trichtian has been published so far. cellent biostratigraphic resolution potential.
Nannoplankton specialists have placed strong
1.1. The Cretaceous^Tertiary boundary emphasis on the global synchronous extinction
of many representatives of this group at the K^
The K^T boundary is the best studied mass T boundary and according to them nothing in the
extinction event in the geological record. This is record predicts the mass extinction event. The cal-
due in part to the attractive discussion that has careous nannoplankton extinction pattern and
gone on since the 1980 suggestion of the bolide magnitude at both high and low-latitude sites
impact scenario (Alvarez et al., 1980). The idea seems very similar (Pospichal, 1996a) and it is
that dinosaurs and many marine organisms were considered as a geologically instantaneous event.

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No latitudinal extinction pattern has been ob-

served among calcareous nannoplankton, the dif-
ferences are in the composition of the assemblages
due to late Maastrichtian and early Danian pro-
vincialism (Pospichal, 1996a, 1996b). There is an
exhaustive literature dealing with detailed calcare-
ous nannofossil studies across the K^T boundary
(Bramlette, 1965; Perch-Nielsen, 1979a,d; Wors-
ley, 1974; Percival and Fisher, 1977; Romein,
1979; Thierstein and Okada, 1979; Thierstein,
1981; Jiang and Gartner, 1986; Monechi, 1977,
1979; Perch-Nielsen, 1981a, 1981b; Romein and
Smit, 1981; Perch-Nielsen et al., 1982; Seyve,
1990; Gorostidi and Lamolda, 1991; Lamolda
and Gorostidi, 1992; Eshet et al., 1992; Pospi-
chal, 1994, 1995; Gartner, 1996; Henriksson,
1996; Gardin and Monechi, 1998).

2. The Elles section

2.1. Previous work

During the ¢eld trip of the ‘K^T boundary

Workshop’ held in Tunis on May 1998, two other
K^T sections were sampled beside the well-known
stratotype of El Kef: The Elles and the A|«n Set-
tara sections. The planktonic and benthic fora-
minifera content of the Elles section had been
previously investigated by Said (1978). Though a Fig. 1. Paleogeographic setting of northern Tunisia and loca-
tion map of the Elles section. Lines indicate palaeo-water-
number of nannofossil studies on the K^T strato- depth.
type at El Kef have been published (Perch-Niel-
sen, 1979b,c, Perch-Nielsen, 1981a, 1981b; Pospi-
chal, 1994; Verbeek, 1977; Gardin and Monechi,
1998), no calcareous nannofossil analysis has been and comprises a well-exposed marly succession
carried out on the Elles section. of late Cretaceous^Eocene age. The palaeogeo-
graphic setting of the Elles section is similar to
2.2. Location and lithology of the Elles section that of El Kef, both localities being situated on
the continental shelf. Elles is located in a slightly
Tunisian sections feature on a high sedimenta- more proximal position than El Kef.
tion rate and an apparently continuous sedimen- The studied interval starts at about 326 m be-
tary record. They are thus well suited to study low the boundary up to the lowest 13 m of the
and to understand the K^T boundary event(s) Danian. The upper Maastrichtian sediments con-
as well as the whole late Cretaceous interval. sist of grey marls, with some siltstone intercala-
The Elles section is located in the Tunisian Cen- tions. The K^T boundary transition is character-
tral Atlas (Fig. 1) 35 km southeast of El Kef. The ised by 20 cm of grey sandy^silty marls, followed
El Haria Formation stretches about 40 km long by a 5-cm, bioturbated, cross-bedded yellow sand-
and 10 km wide within the Elles western syncline stone, which in turn ends with 0.5 cm of green

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clay. The boundary is well marked by a 0.5-cm 3. Results

