TMP 4 F17

Author E-Print 1/3/2012
MAASTRICHTIAN PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY AND

PALEOENVIRONMENT OF BRAZOS RIVER, FALLS COUNTY, TEXAS, U.S.A.
SIGAL ABRAMOVICH
Dept. of Geological & Environmental Sciences, Ben-Gurion University of the Negev, Beer Sheba, 84105, Israel
e-mail: sigalabr@bgu.ac.il
GERTA KELLER
Department of Geosciences, Princeton University, Princeton NJ 08544, U.S.A.
t
ZSOLT BERNER
in
Institute for Mineralogy & Geochemistry, University of Karlsruhe, 76128 Karlsruhe, Germany
MATAN CYMBALISTA AND CARMI RAK

Dept. of Geological & Environmental Sciences, Ben-Gurion University of the Negev, Beer Sheba, 84105, Israel
Pr
ABSTRACT: Investigation of late Maastrichtian faunal and environmental changes in three subsurface wells spanning over 3 km along the Brazos River,
Texas, reveals similar minimum-diversity high-stress assemblages associated with shallow shelf conditions. Upper Maastrichtian sediments recovered
span planktic foraminiferal (CF) zones CF1 to CF4 in well Mullinax-1 to the north and well KT3 at Cottonmouth Creek, and zones CF1-CF2 in
Mullinax-3 at Darting Minnow Creek. Biotic stress conditions are demonstrated by the minimum species richness, near exclusion of larger specialized
species, dwarfing, and dominance of small generalist taxa. Faunal assemblages dominated by alternating abundances of the disaster opportunist
Guembelitria cretacea (Cushman) (particularly in zones CF4 and CF2-CF1) and heterohelicid species [e.g., Heterohelix globulosa (Ehrenberg), H.
E-
planata (Cushman), Paraspiroplecta navarroensis (Loeblich)]. Other small surface and subsurface mixed-layer dwellers are rare to common (e.g.,
hedbergellids, globigerinellids, pseudoguembelinids).
The coincidence of Guembelitria blooms with lithological changes and oxygen and carbon stable isotope excursions may represent discrete episodes
of freshwater runoff related to short-term pulses of the latest Maastrichtian (zone CF1-CF2) global climate warming. Climate warming ended in the upper
part of zone CF1 with the return to a cooler climate, lower sea level, and the formation of incised valleys in a coastal–lagoonal environment. With the
subsequent early transgression, incised valleys were infilled by a sandstone complex with reworked impact spherules, as well as lithified clasts with
impact spherules up to 80 cm below the KT boundary. In this environment the shallow inner-neritic setting superimposed by changes in sea level and
climate is the most probable cause for the observed conditions of high biotic stress preceding the KTB in the Brazos area.
KEY WORDS: Index Words: Brazos Texas, Biostratigraphy, Paleoenvironment, upper Maastrichtian, planktic foraminifera, Faunal Turnover, Depositional
or
environment
INTRODUCTION complex. Smith and Pessagno (1973) reported on the taxonomy and
excellent preservation of some Maastrichtian samples from various
Most studies of the Brazos River area (Falls County, Texas) have localities between Dallas and College Station in north-central Texas.
th
focused on the Cretaceous–Tertiary boundary (KTB) (also called K– No stratigraphic sections or precise localities of the samples are
Pg) transition and in particular the sandstone complex with Chicxulub provided. For the Brazos outcrops in Falls County, eight samples are
impact spherules at the base, which is interpreted by some as the listed, all of which contain Gansserina gansseri (Bolli), which
deposit of an impact-generated megatsunami (e.g., Bourgeois et al., indicates that the age of these samples is probably early late
1988; Hansen and Upshaw, 1990; Hansen et al., 1993a; Hansen et al., Maastrichtian. It was not until 1986 that the first Maastrichtian
1993b; Heymann et al., 1998; Schulte et al., 2006). Consequently, sequences were recovered by Thor Hansen and Earl Kaufman by rotary
Au
these workers placed the KTB at the base of this sandstone complex drilling of two wells (KT1, KT2) approximately 350 m south of the
based on the assumption that the Chicxulub impact caused the KTB Highway 413 Bridge across the Brazos River and a third well (KT3) on
mass extinction and therefore defines the KTB (see Schulte et al., Cottonmouth Creeek (Fig. 1). Few studies have concentrated on the
2008; Schulte et al., 2010; Keller et al., 2008; Keller, this volume, for Maastrichtian interval of these wells. Keller (l989a) reported on the
arguments pro and con of this circular reasoning). Other workers biostratigraphy of the KT3 well, but the age remained uncertain
interpreted the sandstone complex as a submarine-channel infill because the standard late Maastrichtian index species Abathomphalus
associated with the latest Maastrichtian sea-level lowstand and mayaroensis (Bolli) is absent in these shallow-water sequences. In
subsequent rise, with the KTB well above it (e.g., Donovan et al., subsequent studies of El Kef and DSDP Site 525A, Li and Keller
1988; Keller, 1989a; Keller et al., 1989b; Beeson, 1992; Médus, 1992; (l998a, 1998b, 1999) refined the Maastrichtian biozonation and
Gale, 2006; Keller et al., 2007a; Keller et al., 2009a, Keller et al., this subdivided the A. mayaroensis zone into four biozones, CF1 to CF4.
volume). Based on this biozonation, Schulte et al. (2006) assigned the 7.5 m of
Older Maastrichtian sediments from the Brazos area are little known uppermost Maastrichtian sediments below the sandstone complex of
because outcrops are limited to approximately 1 m below the sandstone well KT2 (which they called Brazos core 1) to zones CF1-CF2 and
The End-Cretaceous Mass Extinction and the Chicxulub Impact in Texas

SEPM Special Publication No. 100, Copyright Ó 2011
SEPM (Society for Sedimentary Geology), Print ISBN 978-1-56576-308-1, CD/DVD ISBN 978-1-56576-309-8, p. 123–156.
124 SIGAL ABRAMOVICH, GERTA KELLER, ZSOLT BERNER, MATAN CYMBALISTA AND CARMI RAK
For this study we chose well Mullinax-1, which contains a nearly

complete and undisturbed Maastrichtian subsurface record spanning
36 m below the sandstone complex. In addition, we reanalyzed and
updated the taxonomy of the old well KT3 first published by Keller
(1989a), which spans a 23-m-long subsurface record below the
sandstone complex. Mullinax-3, to the south, spans only the uppermost
Maastrichtian zones CF1-CF2, but is added for completeness of the
Brazos north–south transect showing the shallowing environment
towards the shoreline over a distance of little more than 3–4 km (Fig.
1). These three wells provide insights into the nature and complexity of
shallow water environments in the Western Interior Seaway and how
they were affected by changes in sea level and climate.
t
The main objectives of this study are to: (1) document the age and
biostratigraphy of the Brazos area based on planktic foraminifera, (2)
in
evaluate the climatic and environmental changes during the late
Maastrichtian based on stable isotopes, (3) evaluate paleoenvironmen-
tal conditions based on planktic foraminiferal assemblages, including
species richness, nature and composition of assemblages, and species
dwarfing, and (4) document species assemblages through SEM
Pr
illustrations as guide to future identification of species in shallow
water environments of the Western Interior Seaway.
METHODS
Maastrichtian cores of Mullinax-1 and Mullinax-3 were measured,
described, photographed, and sampled at an average sampling step of
E- 10–20 cm. Planktic foraminifera were processed using standard
techniques (Keller et al., 1995). Samples were soaked overnight in
dilute (10%) H2O2, then gently washed through sieve sizes 38–63 lm
and 63–150 lm to recover very small, small, and large planktic
foraminifera. The washed residues were dried in the oven at 508C. Core
sample size was generally restricted to 5 cm3, except for intervals
where too few specimens were recovered for quantitative analysis and
therefore sample size was doubled. In some intervals even 10 cm3
samples yielded insufficient foraminiferal tests for quantitative
or
analysis. In these cases several samples were grouped to yield
FIGURE 1.—Map of Cretaceous–Tertiary boundary (KTB) outcrops and representative assemblages. Larger samples were provided by Thor
wells drilled along the Brazos River, Cottonmouth Creek, and Hansen for the KT3 cores in 1987, and this has yielded a significantly
larger number of rare species.
Darting Minnow Creek of Falls County, Texas. Wells Mullinax-1,
Quantitative analysis was conducted on two size fractions, . 63 lm
Mullinax-3, and KT3 were analyzed for this study. and . 150 lm, to obtain optimum representation of both smaller and
th
larger species. In Mullinax-3 the . 38 lm size fraction was also analyzed

