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Rapid diversification of planktonic foraminifera in

the tropical Pacific (ODP Site 865) during the late

Paleocene thermal maximum
D. Clay Kelly
Timothy J. Bralower
Department of Geology, University of North Carolina, Chapel Hill, North Carolina 27599
James C. Zachos
Earth Science Board, University of California, Santa Cruz, California 95064
Isabella Premoli Silva
Dipartimento di Scienza della Terra, Universita di Milano, Milan, Italy
Ellen Thomas
Department of Earth Sciences, Wesleyan University, Middletown, Connecticut 06459

The planktonic foraminiferal genera Morozovella and Acarinina rapidly (in ;10 k.y.)
diversified during the late Paleocene thermal maximum (LPTM), giving rise to such mor-
photypes as M. allisonensis (new species), M. africana, and A. sibaiyaensis. Single-specimen
isotopic analysis confirms that M. allisonensis and A. sibaiyaensis are restricted to the LPTM
carbon isotope excursion recorded at Ocean Drilling Program Site 865 (equatorial Pacific
Ocean). The short-lived (50 to several 100 k.y.) excursion taxa attest to the ephemeral
effects of the LPTM on the calcareous plankton. Single-specimen oxygen isotope data show
that evolution of M. allisonensis and A. sibaiyaensis was accompanied by migration to
deeper water depths. Ancestral M. velascoensis and A. soldadoensis were extremely rare or
absent during the early stages of the LPTM, but immigrated back into the study area to
coexist with their descendants in later LPTM horizons. Photosymbiosis may have facili-
tated the morozovellid and acarininid diversifications during the oligotrophic conditions of
the LPTM.

INTRODUCTION and the most dramatic extinction of deep- We report the findings of an investigation
One of the largest and most sudden global sea benthic foraminifera of the past 100 m.y. of the planktonic foraminiferal and calcar-
temperature increases of the Phanerozoic (e.g., Tjalsma and Lohmann, 1983; Thomas, eous nannoplankton responses to the LPTM
Era occurred during the latest Paleocene 1990). The benthic foraminiferal extinction as recorded at Ocean Drilling Program
(;55.5 Ma); Zachos et al. (1993) referred to has been interpreted as resulting from low- (ODP) Site 865. ODP Site 865 (lat
this warming event as the late Paleocene ered levels of dissolved oxygen in oceanic 18826.419N, long 179833.349W) is located
thermal maximum (LPTM). Benthic fora- deep waters (e.g., Thomas, 1990; Kennett atop Allison Guyot in the Mid-Pacific Moun-
miniferal d18O values indicate that oceanic and Stott, 1991). The combined geochemi- tains at a water depth of 1518 m (Bralower
deep waters warmed substantially world- cal and paleontological evidence has led re- et al., 1995). Paleolatitude projections posi-
wide, from ;10 to ;18 8C in high latitudes searchers to postulate that the source area tion Site 865 at ;28N during the late Paleo-
(Kennett and Stott, 1991), and from ;10 to for oceanic deep-water formation tempo- cene (R. Larson, 1994, personal commun.).
;16 8C in the tropics (Bralower et al., 1995). rarily shifted from high- to low-latitude re- Bralower et al. (1995) established that the
Sea-surface temperatures, as recorded in gions during the LPTM (e.g., Miller et al., LPTM d18O and d13C negative excursions
planktonic foraminiferal d18O values, in- 1987). occur over a relatively thin stratigraphic in-
creased from ;14 to ;20 8C at high lati- Although geographic coverage has been terval (;15 cm) in Holes 865B and 865C.
tudes (Kennett and Stott, 1991), but showed adequate to document the global extent of The benthic foraminiferal extinction has
little change at low latitudes (Bralower et the LPTM, the paucity of suitable marine been identified near the bottom of this 15
al., 1995), leading to diminished vertical and sections from equatorial regions has hin- cm interval (102.94 102.96 mbsf [metres be-
latitudinal thermal gradients. dered attempts to identify potential tropical low sea floor] in Hole 865C). Detailed as-
The change from background Paleocene source areas for deep-water formation. This semblage work from this geographically im-
climatic conditions into the LPTM has been shortcoming has been exacerbated by a lack portant site has been combined with
estimated to have taken no more than 10 of information regarding the effects of the measurements of stable isotope ratios of sin-
kyr breakneck speeds by geologic stan- LPTM on surface-water microbiotas. Stud- gle specimens of planktonic foraminifera to
dardswith LPTM conditions persisting for ies (Lu and Keller, 1995) have shown that elucidate the behavior of equatorial Pacific
only 50 to several 100 k.y. (e.g., Kennett and the number of planktonic foraminiferal spe- surface waters during the LPTM.
Stott, 1991; Thomas and Shackleton, 1996). cies increased dramatically during the latest
Coincident with the LPTM was an abrupt Paleocene, but the driving mechanism(s) METHODS
negative isotopic excursion in the global car- and evolutionary mode(s) responsible for We focused on the 100 105 mbsf strati-
bon resevoir (e.g., Kennett and Stott, 1991) this diversification remain obscure. graphic interval of Hole 865C, sampling at