red layer, rich in Fe-oxides and containing Ir-
and Ni-rich spinels. The lowest Danian sediments 3.1. Biostratigraphy
start with grey clay, 38 cm thick, and are be-
coming more marly to the top of the studied in- The Maastrichtian portion of the Elles section
terval. studied here belongs to the Nephrolithus frequens
A sedimentation rate of 4 cm/1000 yr (at least Zone (‘CC26’ of Sissingh, 1977). This taxon is
for the Maastrichtian) was calculated based on quite rare due to its high-latitude preference. C.
biostratigraphic correlation with DSDP Site 525 kamptneri, Litraphidites quadratus were also ob-
and the palaeomagnetic data from that site (Kel- served from the ¢rst sample, at 326 m. The latter,
ler, written communication, 1999; Li and Keller, which is the zonal marker of Zone NC 22 (Roth,
l998c). 1978) and of the L. quadratus Zone of Thierstein
(1976), is commonly present in all the studied
2.3. Sampling and methodology samples. In terms of subzones (CC 26a and CC
26b; Perch-Nielsen 1985a) the studied section be-
For the Maastrichtian portion, samples were longs to the Micula prinsii nannofossil subzone
taken at about every metre from 326 m to 321 (CC26b) from 323 m (the ¢rst sample where M.
m; every 50 cm from 321 m to 33 m and every prinsii was observed). Micula murus and M. prinsii
10 cm from 3 m to 0.1 m. The last 10 cm were are always very rare in the lower part of the
sampled every cm. The boundary level, whose studied section, probably due to poor preserva-
thickness is about 20 cm, was sampled at 1^5 tion. They are more common from sample 18 up-
cm. The Danian sediments were sampled every wards.
50 cm untill 13 m. The position of the M. prinsii subzone varies
Samples were processed as follows : 0.02 g of within C29R according to:
sediment were gently disaggregated in a mortar 1. The preservation of nannofossils (Flores et al.,
with 10 ml of distilled water. The obtained sus- 1990; Gorostidi and Lamolda, 1995; Henriksson,
pension was agitated several times in a tube and 1993), M. prinsii being sensible to dissolution.
left to settle down for 2 min. After this time smear 2. Palaeolatitude, M. prinsii being more abundant
slides were prepared using 0.2 ml of the solution. at low latitudes (Perch-Nielsen et al., 1982)
At least three traverses of the slides, correspond- 3. Sedimentation rate.
ing to 450 ¢elds of view, were examined in order Norris et al. (1998) reported a duration of the
to document species richness (Cretaceous+Ter- M. prinsii subzone of 1 Myr with its FO lying at
tiary species) encountered during the analysis the top of C30n, which is the oldest FO reported
of one slide (see Gardin and Monechi, 1998, by the literature for this taxon. It is not unlikely
for more detail). Relative species abundance that the FO of M. prinsii is diachronous and er-
was calculated as the percentage on 300 (total) rors may occur in using this taxon for sedimenta-
specimens randomly counted with a light micro- tion rate estimates and chronostratigraphic corre-
scope at a magni¢cation ofU1250. The total nan- lations.
nofossil abundance estimate was calculated In terms of planktonic foraminifer data (Abra-
through the ratio between the 300 nannofossils movich and Keller, this volume) the Maastricht-
counted and the number of ¢elds of view in which ian portion of the Elles section stretches from
this total was reached. Some Cretaceous species CF4 to CF1 Zones (Li and Keller, 1998).
such as Arkhangelskiella cymbiformis, Prediscos- The Danian portion of the Elles section extends
phaera grandis, Ei¡ellithus turrisei¡elii, Cribros- from NP1 to NP2 nannofossil zones of Martini
phaerella erhenbergii, exhibit large size ( s 14 (1971) and from the G. cretacea to the S. pseudo-
Wm) specimens. No distinct quantitative counts bulloides planktonic foraminiferal Zones (Keller et
were made of these specimens except for A. cym- al., this volume). The FO of C. primus, F. peta-
biformis. losa, C. tenuis is currently documented by the

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Fig. 2. Calcareous nannofossil biozonation of the K^T boundary interval and early Danian at low latitudes. Magnetochrons
from Cande and Kent (1995). The magnetic as well as the nannofossil bio-horizons are just depicted in relative order.

published literature. The FO of C. pelagicus is Figs. 3 and 4 show abundance and richness curves
reported by Monechi (1985) from the Northwest as well as the relative abundance of the most im-
Paci¢c Ocean (DSDP Leg 86) between the FO’s portant calcareous nannofossil genera and species
of C. primus and C. tenuis. In this work C. pela- which composed the assemblages. The richness
gicus features two morphotypes, one s 5 Wm in shows minor £uctuations along the studied sec-
size and one 6 5 Wm. Micrantholithus attenuatus tion; the abundance curve however, displays
was described by Bramlette and Sullivan (1961) wider £uctuations due to more or less well-pre-
and seems to be a synonym of M. inaequalis of served assemblages in the or more or less shaly
Martini (1961). Its FO has not been reported in lithology.
the literature so far. The lower Danian calcareous The late Maastrichtian assemblages seem to be
nannofossil biostratigraphical data of the Elles characterised by relative stasis with no clear ex-
section are summarised in Fig. 2. tinctions or originations except for the FO of M.
prinsii. Several species (Acuturris scotus, Tranoli-
3.2. Late Maastrichtian calcareous nannoplankton thus exiguus, Ceratholithoides kamptneri Biscutum
assemblages at Elles. Pre K^T environmental notaculum, Ceratolithoides aculeus, Microrhab-
features and palaeoecology dulus decoratus, Discolithus vagus) commonly
present in low-latitude sites during Campanian
The Elles section features an abundant, highly and Maastrichtian stages occur very rare and
diverse, generally well-preserved late Maastricht- spotty. Some others (Quadrum sissinghi, Reinhard-
ian assemblages. The stratigraphic distribution of tites levis, Tranolithus pahecelosus) have been en-
calcareous nannofossils is reported in Fig. A1. countered as one specimen only in a few samples.