because species dwarfing is severe in the latest Maastrichtian and early
nannofossils zone CC26, which span the last ; 300 kyr (time scale of Danian. For each sample of the two size fractions an aliquot of 250–300
Gradstein et al., 2004) of the Maastrichtian. No older Maastrichtian specimens was picked whenever possible, mounted on microslides, and
sediments have yet been studied from well KT2. identified. Benthic specimens were counted in the same aliquots of the .
In 2005 three wells, 22–30 m deep, were funded by the National 63 lm size fraction as an indicator of sea-level changes. The remaining
Au
Science Foundation (Continental Dynamics Program and Sedimentary residues were examined for rare species, and these were noted for data on
Geology and Paleobiology Program) and drilled by DOSECC species range. The 38–63 lm size fraction was examined for very small
(Drilling, Observation and Sampling of the Earth’s Continental Crust) species that may not be present in the larger size fractions, particularly in
with a CS-500 rig under the guidance of Jerry Baum and Gerta Keller. the early Danian. Scanning electron micrographs of Late Cretaceous
The first well, Mullinax-1, is located at the same GPS location (318 planktic foraminifera are illustrated in Plates 1–8 and the quantitative
07.53 0 N, 968 49.30 0 W) as the old wells KT1 and KT2 of Hansen and foraminiferal data are listed in Appendix I.
Kaufman (Brazos core 1 and 2 of Schulte et al., 2006) (Fig. 1). This Stable isotopes are based on well-preserved specimens of the benthic
location was chosen to recover the expanded KTB sequence of the foraminifer Lenticulina spp. in the size fraction 150–250 lm with little
nearby outcrop Brazos-1, which was also evident in the old wells KT1 or no sediment infilling chambers. Pseudoguembelina costulata
and KT2 that encountered drilling disturbance (Thor Hansen, written (Cushman) was analyzed as indicator of surface water conditions.
communication, 1987). Wells Mullinax-2 and Mullinax-3 were drilled Heterohelix globulosa was previously analyzed for the KT3 well
as overlapping cores 1.2 m apart on a meadow approximately 150 m (Barrera and Keller, 1990). Stable isotope analysis was performed
from the Darting Minnow Creek waterfall (GPS Location 318 06.55 0 N, using a fully automated carbonate preparation system (MultiCarb)
968 50.28 0 W). Core recovery at was 95% Mullinax-1 and 100% at connected on-line to an isotope ratio mass spectrometer (Optima,
Mullinax-2 and Mullinax-3. Of these new wells only the KTB Micromas Ltd., UK). Isotope ratio values are reported relative to NBS-
transition of Mullinax-1 has been published (Keller et al., 2007a; Keller 19 with d13C¼ 1.95ø (V-PDB). Precision, assessed on the basis of
et al., this volume). repeated measurements of the carbonate standard, was generally better
MAASTRICHTIAN, BIOSTRATIGRAPHY, PALEOENVIRONMENT, BRAZOS TEXAS 125
than 0.06ø for each analytical batch. Stable isotope data are listed in scoured surface of submarine channels that marks an erosional surface.
Appendix II. Clasts from the underlying mudstone and lithified clasts, some with
spherules and mudcracks, are commonly found above the erosion
LITHOLOGY surface and indicate erosion from an older impact spherule layer
(Keller et al., 2007a; Adatte et al., this volume). Above these large
Upper Maastrichtian sediments of the Brazos area have been clasts are three upward-fining glauconitic, shell-rich, and spherule-rich
recovered in three wells; all three span the entire upper Maastrichtian layers with smaller claystone and phosphatic clasts. An erosional
from zones CF4 through CF1 (wells Mullinax-1, KT3), but only the surface separates these layers from the overlying hummocky cross-
uppermost Maastrichtian zones CF1 and CF2 were analyzed in well bedded sandstone (HCS) with large burrows truncated at the top.
Mullinax-3. Detailed lithologic descriptions are available only for the Laminated dark gray and light gray silty mudstone with small
Mullinax wells, which are briefly described here and illustrated for Chondrites burrows mark the top of the sandstone complex. The 80
Mullinax-1 and Mullinax-3. Photologs and descriptions of each well cm between the top of the sandstone complex and the KTB consists of
t
are provided in Appendix III of this volume. dark gray mudstones with few small shells and burrows, some infilled
with framboidal pyrite. There is no lithological change at the KT
in
Well Mullinax-1 boundary.
The base of well Mullinax-1 stratigraphical succession consists of Wells Mullinax-2 and Mullinax-3
poorly sorted, argillaceous sandstone with a mud matrix, irregular
bedding, and common sand-filled burrows. Sharp contacts mark the A total of 23.77 m of sediments was recovered in Well Mullinax-3,
base (44.32 m) and top of this sandstone (40.73 m; Fig. 2). Above the which is located approximately 150 m from the Darting Minnow Creek
Pr
sandstone (40.75 m to 29.5 m) sediments consist of medium gray waterfall. This well was double cored to ensure complete recovery.
sandy mudstone with abundant fine-grained sand and mud laminae, Only the upper interval of Mullinax-3 was analyzed from 3 m to 11 m
some sand lenses indicating low-amplitude ripples, shell fragments, depth spanning the early Danian and latest Maastrichtian zones CF1
and sand filled burrows. Sand content decreases upwards. A layer of and upper CF2 (Fig. 4). Wells Mullinax-2 and Mullinax-3 are
framboidal pyrite concretions occurs at 29.6 m followed by a sharp important for their very different lithologies across the KTB transition,
basal contact approximately 10 cm above it. Above this disconformity the absence of a sandstone complex with impact spherules, the
sediments consist of dark gray thick-bedded sandy mudstones with occurrence of frequent disconformities and erosion surfaces, and
E-
framboidal pyrite concretions, common sand laminae, small macro-
fossils, sand-filled burrows, and rare ammonites Discoscaphites iris
(Conrad), which is indicative of the uppermost Maastrichtian
common roots indicating a coastal to lagoonal environment.
In the lower part of the analyzed sequence (11.0–7.3 m) the lithology
consists of thick-bedded dark gray mudstone, fractured in layers,
ammonite zone in North America; Neil Landman, written communi- common framboidal pyrite, few small macrofossils, and rare
cation 2005; see also Kennedy et al., 2001. This lithologically ammonites, similar to well Mullinax-1 (Fig. 4). At 7.26 m is a sharp
monotonous interval spans from 29.5 to 16.5 m, but is interrupted at contact followed by another sharp and undulose contact at 6.86 m. The
21.95 m by a 4-cm-thick silty mudstone with abundant shells and shell interval above is a silty mudstone with weathered yellow streaks, few
fragments that may represent a tempestite (Fig. 2). About 25 cm above fossils, common roots, up to 12 cm in length, and pyrite.
is a 12-cm-thick dark gray fine-grained sandstone followed by light Disconformities occur at 5.9 m and 5.6 m. The latter marks the KT
or
gray concretions with carbonate cement. Another thin sandy mudstone boundary, as a major erosion surface and hiatus. Above it is a sandy
with abundant shells (a tempestite?) is present at 20.90 m. At 18.95 m a mudstone with quartz pebbles, clasts, and few small shells with a
gradational contact with a laminated upward-coarsening sandstone gradational contact at 4.5 m. In the upper part of the sequence a
with sand-filled burrows and phosphatic gastropods indicates a very sandstone with gastropods, shells, burrows, rizoliths, and phosphate
shallow depositional environment (Fig. 3). The upper contact with the nodules ends with a sharp contact 3.5 m.
overlying silty mudstone at 18.35 m is gradational, with sand laminae Disconformities and gradational and erosional contacts in Mullinax-
th
and sand-filled burrows. At 16.5 m the silty, sandy mudstone is 3 mark sea-level fluctuations in a shallow coastal to lagoonal
overlain by dark gray, thick-bedded mudstone with small and large environment. The common presence of roots in the upper part of zone
macrofossils. Uppermost Maastrichtian lithology from 16.5 to 8.6 m CF1 indicates temporary subaerial exposure, which appears to be
consists of monotonous dark gray, bedded mudstone with macrofossils, restricted to the southern Brazos area (Darting Minnow Creek, Fig. 1).
rare ammonites, and common burrows. This sequence is interrupted by The unconformity at the KTB also indicates that this area was
changes from dark gray to lighter gray that indicate increased shallower than all other localities analyzed.
Au
terrigenous influx and transitions from dysoxic to oxic conditions on