Geology; May 1996; v. 24; no. 5; p. 423 426; 3 figures. 423

Figure 1. Planktonic foraminiferal and nannofossil assemblage changes in uppermost Paleo-
cene strata from ODP Hole 865C. Biostratigraphy at left is after Bralower et al. (1995); arrow
denotes benthic foraminiferal extinction. Relative abundances of planktonic foraminiferal
groups (middle) and nannofossil genera Ericsonia and Toweius (right).
Figure 2. Scanning electron micrographs of
excursion taxon Acarinina sibaiyaensis (1a,
2 6 cm intervals within the isotopic excur- hering particles and then were roasted in ventral view; 1b, edge view) and proposed
evolutionary sequence (2a7b) from Morozo-
sion and 50 70 cm intervals over the re- vacuo at 380 8C. All specimens were ana-
vella velascoensis (2a, ventral view; 2b, edge
mainder of the study interval (Fig. 1). As- lyzed by using an Autocarb device coupled view) to Morozovella allisonensis (7a, ventral
semblage diversity and relative abundance to a Fisons Prism located at the University view; 7b, edge view). All specimens from
data were determined by picking a minimum of California, Santa Cruz. In this system 102.90 mbsf. Scale represents 100 mm.
of 300 planktonic foraminifera from the each sample is reacted in a common phos-
.125 mm size fraction of each sample. phoric acid bath at 90 8C. Average precision
Morphological and paleoecological crite- for samples smaller than 40 mg, as deter- botina group also exhibits a decrease, albeit
ria were used to divide the planktonic fo- mined from replicate analyses of NBS-19 less marked.
raminifera into 11 broadly defined groups. and CM, a laboratory standard, was better Two of the excursion taxa (Acarinina
For comparative purposes, taxa (e.g., the than 60.01 for both d18O and d13C values. sibaiyaensis [Fig. 2, 1a and 1b] and Morozo-
chiloguembelinids, pseudohastigerinids, All isotopic values are reported relative to vella africana) have been described by El
globanomalinids, and Globorotalia imi- the Peedee belemnite (PDB) standard. Naggar (1966) from the upper Paleocene of
tata) with erratic stratigraphic ranges and the Nile Valley of Egypt. Morozovella alli-
very low abundances have been assigned to LATEST PALEOCENE EXCURSION sonensis (Fig. 2, 7a and 7b), a new species, is
a miscellaneous taxa group. MICROBIOTAS the most abundant of the excursion taxa and
Assemblage studies were also conducted High-resolution sampling of latest Paleo- has no previously published record. Cursory
on calcareous nannofossils in 26 samples cene sediments from Site 865 reveals that examination of deep-sea sediments from
across the LPTM. Three hundred specimens the calcareous plankton underwent marked
other latest Paleocene sections (Deep Sea
were counted in each sample. The species changes during the LPTM (Fig. 1). The gen-
Drilling Project [DSDP] Site 577, DSDP
relative abundance data were then grouped era Morozovella and Acarinina dominate
Site 527, and ODP Site 690) suggests that
into genera to more clearly delineate trends. planktonic foraminiferal assemblages and
this unique assemblage is chiefly restricted
Isotopic analyses were conducted on sin- display striking increases in morphological
to tropical regions.
gle specimens of several species belonging variation. This transient diversification gave
to the planktonic foraminiferal genera rise to a suite of distinctive, short-lived mor- Changes in the nannoplankton further at-
Morozovella, Acarinina, and Subbotina. All photypes that are restricted to the 15 cm test to the impact of the LPTM on the sur-
specimens were picked from three strati- interval in which the carbon isotope excur- face-water ecosystem (Fig. 1). Species of
graphic horizons (102.86, 102.90, 102.94 sion is recorded. These excursion taxa Ericsonia (E. cava, E. ovalis, and E. subper-
mbsf) within the LPTM d13C excursion. To reach their peak abundance (;16.5%) at tusa) and Toweius (mostly etched shields of
ensure internal consistency and preclude 102.90 mbsf. Synchronous with the sudden T. eminens and T. pertusus) dominate assem-
any size-dependent isotopic trends (e.g., appearance of the excursion taxa is a nearly blages, accounting for ;80% of specimens
Shackleton et al., 1985), all stable isotope tenfold increase in the relative abundance of counted. Within the 15 cm LPTM interval, a
measurements were performed on speci- the Acarinina soldadoensis group and de- marked (20%) increase is observed in the
mens picked from a restricted size range creases in the Morozovella subbotinae, Igo- relative abundance of Ericsonia, together
(300 355 mm). Individual foraminifera were rina albeari, and weakly keeled morozovellid with a concomitant (20%) decrease in the
sonicated in distilled water to remove ad- groups; the relative abundance of the Sub- number of Toweius.