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Fig. 3. Relative abundance of selected calcareous nannofossil genera and species in the upper Maastrichtian (326 to 33.00 m).
The dotted area in the abundance curve of A. cymbiformis corresponds to the large forms; the black one corresponds to small
forms (see text for explanation). The abundance curve ‘cool water taxa’ comprises the sum of the relative abundance of K. mag-
ni¢cus, Gartnerago spp., A. octoradiata and N. frequens

The stratigraphic range of Q. sissinghi, R. levis, T. Some Maastrichtian samples are characterised
pahecelosus, is well known from the literature, by a marked decrease of the warm water form W.
since their FAD (¢rst appearance datum) and barnesae and an increase of cool water taxa
LAD (last appearance datum) are used as zonal (Nephrolithus frequens, Gartnerago spp., Kampt-
markers; here they are present above what is usu- nerius magni¢cus and Ahmuellerella octoradiata).
ally considered their extinction level. The strati- This trend is clearly visible from sample 4 to 6 m
graphic range of the former taxa is less precisely (Fig. 3) These taxa are reported by Pospichal
documented, therefore it is di⁄cult to draw any (1996a,b) and Pospichal and Wise (1992) to be
reliable conclusion on these species (if they are common or more abundant in high latitudes as-
vanishing or even reworked). Species as A. scotus semblages, whilst they rarely occur at low-latitude
and B. notaculum may be palaeogeographically sites where W. barnesae is abundant.
controlled, being more abundant at high latitudes In the Maastrichtian high-latitude assemblages,
(Pospichal and Wise, 1992). which are generally less diverse than those at low
Holococcoliths such as Lucianorhabdus cayeuxi, latitudes (Pospichal, 1996), P. stoveri was ob-
Semihololithus priscus, S. bicornis, Russellia bukryi served to show an abundance acme prior to the
and Octolithus multiplus are generally considered boundary. This acme was not observed at other
as marginal or nearshore forms associated with low-latitude sites (Gardin and Monechi, 1998) ex-
shallow water deposits. Surprisingly, they are cept at Brazos ( = P. quadripunctata Jiang and
very rare in the inner neritic section of Elles. Gartner, 1986). At Elles, P. stoveri shows an in-
Large forms ( s 14 Wm) of Arkhangelskiella crease in its relative abundance 1 m below the
cymbiformis, Prediscosphaera cretacea, Ei¡ellithus boundary, with a maximum value of 10% at
turrisei¡eli and Cribrosphaerella ehrenbergii are about 20 cm below the K^T boundary.
very common. Peculiar is the trend observed for Neobiscutum romeinii was found in several sam-
A. cymbiformis. This species features a relative ples though in extremely low numbers. The rare
abundance of about 8^10% at the base of the occurrence in the Maastrichtian of this very small
studied interval, with a higher percentage of large species was previously recorded by several authors
forms (see above). Two levels (313.50 m and (Perch-Nielsen; Pospichal, personal communica-
39.0 m) exhibit a marked drop in its relative tions; Mai et al., 1997; Gardin and Monechi,
abundance, showing the lowest relative abun- 1998). This record is a further con¢rmation of
dance recorded for this species (0.3% and 0.9% the presence of this species below the boundary.
respectively). This fact is not associated with pres- N. romeinii re-entrys and blooms during the early
ervation or due to an artefact of preparation. Danian. A specimen of Chiasmolithus? sp.,
From 39 m upsection onwards the relative abun- (Fig. 5), occurs in some Maastrichtian samples
dance of A. cymbiformis increases slightly, £uctu- (3800, 3250, 3190, 380, 340, 320 cm res-
ating between 3 and 4%, but reaches never again pectively). Up to now no other author reported
the previous values of 10% or 6%. Moreover, the the presence of this genus, considered as Tertiary,
proportion of small versus large specimens begins in upper Maastrichtian sediments.
to increase and is higher in the last 140 cm. The The palaeoenvironmental signi¢cance of calca-
change in dominance from large to small speci- reous nannofossils is not yet completely under-
mens is in agreement with the ¢ndings of Girgis stood. Published palaeoproductivity studies
(1989) and Faris (1995) in the uppermost Maas- mainly concern Tertiary or Quaternary sediments.
trichtian of Egypt. A limited number of reports exist for the Meso-