the sea floor. Each of these lithological changes coincides with major BIOSTRATIGRAPHY
negative d13C and d18O excursions that suggest fresh-water influx at
these times (Fig. 3). The first contact at 12.10 m marks the transition Biozonation
from fissile medium gray mudstone with rare macrofossil shells to
lighter gray mudstone with common shells. The second lithological Maastrichtian biostratigraphy used in this study is based on the high-
change at 11.10 m marks a sharp contact between dark gray and resolution CF (Cretaceous Foraminifera) biozonation of Li and Keller
medium gray mudstone. The third lithological change marks a (1998a, 1998b, 1999) which subdivides this stage into eight biozones
transition from dark gray mudstone and medium gray mudstone with (CF1-CF8; Fig. 5). For the KTB transition the early Danian zonal
common shell fragments. scheme of Keller et al. (1995) is used. For this study the biozonation
The most prominent lithological change of the upper Maastrichtian and index species are illustrated in Figure 6.
is to the sandstone complex between 8.0 and 8.65 m (Fig. 3), which has The CF zonal scheme was originally proposed based on El Kef,
been variously interpreted as an impact0-generated tsunami deposit Tunisia, and DSDP Site 525A (Walvis Ridge, South Atlantic). Since that
(e.g., Bourgeois et al., 1988; Smit et al., 1996; Heymann et al., 1998; time these biozones have been tested, applied, and in some cased refined
Schulte et al., 2006; Schulte et al., 2008), or sea-level lowstand systems in Maastrichtian sequences worldwide (e.g., Abramovich et al., 2002;
tract (e.g., Keller, 1989a; Gale, 2006; Keller et al., 2007a; Keller et al., Abramovich et al., 2003; Abramovich et al., 2010; Abramovich and
2009a; Adatte et al., this volume). The sandstone complex overlies the Keller, 2002, 2003; Keller, 2002, 2003, 2004; Keller et al., 2007a; Keller
t
in
Pr
E-
or
th
Au
FIGURE 2.—Lithology and description of the upper Maastrichtian Well Mullinax-1, zones CF4-CF3.
et al., 2007b; Keller et al., 2009a; Keller et al., 2009b). They have proved which places the KT boundary close to 66 Ma (Sam Bowring, personal
to be reliable age indicators worldwide, although variations may occur, communication, 2009).
particularly in areas of poor fossil preservation, shallow-water sequences, The Maastrichtian biozones of Li and Keller (1998a, 1998b) are
or high latitudes, where some index species may be sporadically present or shown in comparison with biozonal schemes of Caron (1985) and
missing and/or their stratigraphic ranges shortened. In earlier studies the Huber et al. (2008) (Fig. 6). All three biostratigraphic schemes are
ages of the CF Zones were calculated based on the time scale of Cande and based mainly on datum events of large–size keeled globotruncanid and
Kent (1995) and Berggren et al. (1995), which placed the KTB at 65 Ma. racemiguembelinid species. Huber et al. (2008) recently proposed
They are here recalculated based on the chronostratigraphic update of modifications to Caron’s (1985) biozonation based on deep-sea sites on
Gradstein et al. (2004) who placed the KT boundary at 65.5 Ma (Fig. 6). Blake Nose (DSDP Site 390A, ODP Sites 1049, 1050, 1052), where
However, further updates appear likely based on ongoing U/Pb dating, they encountered rare and sporadic occurrences of some index species
t
in
Pr
E-
or
th
FIGURE 3.—Lithology and description of the upper Maastrichtian well Mullinax-1, zones CF3, CF2, and CF1 with stable isotopes, Guembelitria
data and sea-level interpretation.
(e.g., Gansserina gansseri, Globotruncana aegyptiaca Nakkady). based on sections from Egypt. It was later dated by Pardo et al. (1996)
Au
They proposed to subdivide the long Abathomphalus mayaroensis to correspond to the final 300 ky of the Maastrichtian based on
zone, which spans the upper Maastrichtian (equivalent to zones CF1 to calibration with the paleomagnetic record at Agost, Spain. Based on
CF4) into two zones (Pseudoguembelina hariaensis and A. mayar- the new chronostratigraphic update of Gradstein et al. (2004), which
eoensis) and replace the lower Maastrichtian G. gansseri and G. dated the KT boundary at 65.5, Zone CF1 is considerably reduced,
aegyptiaca zones of Caron (1985) (equivalent to CF5-CF8) by spanning just the last 160 ky (between 65.5 and 65.66 Ma) of the
Racemiguembelina fructicosa and P. palpebra zones (Fig. 6). Maastrichtian. The CF1 Zone is now routinely applied in low-latitude
In the Brazos well Mullinax-1, the CF zonation indicates that the regions, particularly of the Tethys Ocean (Keller et al., 2007a; Keller et
entire late Maastrichtian interval from Zone CF4 up to Zone CF1 was al., 2009a; Keller et al., 2009b; Keller et al., this volume; Abramovich
recovered along with the early Danian (Keller et al., this volume). et al., 2010).
Zones CF1 to CF4 correspond to the P. hariaensis and A. mayaroensis In the Brazos well Mullinax-1, as well as the other wells, Plummerita
Zones of Huber et al. (2008). In the Mullinax-1 well, the keeled hantkeninoides (Brönnimann) is rare and sporadically present, with
globotruncanid and racemiguembelinid species are sporadically few specimens exhibiting the characteristics of well-developed asteroid
present or exhibit juvenile or dwarfed morphologies, which introduces tests. Based on the lowest (11.09 m) occurrence of P. hantkeninoides in
some uncertainty into determinations of zonal boundaries. Mullinax-1, Zone CF1 spans 3.85 m; this includes the 80 cm of Zone
CF1 above the sandstone complex (Keller et al., 2007a) (Fig. 7). The
Plummerita hantkeninoides CF1 Zone: This zone was first average sedimentation rate is 2.3 cm/ky (Fig. 8). The presumed
defined by Masters (1993) as the total range of the nominate species, ancestor Rugoglobigerina reicheli Brönnimann is more common and
t
in
Pr
E-
FIGURE 4.—Lithology and description of the upper Maastrichtian well Mullinax-3, zones CF1 and CF2 with stable isotope data.
or
th
Au
FIGURE 5.—Maastrichtian planktic foraminiferal zonal schemes and ages of biozones based on the revised time scale with the KTB at 65.5 Ma.
t
in
Pr
E-
or
FIGURE 6.—Illustration of Late Maastrichtian planktic foraminiferal CF biozones used in this study. (CF ¼ Cretaceous Foraminifera).
first appears earlier at 16.99 m. In well Mullinax-3 Zone CF1 spans 3.2 The Gansserina gansseri acme event was not recognized in the
m but is terminated at the top by a hiatus (Fig. 9). In well KT3 Zone Brazos wells, and this species is rarely present. Its absence may be due
th
CF1 spans 1.4 m (Fig. 10). The shorter stratigraphic range is partly due to the shallow shelf environment of the Brazos area in the Western
to a less complete sedimentary record and partly to the rarity of the Interior Seaway. A hiatus that spans the entire G. gansseri acme event
index species. Zone CF1 in the Brazos sections contains the sandstone (i.e., upper CF5-CF2 interval) can be excluded because the nominate
complex with its reworked Chicxulub impact spherules. This sandstone species of Zone CF3, Pseudoguembelina hariaensis Nederbragt is
complex and its implications for the age of the Chicxulub impact are present. The base of Zone CF2 at well Mullinax-1 was recognized
discussed in Keller et al. (this volume). based on the presence of a single specimen of G. gansseri at 15.95 m.
Au
Based on this occurrence Zone CF2, which lasted 120,000 years, spans
Pseudoguembelina palpebra CF2 Zone: This zone is defined by 4.86 m, with an estimated sedimentation rate of 4.1 cm/ky (Figs. 7, 8).
the last appearance (LA) of Gansserina gansseri at the base and first In well KT3, G. gansseri is more frequently present, with the last
appearance (FA) of Plummerita hantkeninoides at the top (Li and appearance at 10.72 m and spanning 2.37 m, and an estimated
Keller, 1998a). Based on the time scale of Gradstein et al. (2004) the sedimentation rate of 2.27 cm/ky (Fig. 10). The lower sediment
age duration of this biozone is 65.66–65.78 Ma, or 120,000 years (Fig. accumulation rate compared with Mullinax-1 is probably due to more
6). Gansserina gansseri is considered to have been a deep–cold water- condensed sedimentation and/or a short hiatus. The base of Zone CF2
dwelling keeled globotruncanid (Abramovich et al., 2003; Abramovich was not recovered in Mullinax-3.
et al., 2010). This species is also observed in shallow-water sequences,
including Egypt, Argentina (Keller, 2002; Tantawy, 2003; Keller et al., Pseudoguembelina hariaensis CF3 Zone: Zone CF3 was defined
2007b), and the Brazos sections. Its presence in these sequences was by Li and Keller (1998a) as the interval between the FA of P. hariaensis
interpreted as representing the expansion of cold water into shallower at the base and the LA of G. gansseri at the top. This zone is not
environments in the tropical zone. In these localities, the LA of G. equivalent to the P. hariaensis Zone of Huber et al. (2008). In the Blake
gansseri typically marks the termination of an acme event of this Nose localities analyzed by Huber et al. (2008) they did not recognize
species. This acme event was found to coincide with a global cooling the Plummerita hantkeninoides Zone and therefore extended the P.
event between ; 68 and 65.8 Ma from the upper part of Zone CF5 up hariaensis Zone to the end of the Maastrichtian, thus spanning our
to the base of Zone CF2 (Abramovich et al., 2010). zones CF3-CF1 from 66.78 to 65.5 Ma.
t
in
Pr
E-
or
FIGURE 7.—Species ranges of planktic foraminifera in well Mullinax-1 along with species richness (three-point average) of small species (63–150
lm) and larger species (. 150 lm) and planktic/benthic ratio. Gray-shaded bands in CF3 mark intervals with very rare foraminifera and where
several samples were combined to obtain statistically representative samples. Gray-shaded band in CF1 marks the sandstone complex with
reworked impact spherules.
th
In well Mullinax-1, the FA of P. hariaensis occurs at 26.64 m. this biozone spans from 68.36 to 66.99 Ma, for a duration of 1.37 My,
Therefore Zone CF3 spans 10.69 m and lasted 1.21 My based on the (Fig. 6). Zone CF4 is not coeval with the R. fructicosa Zone of Huber et
recalculated age of 65.78 to 66.99 Ma. The average sediment al (2008), who assigned a significantly older age for the FA of this
accumulation rate for this interval is 0.88 cm/ky. In well KT3, the species at 68.72 Ma based on the Blake Nose sections. This age
Au
FA of P. hariaensis occurs at 19.70 m and Zone CF3 spans 8.98 m. The discrepancy may be a taxonomic artifact related to identifying the point
average sediment accumulation rate is thus 0.74 cm/ky (Fig. 7). These of transition from R Racemiguembelina powelli Smith & Pessagno to
sedimentation rates are significantly lower than for Zones CF1 and R. fructicosa. Zone CF4 mostly overlaps with the Abtahomphalus
CF2. Huber et al. (2008) placed the FA of P. hariaensis Zone at 66.78 mayaroensis Zone of Huber et al (2008) (Fig. 6). The index taxon A.
Ma, approximately 200 ky younger then the revised age of Li and mayaroensis is absent in the shallow depositional environment of the
Keller (1998a) (Fig. 6), though this only marginally raises the Brazos area because this species was a deep-water dweller.
sedimentation rate to a maximum of 1 cm/ky. The reason for the Racemiguembelina fructicosa is very rare. This species did not
lower sediment accumulation rate is probably one or more hiatuses. For survive, probably because of the frequently eutrophic conditions which
example, in Mullinax-1, a sandstone bed marks a sharp contact with are unfavorable to photosymbiotic species such as R. fructicosa. This is
the mudstone at 18.5 m. In both wells, the base of CF3 follows an also indicated by the rare occurrences of the other members of this
interval of high abundance in the disaster opportunist G. cretacea, lineage in Brazos sections (e.g., Racemiguembelina powelli and
suggesting high stress conditions. Pseudotextularia intermedia De Klasz). For these reasons, the base of
Zone CF4 in the Brazos River area cannot be determined confidently
Racemiguembelina fructicosa CF4 Zone: This biozone is defined based on the nominate taxon.
by the first appearance (FA) of R. fructicosa (Egger) at the base and the Huber et al. (2008) suggested that Pseudotextularia elegans
FA of P. hariaensis at the top (Li and Keller, 1998a, 1998b). Based on (Rzehak) serves as a secondary marker species in the absence of R.
the placement of the KT boundary at 65.5 Ma (Gradstein et al., 2004), fructicosa. This is not possible in the Brazos River area because P.
elegans is very rare. In addition, P. elegans first appears in zone CF6 in