424 GEOLOGY, May 1996

or A. soldadoensis register excursion d13C
values. This finding suggests that these an-
cestral species emigrated out of the study
Figure 3. Right: Stable carbon
area and/or were too rare to detect during
isotope stratigraphies for upper-
most Paleocene section recov- the early stages of the LPTM. Thus, the ex-
ered from ODP Hole 865C (from cursion group of the 102.94 m horizon is
Bralower et al., 1995). The d13C composed exclusively of M. allisonensis and
curves are based on multispeci- A. sibaiyaensis specimens.
men (8 10) samples of Morozo-
vella velascoensis (circles), Aca-
The local paucity of ancestral forms was
rinina soldadoensis (triangles), temporary, for in subsequent stratigraphic
and Subbotina spp. (pluses). horizons (102.90and 102.86 mbsf), speci-
Left: Bivariate plots of single- mens of M. velascoensis and A. soldadoensis
specimen stable isotope data
with excursion d13C values become more
from three sample horizons
(102.94, 102.90, and 102.86 mbsf) abundant. We infer from this stratigraphic
within carbon isotope excursion. sequence that M. velascoensis and A. solda-
Open circlesM. velascoensis; doensis immigrated back into the study area
solid circlesM. allisonensis; to coexist with their descendants. Similar
open trianglesA. soldadoensis;
solid trianglesA. sibaiyaensis;
evolutionary rates and patterns of paleobio-
and plusesSubbotina spp. geographic variation have been observed in
Shaded areas circumscribe non- the Neogene planktonic foraminifera (Wei
excursion specimens. and Kennett, 1988).
Although M. velascoensis and M. alli-
sonensis coexisted in sample 102.90 m, the
two occupied distinctly different paleoeco-
logical niches. The generally higher d18O
APPLICATIONS OF SINGLE-SPECIMEN merely reflect their deeper depth-habitat values of M. allisonensis (Fig. 3) suggest that
STABLE ISOTOPE DATA and/or lack of photosymbionts (Fig. 3). it inhabited a deeper depth-habitat than M.
Short-lived paleoceanographic events Since the subbotinid d13C values do not sig- velascoensis. The evolutionary trends in both
such as the LPTM are ideally suited for sin- nificantly depart from pre- and postexcur- morphology (axial compression) and ecol-
gle-specimen isotopic analysis (e.g., Stott, sion d13C values (;2.0) reported by ogy (descent to a deeper depth-habitat) re-
1992); it is possible to isotopically distin- Bralower et al. (1995), all 20 specimens from semble those reported by Norris et al.
guish in situ specimens from those that have the 102.90 m sample are considered re- (1993) for the Neogene planktonic forami-
been mixed into the LPTM horizons. In all worked. These findings show that conven- niferal Globorotalia (Fohsella) lineage.
three of the LPTM sample horizons exam- tional counting methods have grossly under- During the later stages of the LPTM
ined, the single-specimen isotopic data sep- estimated the relative abundances of M. (102.86 m in Fig. 3), some of the excursion
arate into two distinct groups offset by allisonensis and associated excursion taxa. specimens of M. velascoensis and most spec-
;2 in their carbon isotope signatures imens of A. sibaiyaensis begin to record rel-