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Fig. 4. Percent abundance of selected calcareous nannofossil genera and species in the uppermost Maastrichtian (33.00 to 0 m).
The dotted area in the abundance curve of A. cymbiformis corresponds to the large forms; the black one corresponds to small
forms (see text for explanation). The abundance curve ‘cool water taxa’ comprises the sum of the relative abundance of K. mag-
ni¢cus, Gartnerago spp., A. octoradiata and N. frequens.

zoic, but most of them do not propose any sig- boundary. The most remarkable feature is the to-
ni¢cant taxa enabling accurate palaeoproductivity tal absence of coccoliths in the Iridium layer,
reconstructions. Exceptions are Watkins (1989); while they are present below and above where
Premoli Silva et al. (1989); Eshet et al., (1992); they essentially include the most common species
Eshet and Almogi Labin (1996), who suggest to found in the Maastrichtian samples. The same
use nannofossil species abundances as palaeopro- pattern was described by Perch-Nielsen (1981)
ductivity proxies. According to their independent from El Kef.
statistical correlations, the latter authors broadly The abundance curve in the early Danian
distinguish two groups of calcareous nannofossils. strongly £uctuates at Elles (Fig. 6). Coccoliths
The one including Ei¡ellithus spp., Predisco- markedly increase within the 5^7-m interval,
sphaera spp. (not including P. stoveri), Litraphi- reaching the same values as found in the Maas-
dites, Vekshinella spp., was regarded as a low-pro- trichtian in the upper part of the section. Richness
ductivity group, whereas the one including shows drastic £uctuations in the ¢rst metre;
Zygodiscus, Biscutum, Discorhabdus and T. saxea above it slightly increases but never reaches the
was thought to represent high-productivity condi- high values of the Maastrichtian.
tions. If these correlations are correct, the studied The relative abundance of Cretaceous species
late Maastrichtian interval, which is characterised (A. cymbiformis, Ahmuellerella regularis, P. creta-
by abundant and diverse calcareous nannofossil cea, E. turrisei¡elii, W. barnesaeT) is rapidly di-
assemblages with common Ei¡ellithus turrisei¡elii,
Prediscosphaera cretacea and Litraphidites carnio-
lensis + L. quadratus, should be regarded as an
interval of relatively low primary productivity.

3.3. Calcareous nannoplankton palaeoecology and

turnover across the K^T boundary and during the
early Danian at Elles

There is no strong evidence for anomalous

Maastrichtian assemblages immediately prior to
the K^T boundary. Species distribution and rela-
tive abundance changes in species populations
show a major turnover only above the boundary,
not below. The calcareous nannofossil distribu-
tion pattern found at Elles is almost the same as
the one found at El Kef: above the clay layer
containing the Ir anomaly and devoid of cocco-
liths there is a drop in abundance of Cretaceous
species and an increase of the relative abundance
of the calcareous dinocyst Thoracosphaera opercu- Fig. 5. 1, 2, 3, 4:Chiasmolithus? sp. (U2500), crossed nicols.
lata. Species richness drops from about 70 species Same specimen rotated to show the structure of the central
in the latest Maastrichtian to 30 just above the crossbar. Sample 3800 cm, Elles section.

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Fig. 6. Relative abundance of selected calcareous nannofossil genera and species in the lowermost Danian.

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minishing throughout the studied interval with no 4. Discussion