deep-sea sections, including DSDP Sites 463 and 525A and in Tunisia
(Li and Keller 1998a, 1998b; Abramovich and Keller, 2003). An
alternative marker for Zone CF4 is an acme of Guembelitria cretacea
(Fig. 11). This disaster opportunist is best known as the dominant
species in the aftermath of the KTB mass extinction, when it thrived in
the absence of most other species (Keller and Pardo, 2004; Pardo and
Keller, 2008). However, G. cretacea thrived in any high-stress
environments, particularly in shallow-water sequences and during the
late Maastrichtian. Two intervals of high Guembelitria abundances are
commonly recognized, one in Zone CF1 and the other in Zone CF5 and
the lower part of CF4 (e.g., Keller et al., 1998; Keller et al., 2007b;
t
Keller, 2002, 2003; Abramovich et al., 1998; Abramovich et al., 2010;
Abramovich and Keller, 2002).
in
In Mullinax-1 and KT3 wells the lower Guembelitria acme appears
to span Zone CF4, a time of relatively warm climate (Fig. 11) (Haq et
al., 1987; Li and Keller, 1999; Adatte et al., 2002). A global cooling
occurs at the transition from the relatively warm climate of CF4 to CF3.
In Brazos sections, this interval is marked by sandstone at the base of
Pr
well Mullinax-1, which suggests a sea-level fall, followed by sandy
silty mudstone with high abundance of Guembelitria and the lowest FIGURE 8.—Age–depth plots for wells Mullinax-1 and KT3, showing
species richness (10–20 species) of the upper Maastrichtian (Fig. 7). In sediment accumulation rates for biozones CF1 to CF4. The KTB is
well KT3 Zone CF4 is also marked by sandy silty mudstone and set at 65.5 Ma; chronology after Gradstein et al. (2004).
abundant Guembelitria, though species richness is higher (20–30
species), probably due to the larger sample size available. The index
species R. fructicosa was observed near the base of this interval (29.5 The differences in d18O values between well Mullinax-1 and Hole
E-
m; Fig. 10), suggesting that the Guembelitria acme is at or near the
base of Zone CF4. Here we tentatively assume that the Guembelitria
acme approximates the base of CF 4, which is also suggested by the
sandstone deposit that marks a sea-level fall in well Mullinax-1. Based
525A can be partially explained by the fact that ocean-bottom
temperatures were considerably cooler than bottom temperatures in the
shallow neritic environment of the Brazos River area. However, other
factors that involved depletion of the isotopic values of foraminifera at
on this placement, sediment accumulation rates for CF4 range from Mullinax-1 must be taken into consideration.
0.82 m/ky (KT3) to 1.35 m/ky (Mullinax-1) (Fig. 8). A cross plot of the d18O and d13C values of the benthic
foraminifera in well Mullinax-1 shows a significant positive
STABLE ISOTOPES correlation at the 99.95% level (r ¼ 0.78; Fig. 5). Such a high
positive correlation is usually attributed to late diagenetic alteration
or
Oxygen and carbon isotopes of benthic and planktic foraminifera of meteoric groundwater that is characterized by both low d18O and
were analyzed to document changes in paleoproductivity and d13C values (Faure, 1986). However, recrystallization and cementa-
paleotemperatures in the upper Maastrichtian of well Mullinax-1 tion caused by meteoric groundwater diagenesis is not evident in the
(Fig. 3). Benthic foraminifera (mixed species) were analyzed from the foraminiferal tests from Mullinax-1 which mostly exhibit excellent
interval between 19.0 to 5.43 meters which corresponds to the upper or pristine preservation.
part of the Maastrichtian zone CF3 to the early Danian zone P1b. Stable The covariance between d18O and d13C and the low isotopic values
th
isotope data of P. costulata, the index taxon for upper surface waters, is can be alternatively explained by lower salinity due to fresh-water
limited to the interval between 14.56 to 11.69 meters, which includes influx (Khim and Park, 2000) The oxygen isotope composition of
most of zone CF2, where sufficient specimens were found for the stable seawater is linked to salinity either as a result of mixing between water
isotope analyses. masses with different salinities and isotopic compositions, or changes
Throughout this interval, benthic d18O values are distinctly higher in the precipitation–evaporation cycle (Wolff et al., 1999; Pierre, 1999;
than those of P. costulata, suggesting the existence of a thermal Rohling and de Rijk, 1999). A correlation between d18O and d13C of
Au
gradient between surface and bottom waters in the shelf environment of foraminifera is to be expected only if salinity changes are due to influx
the Brazos River area. Similarly, benthic d13C are mostly ; 0.5ø of a fresh water source, which is lighter in both d18O and d13C.
higher than those of P. costulata, indicating a surface-to-bottom During the late Maastrichtian in the Brazos River area, a low-
gradient in paleoproductivity. elevation rainfall source of the river water was probably not
Both d18O and d13C values of foraminifera at well Mullinax-1 are sufficiently light in d18O to produce a significant reduction of the
significantly lower than expected for the Maastrichtian and suggest a seawater values. An additional rainfall source of the fresh water
regional signal, as revealed by the comparison with coeval isotopic could have originated from more distant sources. Dettman and
records from oceanic and other continental-shelf localities. For Lohmann (2000) suggested that permanent snowfields existed at
example, benthic d18O values vary mostly between 4ø to 1.5ø. times in the late Maastrichtian and late Paleocene mountains of
In comparison, benthic d18O values at DSDP Hole 525A vary between north-central Wyoming and southwest Montana, and that melting of
0.25ø to 0.5ø in the cold interval between 67 to 65.9 Ma and reached this snow fed the rivers and significantly reduced the d18O values of
a minimum of 0.37ø during the latest Maastrichtian warm event at mollusk fossils. Similarly, Cochran et al. (2003) presented the
65.4 Ma (Li and Keller, 1998a). Similarly, benthic d13C at well possibility of fresh water influx into nearshore Western Interior
Mullinax-1 vary between 1.5ø to 0.4ø, whereas at Hole 525A they Seaway based on stable Sr and O isotopes. It is therefore reasonable
average around 1.5ø, and in the shallow shelf localities of Madagascar to consider snowmelt from distant highlands, for example to the
and Argentina they range between 1.5ø to 0ø (Abramovich et al., west of the Brazos River area as a possible source for the light d18O
2002; Keller et al., 2007b). and d13C values of the foraminifera in well Mullinax-1.
t
in
Pr
E-
or
FIGURE 9.—Species ranges of planktic foraminifera in well Mullinax-3 along with species richness (three-point average) of small species (63–150
lm) and larger species (. 150 lm). Note the decrease in species richness to 10 species is due to shallowing to coastal–lagoonal environment.
One interval with high species richness is due to reworking.
th
Periodic melting of ice caps could also explain the prominent freshwater runoff and provide further support that the negative d18O
negative peaks in both d18O and d13C records, which are particularly peaks represent discrete short-term warming pulses that may have been
frequent in intervals from the upper part of zone CF2 to the middle part part of the global late Maastrichtian warm event and associated sea-
of zone CF1 (; 65.7–65.4 Ma). d18O and d13C values decreased level changes (e.g., parasequences; Fig. 3).
Au
sharply at these episodes by as much as 2ø, relative to background

values (Fig. 3). This time interval marks the globally recognized latest FAUNAL ASSEMBLAGES
Maastrichtian warm events (Li and Keller, 1998c; Premoli Silva and
Sliter, 1999; Olsson et al., 2001; Nordt et al., 2003). The d18O negative Species census data and species ranges are the necessary tools for
peaks could therefore represent a periodic pattern of the warm event biostratigraphy and relative age dating along with unique acme events
that may have consisted of several warm peaks as recently suggested by of species. Paleoenvironmental information is largely obtained from
Abramovich et al. (2010). The negative d18O peaks could in part reflect the total species assemblages and, in particular, the relative abundances
an increase in seawater temperature but also an increase in fresh-water of individual species as obtained from quantitative analysis based on
runoff from a high altitude source. Most of the isotopic peaks coincide representative sample splits. This dataset combined with observations
with peak abundances of Guembelitria (Fig. 3). This correlation of the geographic and latitudinal species distributions, affinity to low-
provides strong evidence that the negative isotopic peaks in the latest oxygen and/or high-productivity environments, and species depth
Maastrichtian at Brazos were induced by environmental–climatic habitats based on stable isotope ranking all provide insight into the
changes that involved shifts in the isotopic composition of the seawater. paleoecology and depositional environment of a particular region. For
The isotopic peaks of foraminifera at well Mullinax-1 are also the Brazos area, relative species abundances of the late Maastrichtian
marked by distinct color changes from dark gray to light gray shales were analyzed for each of the three wells (Mullinax-1, Mullinax-3,
(Fig. 3) that imply transitions from dysoxic to more oxic conditions at KT3) in an attempt to gain insights into the local depositional and
the sea floor. Such transitions are expected during times of increased paleoenvironmental conditions.
t
in
Pr
E-
or
FIGURE 10.—Species ranges of planktic foraminifera in well KT3 along with species richness (three-point average) of small species (63–150 lm)
and larger species (. 150 lm). Note that the more consistent presence of rare species is probably due to larger samples analyzed.
th
Ecological Opportunists Persistent and Sporadic Taxa