(Fig. 3). This bimodality signifies the pres- Evolutionary and Paleoecological atively higher d18O values. This indicates
ence of both reworked specimens with high Implications that parts of these populations migrated
d13C values (.3) and in situ specimens The responsiveness of the surface-water downward to occupy the same depth-habitat
with anomalously low d13C values (,2). microbiota to oceanographic change is ex- as M. allisonensis. Thus, planktonic forami-
For example, of the 29 randomly picked emplified by the rapid (;10 k.y.) evolution niferal groups (morozovellids and acarini-
specimens of M. velascoensis from the of Morozovella allisonensis from M. velasco- nids), typically regarded as inhabitants of
102.90 m horizon (Fig. 3), only 7 have ex- ensis (Fig. 2, 2a to 7b). The rapidity at which the shallow mixed layer (Boersma et al.,
cursion d13C values. In other words, nearly the short-lived (;50 to several 100 k.y.) M. 1987), descended through the water column
75% of M. velascoensis are contaminants allisonensis and associated excursion taxa to live at depths previously inhabited by the
mixed into the 102.90 m sample; this rough evolved ranks among the fastest known in genus Subbotina. A notable exception to this
estimate suggests that the M. velascoensis the pelagic protista. The suddenness of this paleoecological change is A. soldadoensis,
group composes considerably less of the evolutionary event raises the possibility that which appears to have retained its near-sur-
102.90 m assemblage than the reported these excursion taxa evolved elsewhere and face habitat, as inferred from its relatively
15%. Likewise, the Acarinina soldadoensis immigrated into the study area. However, low d18O values.
group appears to be about half as abundant the presence of transitional forms linking
in the 102.90 m sample as conventional as- M. allisonensis to M. velascoensis and A. PALEOCEANOGRAPHIC
semblage counts indicate. Thus, all three sibaiyaensis to A. soldadoensis argues against INTERPRETATIONS
nonexcursion groups are composed solely of such a scenario. The unique microfossil assemblages asso-
specimens of ancestral species that pre- and The stratigraphic succession of single- ciated with the LPTM interval at Site 865
postdate the LPTM carbon isotope excur- specimen isotopic data (Fig. 3) taken from beg the question, What paleoceanographic
sion. All specimens of M. allisonensis (44 of within the carbon isotope excursion dis- conditions triggered the transient biotic re-
44) and A. sibaiyaensis (28 of 28) recorded closes interesting paleobiogeographic and sponses seen in the calcareous plankton?
anomalously low d13C values, confirming paleoecologic trends. In the lowermost ex- The diverse but impoverished benthic fo-
that these morphotypes are restricted to the cursion sample analyzed (102.94 mbsf, raminiferal faunas at Site 865 suggest that
LPTM carbon isotope excursion interval. which corresponds to the benthic foraminif- the overlying surface waters were highly oli-
The relatively low d13C values measured eral extinction horizon), none of the ran- gotrophic during the late Paleocene. Al-
from members of the genus Subbotina domly picked specimens of M. velascoensis though depauperate, these assemblages do