clear LO except for Micula prinsii, Ahmuellerella
octoradiata, Microrhabdulus undosus, Predisco- The interpretation of the extinction pattern of
sphaera stoveri, Cribrocorona gallica and Creta- calcareous nannofossils across the K^T boundary
rhabdus crenulatus. Other Cretaceous species remains controversial. Though nannopalaeontolo-
seem to reappear at about 60 cm above the K^T gists are quite in agreement that no pre-boundary,
boundary (mainly Discorhabdus rotatorius, Pa- gradual or stepwise extinction happened during
laeopontosphaera sp., P. stoveri, Tranolithus exi- the late Maastrichtian, disagreement exists in con-
guus). K. magni¢cus, N. frequens, Gartnerago sidering the Cretaceous taxa (sensu Jiang and
spp. show the same relative abundance they had Gartner, 1992) found in the Danian samples ei-
in the Maastrichtian. ther as totally (Pospichal, 1994) or as only partly
Several Danian samples (+5 +40, +60 +150, reworked (Perch-Nielsen et al., 1982; Mai et al.,
+300 cm) are characterised by a peculiar increase 1997). Because of their very small size, calcareous
of dwarf specimens of W. barnesae, M. decussata nannofossils are easily reworked and this fact
and C. ehrenbergii with respect to those of normal hampers the interpretation of their extinction pat-
size. Several coccospheres of W. barnesae and D. tern across the K^T boundary. Up to date there is
rotatorius were found in Danian samples, together no way to prove whether a nannofossil specimen
with coccospheres of B. bigelowii and M. inversus. is reworked or in situ. The concept of a mass
Some of these coccospheres are present well above extinction largely depends on the interpretation
the K^T boundary (+7.5 m). of these species, whether reworked or survivors,
Calcareous nannofossil productivity in the low- in lower Danian sediments.
er Danian samples at Elles shows a pattern very Besides the quality of the fossil record, the oc-
similar to that observed at El Kef and at other currence or the non-occurrence of a calcareous
low-latitude sites (Gardin and Monechi, 1998); it nannofossil species prior to the boundary at Elles
is re£ected by the succession of blooms of surviv- (as well as at other sites) can be ambiguously in-
ing and newly evolved nannoplankton taxa terpreted because of three major factors:
(Fig. 5.). The ¢rst bloom was that of Thoraco- (a) Species which were rare during the late Maas-
sphaera, which started a few cm above the bound- trichtian and whose stratigraphic range is poorly
ary. This is in turn followed by successive blooms known (namely T. exiguus, M. decoratus, L.
of N. romeini, N. parvulum, F. petalosa, C. pelagi- cayeuxii, Ceratoilithoides arcuatus, Ceratolithoides
cus. Some taxa which exhibit blooms in other low- aculeus, Watznaueria quadriarcata, Discolithus va-
latitude sections, such as B. bigelowii, C. reinhard- gus), can produce the Signor^Lipps e¡ect (Signor
tii, M. inversus and Neocrepidolithus spp., are and Lipps, 1982), showing a false exinction. If we
present only with very low relative abundances order the stratigraphic distribution of calcareous
at Elles: less than 10% for B. bigelowii or even nannofossils in the Maastrichtian by LO’s of taxa
less than 2% for M. inversus. (Fig. A1), the resulting pattern shows a trend that
The ¢rst, more complex Danian species appear- can be interpreted as a disappearance of taxa be-
ing at Elles is C. primus ( 6 5 Wm), indicating the fore the boundary. However, several of these spe-
beginning of the long return to more stable envi- cies are considered to have become extinct before
ronmental conditions (Gardin and Monechi, (and then to be reworked), whereas others are
1998). It ¢rst occurs at 6.50 m during the ‘dwarf known to have survived into the Danian or are
Biscutum bloom’ (Gardin and Monechi, 1998) still living (B. bigelowii).
and within the P. pseudobulloides planktonic for- (b) Regional palaeocological factors control the
aminifer Zone, as observed at El Kef and at other maximal stratigraphical range of most species,
low-latitude sites (Gardin and Monechi, 1998), with the result that LO’s and FO’s have only local
about 200 kyr after the K^T boundary. During or regional signi¢cance.
this succession of blooms new taxa appeared/dis- (c) Any controversy about calcareous nanno-
appeared upsection (Fig. A2). plankton taxonomy results in contradictions con-

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222 S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231

cerning the stratigraphic distribution of calcare- re-entry from 60 cm above the K^T boundary
ous nannofossils. above an interval with no records (Fig. A2).
Low diversity, spotty occurrence and reappear-
4.1. Reworking versus surviving: some criteria ance of taxa might be a real and useful palaeon-
tological indication of lowered abundance in the
The interpretation of a calcareous nannofossil aftermath of an extremely perturbating event, and
species found above its presumed extinction level not just the e¡ect of reworking.
and above the K^T boundary, is not easy. Several Mai et al. (1997) found intact coccospheres of
species can be recorded even above their pre- Cretaceous taxa (K. magni¢cus, B. rotatorius, W.
sumed extinction level, especially in sediments barnesae and many others) in sediments of the
with a detrital component characterised by high Gullemberg section, 20 cm above the K^T bound-
sedimentation rates. Thus, some taxa considered ary, to coexist with the newly evolved species.
extinct as Aspidolithus parcus, Tranolithus phace- Based on the excellent state of preservation of
losus, Quadrum tri¢dum, Reinhardtites levis and these coccospheres, they suggested these species
Ei¡ellithus eximius were found, as one specimen to have survived at least until the evolution of
in some Maastrichtian and in some Danian sam- new species, such as C. primus. At Elles, several
ples at Elles. It is reasonable to consider them as coccospheres of W. barnesae and B. rotatorius
reworked though they have not been observed were found in the Danian samples together with
occurring together as a distinct reworked compo- coccospheres of survivors, some of them having
nent. The simple statement of reworking however, been found even 7.5 m above the boundary. Un-
can lead to a loss of precious palaeontological less the possibility that coccospheres can be en-
information : the occurrence of these taxa, though tirely reworked, this observation can be a further
extremely rare, could in fact be the result of support to the hypothesis of some species survi-
sporadic reappearance and preservation of the vorship, as invoked by Mai et al. (1997). We can
species itself. In the aftermath of extinction or not completely rule out in fact that, at least a part
of severe palaeoenvironmental changes, such as of them, is the result of in situ, episodic calci¢ca-
a sea-level drop, some population constituents tion.
can be drastically reduced in number and their Thus, the very complex stratigraphic pattern of
production can be much reduced. calcareous nannofossils above the boundary can
The Cretaceous taxa found in the very ¢rst level not be interpreted in terms of reworking only.
of the Danian consist of the most common species Survivorship of a few species with ecological com-
found in the Maastrichtian; they are often, but petition with opportunists and newly evolved taxa
not always, poorly preserved with an empty cen- should be considered as well. These species, which
tral area and they do not exhibit a clear LO (last are not thought to have given rise to the new
occurrence) along the section (Fig. A2). This pat- Danian taxa, may represent ‘short-time survivors
tern can be reasonably interpreted as reworking ; without progeny’ (Harries et al., 1995), which did
a great part of them is surely reworked. However, not take advantage of the conditions that allowed
some Cretaceous species do not exhibit a clear LO the new species to evolve and became extinct.
datum in the Danian just because they are rare They survived the crisis in very low number but
both below and above the boundary (namely Cor- did not adjust to the subsequent reorganisation of
ollithion completum, Gartnerago spp., K. magni¢- the earth’s physical and biotic system, probably
cus, N. frequens). Others instead (P. stoveri, M. because of di¡erent feeding strategies not met by
undosus, C. crenulatus, Cribrocorona gallica) exibit the changed availability of nutrients.
a more continuous record above the boundary Size is another criterion used to question re-
and a more evident LO along the section, so there working/versus survivorship of taxa. According
are no strong reasons to consider them as entirely to Keller and Von Salis-Perch-Nielsen (1997),
reworked. Moreover, D. rotatorius, Paleoponto- ‘measurements of size distribution of several
sphaera sp., P. stoveri, T. exiguus seem to clearly Maastrichtian species from samples just below