Late Maastrichtian faunal assemblages are very similar in the three Persistent and sporadic occurrences of taxa vary between Mullinax-1
Au
wells of the Brazos area, which span a distance of approximately 3.5 and KT3. It is unclear whether this is due to local environmental
km (Fig. 1). Species patterns show four groups characterized by conditions or to the smaller sample size analyzed in Mullinax-1. It may
dominant (up to ; 80%), common (; 10–30%), persistent (1–10%), to primarily be the latter that accounts for the lower species richness and
sporadic and rare occurrences (, 1%, Figs. 13–15). In each well the fewer specimens. This is evident in the upper part of zone CF4, where
same three species dominate (H. globulosa, H. planata, G. cretacea; specimen counts of several samples had to be combined in order to obtain
Plates 1, 2), and the same five species are relatively common (; 10%) a statistically significant representation of the assemblages (see gray-
shaded intervals in Fig. 12). This is also evident by the extremely low
[Hedbergella spp., P. navarroensis, Globigerinelloides asper (Ehren-
number of foraminifera per gram of sediments through most of the section
berg), P. costulata, Pseudoguembelina costellifera Masters; Plates 1,
(this data are not available for KT3). High specimen numbers are confined
3]. For most of these species, variations in relative abundances are to the sandstone at the base and to some isolated samples below the
relatively minor with no long-term trends, except for G. cretacea, H. sandstone complex near the top of the Maastrichtian (Fig. 12). In both
globulosa and H. planata (Figs. 14–16). Globigerinelloides asper cases, these erratic peaks may simply reflect reworking and winnowing of
tends to be more common in Zone CF4 (Figs. 14, 15), whereas P. older sediments. Therefore, we assume that well KT3 is more
navarroensis is more abundant in Mullinax-1 than in the two localities representative of late Maastrichtian assemblages in the Brazos area.
to the south. All of these species can be considered ecological In well KT3, persistent and sporadic occurrences of taxa encompass
opportunists (r-strategists) that thrived in open marine as well as a number of small heterohelicids [e.g., Heterohelix striata (Ehrenberg),
shallow-water environments. Heterohelix labellosa Nederbragt, Fleisherites glabrans (Cushman);
t
in
Pr
E-
FIGURE 11.—Stratigraphic correlation of Guembelitria bloom events in the upper Maastrichtian of the eastern Tethys and the Western Interior
Seaway correlated with the climate record of South Atlantic DSDP Site 525A. Note that there are two major Guembelitria blooms in zones
CF4 and CF2-CF1.
or
Plate 3] , pseudoguembelinids (e.g., P. palpebra Brönnimann & Brown, marine waters into the shallow coastal to lagoonal environment of
P. hariaensis, P. kempensis Esker; Plate 4), pseudotextularids (P. Mullinax-2 and Mullinax-3.
elegans ¼ P. nutalli, P. elegans (¼ deformis); Plate 4), rugoglobigerinids The K-strategists in all three wells include all globotruncanids and
(e.g., R. rugosa Plummer, R. macrocephala Brönnimann, R. hexaca- the large multiserial taxa (racemiguembelinids, planoglobulinids; Plate
merata Brönnimann Kuglerina rotundata. Brönnimann, Trinitella 4), although they are represented by only a few specimens (Plates 7, 8)
scotti (Brönnimann); Plates 5, 6), and globigerinellids [G. volutus and are generally poorly developed. The rarity of this group and their
th
(White), G. yaucoensis (Pessagno)] (Fig. 13). In open marine underdeveloped nature reveals high-stress conditions related to the
environments of low to middle latitudes most of these species are shallow-water environment of the Brazos area (Keller and Abramo-
persistently present if not common. Their minor presence (, 15% of vich, 2009).
total assemblage) in the Brazos area most probably reflects adverse
conditions due to shallow water depth and high terrigenous influx ENVIRONMENTAL INDICES
(predominantly mudstone and claystone deposition).
Au
Guembelitria Blooms: Disaster Opportunists

Ecologic Specialists Guembelitrids are a group of small-sized (38–100 lm) triserial
planktic foraminiferal species that are sporadically known from the
All larger specialized (K-strategy) species (. 150 lm) are rare or
middle Cretaceous to the recent and thus form one of the longest-
absent in the Brazos sections, and they gradually disappeared beginning ranging plankton groups. Despite this long history, their stratigraphic
in zone CF2 (Figs. 12–14). An abundance plot of the . 150 lm size record is poorly documented, largely because routine foraminiferal
fraction in Mullinax-3 shows the gradual disappearance beginning below analysis concentrates on larger species. However, in recent years as
CF1 and near absence in the top 1 m below the KT boundary (Fig. 15). In more studies have concentrated on mass extinctions and biotic stress
this uppermost Maastrichtian interval only some sample intervals conditions the importance of small species as environmental indicators
contained any specimens . 150 lm and generally insufficient in has become increasingly recognized, and the dataset has grown
numbers (, 100 specimens) for a quantitative abundance assessment. For accordingly. Guembelitria is particularly common in shallow-water
this reason, they are plotted as present in Figure 15. Moreover, the species environments such as the Brazos area, where they vary in abundance
most common in the . 150 lm size fraction are the ones that are most and size range.
abundant in the 63–150 lm size fraction (e.g., H. globulosa, H. labellosa, The most frequent small species is G. cretacea, a well-known
P. costulata, R. rugosa). The sporadic occurrence of these species and low disaster opportunist that thrived in eutrophic surface waters, and their
specimen numbers suggests reworking and/or intermittent influx of blooms are best known from the aftermath of the KTB mass extinction
t
in
Pr
E-
FIGURE 12.—Relative species abundances of upper Maastrichtian planktic foraminifera (. 63 lm) in well Mullinax-1, along with the planktic/
benthic abundance and the number of foraminifera per gram of sediment. Note that only in the sandstone at the base is there an appreciably
or
high number of specimens per gram; in the rest of the sequence foraminifera are relatively rare. See Figure 8 for complete caption.
(review in Keller and Pardo, 2004). Abramovich et al. (1998) first show three short-term but strongly negative excursions of 2.5ø in
reported Guembelitria blooms in the upperr Maastrichtian of Israel, d18O and 1.5ø in d13C during the latest Maastrichtian maximum
and since then they have been documented in many localities and climate warming, beginning near the top of CF2 and continuing
th
across latitudes globally (Fig. 17; Pardo and Keller, 2008). Guembeli- through the lower half of CF1. These negative excursions are of almost
tria blooms indicate that conditions of high biotic stress occurred the same magnitude as at the KT boundary, and each coincides with a
intermittently, and they are widespread during zone CF1 and CF4 of lithologic change. A smaller 1ø negative excursion is present at the
the late Maastrichtian (e.g., Abramovich et al., 2010), but they have CF2-CF3 transition. (No stable isotope data are available for the
also been observed in CF3 (Pardo and Keller, 2008). These blooms are Guembelitria bloom in zone CF4). Guembelitria blooms (50–60%
Au
thus useful biostratigraphic markers, especially in shallow-water abundance) coincide with the KT boundary and the two zone CF1
environments, such as the Brazos River sections in Texas. excursions but drop to lower values after the excursions. The
Association of Guembelitria with mass extinction is considered Guembelitria blooms, lithological changes and the coincidence with
evidence for the opportunistic nature of this species which proliferated CF2-CF1 maximum global climate warming are real events that may
during times of enhanced food resources and absence of normal represent discrete episodes of freshwater runoff that were induced by
consumers (Keller, 1996; Keller and Pardo, 2004; Pardo and Keller, short-term warming pulses.
2008). In the Maastrichtian, Guembelitria bloomed in the uppermost
surface water primarily above shelf and slope environments. These Dominance of Heterohelix: Common generalists
blooms were found to coincide with global warming and suggest
similar opportunistic behavior (Abramovich et al., 2010). Unlike the The most typical example of generalist strategy among the
KTB, the Maastrichtian blooms appear to have failed to reach the open Maastrichtian planktic foraminifera are the biserial heterohelicids,
ocean. Moreover, the presence of other species during these bloom namely H. globulosa and H. planata (Leckie, 1987; Keller, 1989a,
events indicates that global warming did not involve significant 1996, this volume; Nederbragt and Koning, 1994; Abramovich et al.,
elimination of species habitats, unlike the KTB catastrophe. 1998; Abramovich et al., 2002, Pardo and Keller, 2008). Species of this
In well Mullinax-1, the Guembelitria blooms in Zone CF1-CF2 genus typically display great variation in test size and chamber shape,
show an erratic pattern that correlates with a similar pattern in d13C and which implies the ability to adjust to a wide range of environmental
d18O isotopes and the benthic/planktic ratio (Fig. 3). Stable isotopes conditions. Highest abundance of Heterohelix (. 50%) is found in
t
in
Pr
E-
or
FIGURE 13.—Relative species abundances of upper Maastrichtian planktic foraminifera (. 63 lm) in well KT3, along with stable isotope data.
shelf–slope environments such as Spain (Pardo et al., 1996), El Kef and assemblages is an important global marker of OAE2 (Leckie et al.,
Elles, Tunisia (Keller, 1988; Li and Keller, 1998b; Abramovich and l998; Luciani and Cobianchi, l999; Nederbragt et al., 1998; Nederbragt
th
Keller, 2002), and Negev, Israel (Abramovich et al., 1998; Abramovich and Fiorentino, l999; Keller et al., 2001; Keller and Pardo, 2004;
et al., 2010). In deep-sea localities, particularly in middle latitudes, Coccioni and Luciani, 2004; 2005). Frank et al. (2005) reported a
populations of biserial species are often smaller than those of shelf similar response of heterohelicids during the early Maastrichtian
environments (; 30%), yet considerably larger than those of highly warming (; 70–68 Ma) in the tropical Pacific at Shatsky Rise. This
specialized species in the associated assemblage (Abramovich and event was manifested by the brief appearance of abundant inoceramid
Keller, 2003; Frank et al., 2005). The high abundance of these species bivalves over shallow portions of Shatsky Rise, and coincided with a
Au
in different environments and the great variation in their test distinct increase in heterohelicids, which suggested higher productivity
morphology suggest the ability to adjust to rapid environment changes. at the time of the warming. In contrast, in the late Cretaceous upwelling
However, in contrast to Guembelitria species, those of Heterohelix system of the southern Tethys Almogi-Labin et al. (1993) found that
were generalists and not opportunists, as suggested by their continuous very high TOC levels (. 10% wt) in association with dominant
high occurrences in stable marine environments. Globigerinelloides, whereas dominance of heterohelicids was found in
Small heterohelicids are also often considered to be tolerant of low- intervals with lower TOC levels and higher-diversity assemblages that
oxygen conditions, thriving within the oxygen-minimum zone (e.g., include keeled globotruncanids.
Kroon and Nederbragt, 1990; Barrera and Keller, 1990; Leckie, 1987; Pardo and Keller (2008) summarized late Maastrichtian Heterohelix
Leckie et al., l998; Nederbragt et al., l998; Premoli Silva and Sliter, and Guembelitria blooms from nutrient-rich to eutrophic environments
1999; Keller and Pardo, 2004). A relative abundance increase in these from the Tethys to high latitudes and related these to volcanism and
species was interpreted as indicating an expanded oxygen-minimum high terrestrial runoff, particularly in Argentina (Keller et al., 2007b)
zone due to increased surface productivity and the depletion of oxygen and Indian Ocean Ninetyeast Ridge DSDP Site 216 (Keller, 2003,
in subsurface waters due to oxidation of organic carbon (Hart and Ball, 2005). In all of these environments Heterohelix globulosa and other
l986). Heterohelicids appear to have evolved to take advantage of low- small biserial species (e.g., P. navarroensis, H. dentata, (¼ H. planata
oxygen conditions, as evident in the oceanic anoxic event (OAE2) of in this study; heterohelicids with costate ornamentation over most of
the Cenomanian–Turonian transition, where this group thrived to the the chamber surface) invariably are the first species to thrive after
exclusion of most other species, and the shift to Heterohelix dominated environmental conditions improved and Guembelitria blooms de-
t
in
Pr
E-
FIGURE 14.—Relative species abundances of Upper Maastrichtian planktic foraminifera (. 63 lm) in well Mullinax-3, along with species richness
per sample and three-point average.
or
creased in intensity. This is well illustrated by the alternating KT3, and in the equivalent interval below the KTB unconformity at
Guembelitria and Heterohelix blooms at Site 216 in association with Mullinax-3 (Figs. 12–15). This may also account for the relatively high
volcanic inputs during the late Maastrichtian (Fig. 16). The same but erratic peaks in the number of specimens per gram of sediment in
alternation is observed in the Brazos sections. the sandstone at the base of well Mullinax-1.
At Mullinax-1, where H. globulosa and H. planata have been Plots of species richness for the Brazos sections are shown as three-
th
plotted together, the alternation of Guembelitria and Heterohelix peaks point sample averages, to smooth out closely spaced sample variations
is particularly pronounced (Fig. 12). But the same pattern is evident in and reveal long-term trends. Significant variations are evident between
KT3 and Mullinax-3, although H. planata and H. gobulosa are plotted the three localities (Fig. 17). Well KT3 shows an average total of 35
separately (Figs. 13, 14). In all three Brazos sections Guembelitria and species, with a maximum of 42 and a low of 27 species for the late
Heterohelix peaks frequently exceed 80% during the late Maastrich- Maastrichtian. Larger species (. 150 lm) are plotted separately to
tian, which indicates persistent high-stress conditions, which are show their relative proportion; they average between 12 and 15 species
Au
typical of shallow marine environments with possibly high nutrient with only minor variations. The total species richness shows regular
input as indicated by the dark claystones, mudstones, and burrows variations that may reflect environmental changes. However, this
infilled with pyrite framboids. These conditions are superimposed by cannot be determined because only a generalized sediment description
the global warming episodes during the CF4 and CF2–1 intervals, is available for these old cores, which permits no evaluation of the
which are indicated by the bloom events of Guembelitria. changes in the sedimentary environment.
The species richness pattern is quite different in Mullinax-1,
Species Richness—Environmental Stress Index although the average number of species is approximately the same,
with a minimum of 15 species and a maximum of 39 species (Fig. 17).
Species richness is the most easily identified proxy for conditions of Larger species (. 150 lm) vary from 0 to 15. This extreme variation is
high biotic stress. It assumes that preservation is good and that species due to an interval of low species richness in upper zone CF4 to lower
are not lost to breakage and dissolution. In the Brazos area, zone CF3 that is characterized by sandy mudstone with very rare
preservation is mostly excellent, as evident by the species illustrations planktic foraminifera. In contrast, the lower interval, consisting of fine
(Plates 1–8), and there is no evidence of preservational bias. sandstone, contains relatively common specimens and a high of 33
Nevertheless, there appears to be another type of bias, an influx of species, including 8–12 larger species (. 150 lm). It is unclear why
reworked specimens that enhances primarily the number of large thick- the species richness and number of species per gram of sediment are
shelled species. This is evident in all three sections below the higher in the sandstone than in the sandy mudstone. One possibility is
unconformity at the base of a sandstone complex in Mullinax-1 and an influx of reworked specimens along with the sand. Most probably
t
in
Pr
E-
FIGURE 15.—Relative species abundances of upper Maastrichtian planktic foraminifera (. 150 lm) in well Mullinax-3, along with species
richness per sample and three-point average.
or
this difference is due to local topographic variations and associated
sediment drift. Only zones CF1 to upper CF3 show species richness
similar to that in KT3 (Fig. 17).
Mullinax-3 shows a maximum of 30 species and a minimum of 18
species for the upper Maastrichtian zones CF2 to the lower half of CF1,
with larger species (. 150 lm) varying from 2 to 10 (Fig. 15). Most
th
significant in this section of the . 63 lm size fraction is the