GEOLOGY, May 1996 425

exhibit a modest increase in accumulation the euphotic zone to deepen at Site 865, 1991, Similarities between planktonic and
rates and become dominated by Tappanina allowing the photosymbiont-bearing moro- larger foraminiferal evolutionary trends
selmensis and several buliminid taxa within zovellids and acarininids to explore deeper through Paleogene paleoceanographic
changes: Palaeogeogeography, Palaeocli-
the 15 cm LPTM interval. These changes depth-habitats. matology, Palaeoecology, v. 83, p. 49 64.
could signify an increase in surface-water Kennett, J. P., and Stott, L. D., 1991, Abrupt
productivity, leading to an increased flux of CONCLUSIONS deep-sea warming, paleoceanographic
particulate organic matter to the sea floor. The near-surface dwelling planktonic fo- changes and benthic extinctions at the end of
raminifera rapidly (;10 k.y.) diversified the Palaeocene: Nature, v. 353, p. 225229.
However, we attribute the benthic forami- Lu, G., and Keller, G., 1995, Ecological stasis and
niferal assemblage changes to increased during the late Paleocene thermal maximum
saltation: Species richness change in planktic
preservation of particulate organic matter (LPTM) at ODP Site 865. The short-lived foraminifera during the late Paleocene to
on the sea floor due to decreased oxygen- (50 to several 100 k.y.) taxa Morozovella al- early Eocene, DSDP Site 577: Palaeogeog-
ation in temporarily warmer waters. lisonensis (new species), M. africana, and raphy, Palaeoclimatology, Palaeoecology,
Acarinina sibaiyaensis originated during this v. 117, p. 211227.
In nannoplankton assemblages, Peleo- Miller, K. G., Janacek, T. R., Katz, M. E., and
Alampay and Wei (1993) noticed an in- transient episode of paleoceanographic Keil, D. J., 1987, Abyssal circulation and
crease in the abundance of the warm-water change. Single-specimen isotopic data show benthic foraminiferal changes near the Pa-
genus Discoaster within the LPTM at Sites that all specimens of M. allisonensis and A. leocene/Eocene boundary: Paleoceanogra-
sibaiyaensis are restricted to the LPTM car- phy, v. 2, p. 741761.
577 and 690. The observed increase in Norris, R. D., Corfield, R. M., and Cartlidge, J. E.,
Ericsonia, a group that is more common bon isotope excursion. Single-specimen iso-
1993, Evolution of depth ecology in the
in low-latitude water masses, and decrease topic analysis also reveals that the morozo- planktic foraminifera lineage Globorotalia
in Toweius, which is more abundant in mid- vellids and acarininids underwent parallel (Fohsella): Geology, v. 21, p. 975978.
to high-latitude oceans, is also consistent patterns of paleoecological and paleobioge- Okada, H., and Bukry, D., 1980, Supplementary
ographical variation during the evolution of modification and introduction of code num-
with warming (Fig. 1). However, because bers to the low latitude coccolith biostrati-
sea-surface temperatures at Site 865 only in- the LPTM taxa. We postulate that this sud- graphic zonation (Bukry, 1973; 1975): Ma-
creased by ;1 8C (Bralower et al., 1995), it den diversification was related to extreme rine Micropaleontology, v. 5, p. 321325.
is conceivable that another paleoecological oligotrophy caused by changes in the ther- Peleo-Alampay, A., and Wei, W., 1993, Nanno-
mal structure of the water column. The fossil response to the abrupt warming event
change (e.g., oligotrophy) caused the nan- near the Paleocene/Eocene boundary: Its pa-
noplankton assemblage response. Indeed, ephemeral effects of the LPTM on the cal-
leoceanographic and paleoclimatic implica-
Ericsonia and Discoaster are considered to careous plankton are in striking contrast to tions: International Nannoplankton Associ-
have oligotrophic affinities and Toweius eu- the coeval, global mass extinction suffered ation Newsletter, no. 15, p. 83.
trophic affinities (M.-P. Aubry, 1995, per- by the deep-sea benthic foraminifera. Shackleton, N. J., Corfield, R. M., and Hall,
M. A., 1985, Stable isotope data and the on-
sonal commun.). ACKNOWLEDGMENTS togeny of Paleocene planktonic foraminif-
The preponderance of morozovellids and Supported by National Science Foundation era: Journal of Foraminiferal Research,
acarininids, plus the dearth of subbotinids, grants to Bralower (EAR-94-05784) and Zachos v. 15, p. 321336.
and Thomas (EAR-94-06099). Financial assist- Stott, L. D., 1992, Higher temperatures and lower
within the LPTM interval at Site 865 is con- ance also provided by JOI/USSAC and Martin pCO2: A climate enigma at the end of the
sistent with oligotrophic conditions. DHondt Research Fellowships to Kelly. We thank R. Nor- Paleocene epoch: Paleoceanography, v. 7,
et al. (1994) noted that the stable isotope ris and M. Leckie for critically reviewing the man- p. 395 404.
signatures of the morozovellids and acari- uscript, R. Olsson and W. Berggren for helpful Thomas, E., 1990, Late Cretaceous early Eocene
ninids are comparable to those of modern, discussions, and K. Billups for laboratory mass extinction in the deep-sea, in Sharpton,
assistance. V., and Ward, P., eds., Global catastrophes:
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426 Printed in U.S.A. GEOLOGY, May 1996