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S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231 223

and just above the boundary indicate no signi¢- and optical properties strongly point to Chiasmo-
cant di¡erences in the size of the specimens.’ At lithus/Sullivania characteristics. Possibly, this ge-
Elles several Danian samples are characterised by nus (or its ancestor?) was already present in the
a noticeable increase of smaller than normal latest Maastrichtian as N. romeinii. It has never
specimens of W. barnesae and M. decussata. How- been reported because its is extremely rare (the
ever, dwarf and small size of W. barnesae and M. specimens observed have been encountered at
decussata are present also in the Maastrichtian least after 300 ¢elds of view), and, probably, be-
samples, and an accurate ratio between small cause it was thought to be contaminant. Chiasmo-
and big specimens of these species has not been lithus ? sp. was observed also in the near El Kef
established in the present report. section (Gardin, unpublished data). Its rarity and
spotty occurrence, as well as the smallest size of
4.2. Are ‘new Tertiary species’ new? N. romeinii do not allow ¢xing reliable FO’s of
these taxa in the samples studied.
Beside the preservation e¡ects, the occurrence/ The ¢nding of Chiasmolithus ? sp. in Cretaceous
non-occurrence of calcareous nannofossils in the sediments rises new questions about the calcare-
fossil record can be related to their calci¢cation ous nannofossil’s mode of transition between the
potential (Janin et al., 1996). The absence of a Maastrichtian and the Danian stages, though its
nannofossil species in a sample in fact, does not diagnosis needs to be veri¢ed. It is not unlikely
straightforwardly mean that it has not biologi- that some genus and species, which were abun-
cally appeared or disappeared. Species may be dant in Danian sediments, were already present
present in much reduced numbers, thus beneath in the Cretaceous sediments but beneath the de-
the detection level of the fossil record, and they tection level of the species. They became more
can reappear afterwards, for sporadic occurrences abundant when the environmental conditions
as well as for blooms, as the result of environ- were more favourable for their development and
mental conditions more favourable to the life of evolution. This hypothesis corroborates that of
that species. In the special case of calcareous nan- Perch-Nielsen et al. (1982) and Mai (1999) and
nofossils, their absence or lowered diversity might leads to a more subtle interpretation of the calca-
be due to the reduced calci¢cation capability of reous nannofossil turnover across the K^T
some species, which is re£ected as well in the fos- boundary.
sil record (Gardin and Monechi, 1998). Calcare-
ous nannofossil populations are not homogene- 4.3. Was the late Maastrichtian cooling
ous, being formed by coccolithophorids and responsible for the K^T boundary biotic event?
other nannoliths incertae sedis; their peculiar
and still not clear biology can result in an ambig- The late Maastrichtian was characterised by
uous interpretation of their record in the geolog- latitudinal provincialism and the equatorward mi-
ical past. gration of N. frequens (Worsley and Martini,
This fact can explain the rare and spotty occur- 1970; Worsley, 1974) may re£ect these long-term
rence/preservation of N. romeinii in the upper changes. The presence of N. frequens at low lat-
Maastrichtian and its post K^T bloom as well itudes in fact, could be the result of following the
as the presence in several samples of Chiasmoli- cooler water masses as they expanded toward the
thus? sp., whose entry was thought to be exclu- equator and vice-versa. Decreasing diversity at
sively Tertiary (NP3 Zone of Martini, 1971). The high latitude and increasing latitudinal biogeo-
observed specimens of Chiasmolithus? sp. are well graphic provincialism among calcareous nanno-
preserved, they do not seem to be transported by fossils re£ected an increased ocean mixing as re-
water circulation or burrowing ; laboratory con- sponse to global cooling (Watkins et al. 1996).
tamination seems highly unlikely too. Unfortu- Slight cooling at the end of the Maastrichtian
nately its rarity prevented the diagnosis of this may be indicated by the nannofossil assemblages
taxon by SEM, but its central crossbar structure of other sections. At Bidart (SW France) Goros-