disappearance of larger species at an unconformity in the middle of
zone CF1 and the decrease to just 11 smaller species, including the
group of eight common and dominant species in the Brazos area. Even
in the counts of the . 150 lm size fraction, specimens are rare in the
top meter (Fig. 15) which reveals the high-stress conditions. This
Au
interval consists of a sandy, glauconitic mudstone with clasts, shell

fragments, oysters, desiccation cracks, and rootlets that suggest a
coastal environment, possibly of mangroves. The same lithology
continues for 1.0 m, followed by a 25-cm-thick glauconitic sandstone.
In the upper 50 cm of this interval is a temporary influx of diverse
species that are not present below and above (Figs. 14, 15, 17). These
species probably mark the occasional influx of marine waters
(tempestites?) carrying more diverse planktic foraminiferal assem-
blages into this coastal area. The uppermost part below the KTB
FIGURE 16.—Guembelitria and Heterohelix blooms during the late

Maastrichtian at the Indian Ocean Ninetyeast Ridge DSDP Site 216
are related to mantle-plume volcanism (Modified after Keller, 2003,
2005).
t
in
Pr
E-
or
FIGURE 17.—Species richness of planktic foraminifera in wells Mullinax-1, Mullinax-3, and KT3 based on histograms of three-point sample
average of small species (63–150 lm) and larger species (. 150 lm). Gray shading marks the latest Maastrichtian zone CF1. Data on species
richness show shallowing from north to south, with coastal–lagoonal conditions in the south during the upper part of CF1.
th
unconformity returns to the minimum species richness characteristic of approximately one-third (e.g., Madagascar; Abramovich et al., 2002).
the lower part of the sandy glauconitic interval. Further species reduction is observed in mesotrophic or restricted
Variations in species richness and lithology of the three Brazos wells basins, such as in Israel and Egypt, where species richness varies
indicate significant shallowing from north to south over a distance of between 25 and 35 species, regardless of water depth (Keller, 2002,
approximately 3 km, with the southernmost well Mullinax-3, located in 2004). In very shallow inner-neritic environments species richness
Au
a coastal area marked by temporary emergence in the upper part of varies between 20 and 35 and in coastal to lagoonal settings decreases
zone CF1. This emergence is coeval with deposition of the spherule- to approximately 10 species (e.g., Mullinax-3 in CF1; Fig. 18). A
rich and shell-rich glauconitic sandstone that infills incised valleys in similar low diversity is observed at Seldja at the edge of the Sahara,
wells KT3 and Mullinax-1, as well as the Darting Minnow Creek (e.g., Tunisia; Keller et al., l998). Lowest species richness was
outcrop, only approximately 150 m away (Fig. 1). The overall low observed in areas of volcanic activity, such as DSDP Site 216 and in the
species diversity of the late Maastrichtian (25–40 species) is Neuquén Basin of Argentina (Keller, 2003; Keller et al., 2007b). In
characteristic of middle-shelf to inner-shelf conditions. Based on the both regions species richness varies between 6 and 12 species, but it
faunal assemblages and lithology we estimate deposition during the increased to approximately 20 in Site 216 when volcanic activity
late Maastrichtian to have occurred at a depth much less than 100 m ceased (Figs. 16, 18).
and approaching 0 to 30 m at the time of the sea-level fall and The progressively decreasing trend in species richness towards
deposition of the sandstone complex with impact spherules in zone shallower and restricted environments thus marks increasing biotic
CF1 in Mullinax-1 and KT3, but emersion in Mullinax-3 to the south. stress. In such environments, volcanic activity or volcanic influx from
Diversity reduction is the most commonly noticed response to weathering (e.g., Neuquén Basin) results in extreme biotic stress and
environmental stress conditions. In continental-shelf settings, highest the survival of only the most resistant opportunistic–generalist species.
species richness (; 65 species) occurs in deeper outer-shelf to upper- Across diverse continental-shelf environments, the reduction in species
slope or open marine environments (Fig. 21). In shallower, middle- richness seems to follow a uniform pattern of morphologic selectivity,
shelf conditions (; 100–200 m), species richness is reduced by eliminating large species first and smaller species surviving. The
t
in
Pr
E-
FIGURE 18.—Species richness decreases with increasing biotic stress conditions that parallel environments from outer shelf to inner shelf.
Maximum biotic stress is associated with very shallow inner-neritic to coastal and lagoonal environments with high nutrient influx leading to
eutropic and dysoxic conditions. In these high-stress environments, some or all species may be dwarfed. Mullinax-3 shows high biotic stress
similar to environments stressed by volcanic activity in the Indian Ocean and Argentina. (Modified after Keller and Abramovich, 2009).
or
selective nature of the diversity reduction can be illustrated by The depth distribution deduced from Brazos d18O records of
examining the r–k continuum and depth-ranked species in optimum, individual planktic species is generally in agreement with the oceanic
high-stress, and disaster assemblages. records (Fig. 20). Seven of the eight species listed as surface mixed
layer in the oceans are rare, persistently present, or common in the
Species Depth Ranking Brazos area. Also present in the surface mixed layer are Guembelitria
species. These groups have relatively low d18O values, but most
th
Stable isotope depth ranking of planktic foraminiferal species distinctly ranked very low in d13C values (0.5–0ø) that overlap the
provides insights into the nature of species assemblages, prevailing benthic records (Rak et al., personal communication; Fig. 20). This
watermass characteristics, and probably paleodepths in shelf areas. suggests that the episodic blooms of this species occurred in the
d18O values of foraminiferal species generally increase with depth due uppermost water-layer where photosynthesis is inhibited by high UV
to decreasing temperatures. In contrast, d13C values decrease with and nutrients are nearly absent (see further discussion in Pardo et al.,
depth due to selective removal of 12C by photosynthesis in the photic 2008; Abramovich et al., 2010). Pseudoguembelina costulata and
Au
zone and accumulation of 12C in deeper water due to organic matter Paraspiroplecta navarroensis were also found to inhabit the surface
decay (e.g., Fairbanks et al., 1982; Abramovich et al., 2003). Surface water judging by their relatively low d18O values. Planoheterohelix cf.
dwelling planktic foraminifera are therefore expected to display the planata, a species with lighter striations than P. costulata, is relatively
lightest d18O and heaviest d13C values, whereas deep dwellers display common in Brazos sections and was probably also a surface dweller.
heaviest d18O and lightest d13C values. The largest species group ranks slightly deeper in the mixed layer, or
Stable isotope depth ranking of late Maastrichtian species from the subsurface layer. All but two of the 17 species isotopically ranked
equatorial sites (DSDP Sites 577A, 463) and middle-latitude Atlantic by Abramovich et al. (2003) are present in the Brazos sections (Table
(DSDP Site 525A) by Abramovich et al. (2003) is shown in Table 1. 1). Most of these species are rare and/or sporadically present at Brazos.
This dataset is integrated with new stable isotope data of individual The only exception is the dominant species H. globulosa, and common
species analyzed from three selected intervals in wells Mullinax-1 and species R. rugosa and G. asper. The latter typically display slightly
Mullinax-3 by Carmi et al., (personal communication; Fig. 20). higher d18O and d13C values (2.5ø and 1.2–1.6ø, respectively)
Differences between the d18O values of benthic and planktic which imply that it inhabited somewhat cooler water with higher
foraminifera from Brazos vary between 0.5 and 1 ø, which indicates primary productivity (Fig. 20). There is an additional group of 14
that the water column in this environment was thermally stratified but species at Brazos that are also probably subsurface dwellers. Most of
probably not sufficiently deep to support intermediate-depth and deep- these are small heterohelicids, rugoglobigerinids, globigerinellids, and
water species (i.e., keeled globotruncanids) which are extremely rare hedbergellids. Among these only H. planata and Hedbergella species
and, if present, have dwarfed or juvenile morphologies. are common to abundant.
t
in
Pr
E-
FIGURE 19.—Patterns of test-size variations in three planktic and one benthic species during the late Maastrichtian to early Danian at well KT3
(data from MacLeod et al., 2000). Note that test-size reduction begins below the sandstone complex and coincides with a major sea-level fall
that marks a sequence boundary. (Modified after Keller and Abramovich, 2009).
Most globotruncanids and racemiguembelinids are listed as living at tria, Pseudoguembelina costulata, Planoheterohelix cf. planata,
or
thermocline depths. All but two of the 12 species listed are very rarely Paraspiroplecta navarroensis) and subsurface (H. globulosa, H.
found in the Brazos sections. In addition to these, rare Globotruncana planata, R. rugosa, G. asper) of the mixed layer (Table 1). Almost
duwi, G. esnehensis, and G. insignis are recorded at Brazos (Table 1).
None of the deep dwellers are present. all other species are rare and sporadically present. This indicates a
Stable isotope depth ranking of species reveals that at Brazos the high-stress shallow-water environment, as also indicated by species
bulk of species populations (; 90%) lived in the surface (Guembeli- richness, lithology, and macrofossils.
th
Au
FIGURE 20.—The main features of the planktic foraminiferal assemblages of the Brazos are domination of small, simple morphotypes
opportunists, generalists, and high-stress specialist morphologies.
TABLE 1.—Isotopic depth ranking of late Maastrichtian species based on Abramovich et al. (2003) and Rak et al. (personal communication).
Late Maastrichtian Stable Isotope Depth Ranking of Planktic Foraminifera
mixed layer Thermocine Subthermocline
Surface Subsurface intermediate Deep
Pseudoguembelina excolata Globotruncana aegyptiaca Globotruncana arca Planoglobulina multicamerata

P. kempensis G. mariei G. dupeublei Heterophelix rajagopalani
P. costulata G. rosetta G. falsostuarti Abathomphalus mayaroensis
P. hariaensis Rosita walfishensis G. mariei Gublerina cuvilieri
t
P. palpebra Rugogloberina rugosa G. orientalis Gansserina gansseri
Rugoglobigerina pennyi** R. hexacamerata G. ventricosa Lipsonia lipsonae
in
R. hexacamerata R. milamensis Globotruncanita angulata/pettersi
Planuglobulina acervulinoides R. rotundata G. stuarti
** also listed in thermocline Globigerinelloides asper* Rosita patelliformis
* also listed in subsurface G. subcarinatus Racemiguembelina fructicosa
Gublerina acuta R. intermedia
Pr
Pseudotextularia deformis R. powelli
P. elegans Rugoglobigerina pennyi
Heterohelix globulosa* Globotruncanella havananensis
H. labellosa
Planoglobulina carseyae
P. brazoensis E-
* also listed in thermocline
Species also present in Brazos area (this study)

Paraspiroplecta navarroensis Globotruncana duwi Globotruncanella petaloidea
Guembelitria cretacea G. esnehensis Globotruncanella havanensis
Guembelitria irregularis G. insignis
Guembelitria trifolia Rugoglobigerina macrocephala
Guembelitria danica R. scotti
or
Paraspiroplecta navarroensis R. reicheli
Plummerita hantkeninoides
Heterohelix planate
H. dentate
H. punctulata
Globigerinelloides multispina
th
G. rosebudensis
G. volutes
G. yaucoensis
Hedbergella monmouthensis
H. holmdelensis
Au
DISCUSSION diversity and morphologically smaller species (, 150 lm). In these

assemblages, relative species abundances of surviving larger species
At times of high environmental stress, species diversity is reduced are very low and may be present only sporadically. In contrast, diversity
across latitudes. The size of the diversity decrease depends on the of morphologically small species tends to remain steady and their
severity of biotic stress, though the largest reduction occurs in low to relative abundances tend to increase to 60–80% as documented in
middle latitudes, shallow shelf areas, and restricted basins. A diversity Madagascar, Israel, Egypt, Tunisia, and Denmark (Abramovich and
reduction of 30% to 50% is not uncommon. Species reduction is Keller, 2002; Abramovich et al., 2002; Keller, 2002, 2004; Keller et al.,
selective, eliminating the largest and most specialized species first 1993; Keller et al., 1998; Hart et al., 2005).
(e.g., Racemiguembelina, Contusotruncana, Planoglobulina, most During the late Maastrichtian, the foraminiferal assemblages of the
keeled globotruncanids). These taxa are predominantly intermediate Brazos area were dominated by small, simple morphotypes, oppor-
dwellers (e.g., subsurface and thermocline; Table 1) (Abramovich et tunists, generalists, and high-stress specialist morphologies. Most
al., 2003). The resulting survivor assemblages consist of lower- abundant species at Brazos are small heterohelicid species (H.
globulosa, H. planata, P. navarroensis) and Guembelitria species (Fig. cids species (H. globulosa, H. planata, P. navarroensis). These
20). Other small surface and subsurface dwellers are rare to common assemblages account for approximately 80% and reveal high biotic
(e.g., Hedbergella, Globigerinelloides, P. costulata, Planoheterohelix stress through the late Maastrichtian due to high nutrient influx and
cf. planata). Larger specialized species (e.g., globotruncanids, dysoxic subsurface waters. All specimens are smaller than 150 lm.
racemiguembelinids, planoglobulinids, pseudoguembelinids, pseud-
The remaining faunal components include common surface and
textularids) are rare and sporadically present (Figs. 12–15). Their
combined abundance is less than 2% of the total planktic foraminiferal subsurface dwellers (hedbergellids, globigerinellids, heterohelicids,
assemblages. pseudotextularids, and rugoglobigerinids) which account for ap-
MacLeod et al. (2000) studied the intraspecies variations in three proximately 15% of the total assemblages and are generally dwarfed,
planktic and one benthic species from the Brazos well KT3 and with specimens smaller than 150 lm.
demonstrated that ‘‘all four species experienced statistically significant Larger (. 150 lm) specialized species are very rare and sporadically
test size decreases in the latest Maastrichtian’’ and continuing into the present, including globotruncanids, most rugoglobigerinids, pseud-
t
earliest Danian (Fig. 19). The maximum mean test size in Heterohelix textularids, pseudoguembelinids, planoglobulinids, and racemi-
globulosa and P. navarroensis was reached ; 50 cm below the KTB or guembelinids. Specimens present are generally dwarfed attesting
in
approximately 10 cm below the unconformity at the base of the to high-stress conditions. Most of these species lived in deeper
sandstone complex. A similar pattern is observed in the benthic waters at or below the thermocline or required photosymbionts.
foraminifer Anomalinoides newmanae. However, G. cretacea shows Their rarity, dwarfed size, and frequent absence reflects high biotic
maximum size just below the KTB coincident with minimum test size stress due to the shallow depths, high nutrients, and muddy (green)
in heterohelicid species. Similar patterns of test size are observed at waters.
Pr
Nye Klov, Denmark, and ODP Site 738 (Kerguelen Plateau, southern
Indian Ocean). In all three localities the test size changes parallel a The depositional environment fluctuated between inner-neritic and
strong decrease in species richness and reflect severe biotic stress middle-neritic in the Brazos River area, but frequent disconformities
(Keller et al., 1993; MacLeod et al., 2000; Hart et al., 2005; Keller and and erosional surfaces with abundant shell fragments, sand layers,
Abramovich, 2009). snails, and rare planktic but abundant benthic foraminifera suggest
The most probable causes for the biotic stress in the Brazos area temporary emergence, particularly near the top of zone CF2 and in
include the shallow shelf setting superimposed by global climate CF1. Emergence at the time of deposition of the sandstone complex
is indicated in well Mullinax-3 by a karst surface, rizoliths, clasts,
E-
changes and sea-level fluctuations. During the late Maastrichtian zones
CF4 to CF2, sediment deposition fluctuated between middle-neritic
and inner-neritic environments. The maximum climate warming at the
end of CF2 and in CF1 was accompanied by sea-level fluctuations that
and mudcracks.
Stable isotope data show a short-term negative excursion associated
with disconformities and lower sea levels near the top of CF2 and in
led to coastal–lagoonal environments punctuated by temporary CF1. These negative excursions could in part reflect an increase in
emergence marked by high benthic abundance (. 60%), decreased seawater temperature but also an increase in fresh-water runoff.
planktics, karstified erosion surfaces, roots, and mudcracks (Gale,
2006; Keller et al., 2007a; Adatte et al., this volume). The return to
cooler climate in the upper part of zone CF1 was accompanied by a ACKNOWLEDGMENTS
or
sea-level fall leading to major erosion and the formation of incised
valleys, which were subsequently infilled with clasts, glauconitic, We are very grateful to Jerry Baum, who directed the drilling effort,
shell- and spherule-rich sand eroded from exposed areas nearshore. and Tom Yancey for splitting and describing the cores. A special thank
The biotic stress caused by the climate warming and shallowing you to the owners of the Brazos Rose Ranch, Mr. and Mrs. Ronnie and
environment resulted in the exclusion of deeper dwelling species, Jackie Mullinax, who not only permitted drilling on their land but also
dwarfing, and assemblages dominated by alternating abundances of took intense interest in the geology and so graciously hosted our many
Guembelitria and Heterohelix blooms. visits. We gratefully acknowledge the drilling crew of DOSECC and
th
logging support from Schlumberger during two drilling phases. We