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224 S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231

tidi and Lamolda (1995) reported as reliable the collapse. According to the previous authors the
variations of the Micula spp. /W. barnesae ratio as global environmental and climatic deterioration
indicator of slight cooling in the upper part of the which started in the late Maastrichtian preferen-
Maastrichtian, in agreement with Wind (1979). In tially eliminated deeper and intermediate habitats
the late Maastrichtian interval of Site 527 (re- ¢rst, favouring survival in surface waters. This is
ported by Pospichal, 1996a) the cool water species well evident from their planktonic foraminifer re-
K. magni¢cus, N. frequens, Gartnerago spp. exibit sults, where the deep-dwelling K-strategists de-
a slight increase while the warm water species W. clined during the deposition of the A. mayaroensis
barnesae shows a sensible decrease. Oxygen iso- Zone. Such a behaviour is far less evident for
tope values corroborated this trend (Shackleton calcareous nannoplankton. However, this fact is
and Hall, 1984). The trend found for the late not in contradiction with the trend observed for
Maastrichian at Elles is similar to that observed the planktonic foraminifers because calcareous
at Site 527, though oxygen isotope studies are nannoplankton is less sensitive to depth than
needed to validate this observation. These samples planktonic foraminifers.
where the cool water taxa are present might rep- The early Danian calcareous nannoplankton
resent short surface water cooling pulses, prob- blooms may have played an important role in
ably related to episodic upwellings. Also, though early Danian climate instability, by regulating
the increase in abundance and the palaeo-signi¢- the dissolved pCO2 (Hollander et al., 1993).
cance of P. stoveri just prior to the boundary is Blooms of a few survivor species followed the
not yet elucidated, it might re£ect lower sea-sur- bloom of T. operculata, though these did not
face temperatures during the latest Maastrichtian show blooms in all low-latitudes sections. B. bige-
(Pospichal, 1996). lowii, which is believed to occur preferentially in
According to Girgis (1989) and Faris (1995) the more marginal settings and eutrophic environ-
drop in abundance observed for A. cymbiformis ments (Cunha and Shimabukuro, 1997) rather
might re£ect a decline in surface water palaeotem- than at deep-sea sites (except Site 384) from low
perature in the late Maastrichtian. However, the latitudes, is less than 10% at Elles as well as at El
increase of cool water taxa is not associated with Kef, thus being quite rare in these two marginal
the A. cymbiformis minima. Therefore the drop of settings and never being a dominant species as at
A. cymbiformis relative abundance and its return Caravaca, Gubbio and Bidart (Gardin and Mon-
to smaller forms is probably more linked to echi, 1998; Gorostidi and Lamolda, 1995). It has
change in productivity and water masses charac- never been observed in the Maastrichtian samples
teristics rather than to lowered temperature only. of Elles. A combination of di¡erent factors con-
Gorostidi and Lamolda (1995) suggested decreas- trolled by palaeogeography and palaeodepth and
ing of calcareous nannofossil productivity in the di¡erent distribution of nutrients is thought to be
latest Maastrichtian, based on reduced nannofos- responsible for this.
sil abundance for the Bidart section. Such a de-
crease in abundance was not observed at Elles,
probably because of di¡erent methods of obser- 5. Conclusive remarks
vation; or, it can be due to local palaeoecological
features. The calcareous nannofossil study of the Elles
According to Premoli Silva and Sliter (1995); section encompassed the K^T boundary within
Abramovich et al. (1998); Keller (1996) the K^T about 1 Myr of palaeoecological changes, origina-
exinction operated on a stressed, impoverished, tions, extinctions, and species turnover (Fig. 7).
declining marine ecosystem. Marine regression at ^ The late Maastrichtian portion of Elles revealed
the end of the Maastrichtian and the rapid trans- no clear nannoplankton pre-boundary extinction
gression that followed in the Paleogene could or demise, however, the real stratigraphic oc-
have played a major role as contributor, not as currence of some nannofossil species is hampered
a cause, to the calcareous plankton demise and by palaeoprovincialism, palaeogeography and pa-

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S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231 225

Fig. 7. Summary of the main calcareous nannofossil biostratigraphical and palaeoecological horizons found at Elles.

laeoecology as well as ambiguous taxonomic con- inferred from the planktonic foraminifer record
cepts. M. prinsii ¢rst occurs about 23 m below the from others sites are probably re£ected, though
boundary ; N. romeinii and Chiasmolithus ? sp. are less obviously, by nanno£oral assemblages at El-
present, though extremely rare, in some the Maas- les. These latitudinal gradients which were present
trichtian samples. The presence of Chiasmolithus ? during the latest Maastrichtian especially at high
sp. before the K^T boundary is not considered as latitudes, probably continued during the early Pa-
a contaminant, but its diagnosis must be veri¢ed. leocene. These cooling events are likely responsi-
This ¢nding could lead to the conclusion that ble for a considerable decrease in relative abun-
several Tertiary newcomer species already origi- dance and the return to smaller specimens of A.
nated during the Late Cretaceous. cymbiformis. This observation, in agreement with
^ Late Maastrichtian, surface water cooling pulses previous reports (Girgis, 1989; Faris, 1995) can be
or episodic upwelling are revealed by calcareous the result of water mass and productivity changes
nannofossil assemblages. Short-term climate in the latest Maastrichtian.
changes preceding the K^T boundary event and ^ As in the nearby section of El Kef and other

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226 S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231

low-latitude sites, the Danian nanno£oral assem- event. This investigation does not allow to state if
blages at Elles are characterised by successive the observed changes signi¢cantly contributed to
blooms of surviving and newly evolving species. declining nannoplankton populations and predis-
The point of equilibrium or the complete stabili- posed them to extinction, even considering that
sation of nannoplankton assemblages probably the Earth’s life system during Cretaceous time
took place above NP2 Zone. Though new taxa was much less resilient and more delicately bal-
do appear during the successive blooms, diversity anced than during the Cenozoic or more recent
remained low, pointing to a still mesotrophic, times (Kau¡man, 1988).
poorly strati¢ed marine environment. ^ The rapid and massive nanno£oral turnover at
^ The reworking versus survivorship question is Elles seems strikingly compatible with an extra-
unresolved and will be still a matter of debate. terrestrial impact. However the adequacy of the
The pattern of the Cretaceous taxa found in the calcareous nannofossil record and its correct in-
Danian can, however, not be seen in terms of terpretation across the K^T boundary remains
reworking only and possibly several Cretaceous questionable.
species survived the K^T event in very low num-
bers but never reached high abundance again.
These taxa can represent ‘short-time survivors Acknowledgements
without progeny’ (Harries et al., 1995); the pres-
ence of several coccospheres of W. barnesae and I wish to thank Prof. Gerta Keller (Princeton
D. rotatorius, some of them occurring well above University, NJ, USA) for inviting me to partici-
the boundary can be a further suppport to this pate to the K^T Workshop in Tunis (May 1998).
hypothesis. However, even if we consider that Dr Thierry Adatte (University of Neuchatel,
some of the Cretaceous taxa in Danian sediments Switzerland) is greatly acknowledged for providing
are not reworked, the entity of the turnover is still the Elles log. This manuscript benefited of the
a massive and geologically rapid event, unique in precious and constructive criticisms of Katharina
the calcareous nannofossil evolutionary history. Von Salis (ETH Zentrum, Switzerland) and
^ Calcareous nannoplankton assemblages were Simonetta Monechi (University of Florence,
probably changing in response to various environ- Italy). Mme Isabelle Bauwe and Mr Claude Abrial
mental and climatic stresses which started in the (University of Paris 6) are acknowledged for
late Campanian (Li et al., 1999) and continued technical assistance in the laboratory. CNRS
throughout the Maastrichtian. At this stage of ESA 7073 funds and CNRS Project ‘CRISE-
research, these perturbations may be reasonably VOLE’ supported this work.
interpreted as background climatic/environmental
oscillations rather than foreshadowing the K^T Appendix

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S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231 227

A1. Stratigraphic distribution of calcareous nannofossils. (A) Ranged by FO. (B) Ranged by LO) in the early Danian at Elles.

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228 S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231

A2. Stratigraphic distribution of calcareous nannofossils (ranged by LO) in the late Maastrichtian at Elles.

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S. Gardin / Palaeogeography, Palaeoclimatology, Palaeoecology 178 (2002) 211^231 229

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