CONCLUSIONS greatly thank reviewers Marius Dan Georgescu and Emma Sheldon for
their contribution towards improvement of this paper We also thank
Upper Maastrichtian sediments recovered from three subsurface Marius Dan Georgescu for his comments on the taxonomical
wells along the Brazos River, Cottonmouth Creek, and Darting identifications. The material of this study is based upon work
Minnow Creek span Cretaceous foraminiferal zones CF1-CF4. The supported by the US National Science Foundation through the
Au
KTB is 80 cm above a sandstone complex with reworked impact Continental Dynamics Program and Sedimentary Geology Program
spherules. under NSF Grants EAR-0207407 and EAR-0447171, and BSF grant
2004045.
At the base of CF4, sediments consist of sandstone followed by
sandy mudstone with common sand laminae into the lower part of REFERENCES
CF3 and grading into silty mudstone through CF3 and mudstones–
claystones in CF2 through CF1, except for the sandstone complex in ABRAMOVICH, S., AND KELLER, G., 2002, High stress late Maastrichtian
CF1. Paleoenvironment in Tunisia: Inference from planktic foraminifera: Palae-
ogeography, Palaeoclimatology, Palaeoecology, v. 178, p. 145–164.
The Chicxulub impact spherules in the sandstone complex in zone ABRAMOVICH, S., AND KELLER, G., 2003, Planktonic foraminiferal response to the
CF1 are reworked, as evident by lithified clasts with spherules and latest Maastrichtian abrupt Warm Event: A case study from South Atlantic
mudcracks. DSDP Site 525A: Marine Micropaleontology, v. 48, p. 225–249.
ABRAMOVICH, S., ALMOGI-LABIN, A., AND BENJAMINI, C., 1998, Decline of the
Planktic foraminiferal assemblages of the Maastrichtian at Brazos
Maastrichtian pelagic ecosystem based on planktic foraminifera assemblage
are of low diversity and dominated by alternating high abundances of changes: Implication for the terminal Cretaceous faunal Crisis: Geology, v.
the disaster opportunist G. cretacea (zones CF4 and CF2-CF1) 26, p. 63–66.
which thrived in uppermost water during periods of global warming ABRAMOVICH, S., KELLER, G., ADATTE, T., STINNESBECK, W., HOTTINGER, L.,
and the subsurface-dwelling, low-oxygen tolerant small heteroheli- STUEBEN, D., BERNER, Z., RAMANIVOSOA, B., AND RANDRIAMANANTENASOA, A.,
2002, Age and paleoenvironment of the Maastrichtian to Paleocene of the GEORGESCU, M.D., 2009, Taxonomic revision and evolutionary classification of
Mahajanga Basin, Madagascar: A multidisciplinary approach: Marine the biserial Cretaceous planktic foraminiferal genus Laeviheterohelix
Micropaleontology, v. 47, p. 17–70. Nederbragt, 1991: Revista Mexicana de Ciencias Geológicas, v. 26, p.
ABRAMOVICH, S., KELLER, G., STÜBEN, D., AND BERNER, Z., 2003, Characteriza- 315–334.
tion of late Campanian and Maastrichtian planktonic foraminiferal depth GEORGESCU, M.D., AND ABRAMOVICH, S., 2008, Taxonomic revision and
habitats and vital activities based on stable isotopes: Palaeogeography, phylogenetic classification of the Late Cretaceous (upper Santonian–
Palaeoclimatology, Palaeoecology, v. 202, p. 1–29. Maastrichtian) serial planktic foraminifera (Family Heterohelicidae Cushman
ABRAMOVICH, S., YOVEL-COREM, S. ALMOGI-LABIN, A., AND BENJAMINI C., 2010, 1927) with peripheral test wall flexure: Revista Española de Micro-
Global climate change and planktic foraminiferal response in the Maas- paleontologia, v. 40, p. 97–114.
trichtian: Paleoceanography, v. 25, PA2201. GEORGESCU, M.D., AND ABRAMOVICH, S., 2009, A new Late Cretaceous
ADATTE, T., KELLER, G., AND STINNESBECK, W., 2002, Late Cretaceous to early (Maastrichtian) serial planktic foraminifer (Family Heterohelicidae) with
Paleocene climate and sea-level fluctuations: Palaeogeography, Palaeoclima- early planispiral coil and revision of Spiroplecta Ehrenberg, 1844: Geobios,
tology, Palaeoecology, v. 178, p. 165–198. doi:10.1016/j.geobios.2009.06.001.
t
ALMOGI-LABIN, A., BEIN, A., AND SASS, E., 1993, Late Cretaceous upwelling GRADSTEIN, F., OGG, J., AND SMITH, A., 2004, A Geologic Time Scale:
system along the southern Tethys margin (Israel)—Interrelationship between Cambridge, U.K., Cambridge University Press, 589 pp.
in
productivity, bottom water environments, and organic-matter preservation: HANSEN, T.A., AND UPSHAW, B., 1990, Aftermath of the Cretaceous/Tertiary
Paleoceanography, v. 8, p. 671–690. Extinction: Rate and nature of early Paleocene molluscan rebound, in
BARRERA, E., AND KELLER, G., 1990, Stable isotope evidence for gradual Kauffman, E.G., and Walliser, O.H., eds. Extinction Events in Earth History:
environmental changes and species survivorship across the Cretaceous/ New York, Springer-Verlag, p. 402–409.
Tertiary boundary: Paleoceanography, v. 5, p. 867–890. HANSEN, T., FARRAND, R.B., MONTGOMERY, H.A., BILLMAN, H.G., AND
BEESON, D.C., 1992, High Resolution Palynostratigraphy across the Cretaceous– BLECHSCHMIDT, G., 1987, Sedimentology and extinction patterns across the
Pr
Tertiary Boundary Interval, Falls County, Texas: Ph.D. thesis, The Cretaceous–Tertiary boundary interval in east Texas (USA): Cretaceous
Pennsylvania State University, University Park, 341 p. Research, v. 8, p. 229–252.
BERGGREN, W.A., KENT, D.V., SWISHER, C.C., III, AND MILLER, K.G., 1995, A HANSEN, T.A., UPSHAW, B., III, KAUFFMAN, E.G., AND GOSE, W., 1993a, Patterns of
revised Paleogene geochronology and chronostratigraphy, in Berggren, W.A., molluscan extinction and recovery across the Cretaceous–Tertiary Boundary
Kent, D. V., and and Hardenbol, J., eds., Geochronology, Time Scales, and in east Texas: Report on new outcrops: Cretaceous Research, v. 14, p. 685–
Global Stratigraphic Correlations: A Unified Temporal Framework for an 706.
Historical Geology. SEPM Special Publication, v. 54, p. 129–213. HANSEN, T.A., FARRELL, B.R., UPSHAW, B., III, 1993b, The first 2 million years
after the Cretaceous–Tertiary boundary in east Texas: Rate and paleoecology
241, p. 567–570.
E-
BOURGEOIS, J., HANSEN, T.A., WIBERG, P.L., AND KAUFFMAN, E.G., 1988, A
Tsunami deposit at the Cretaceous–Tertiary Boundary in Texas: Science, v.
CANDE, S.C., AND KENT, D.V., 1995, Revised calibration of the geomagnetic
of the molluscan recovery: Paleobiology, v. 19, p. 251–265.
HART, M.B., AND BALL, K.C., 1986, Late Cretaceous anoxic events, sea-level
changes and the evolution of the planktonic foraminifera, in Summerhayes,
C.P., and Shackleton, N.J., eds., North Atlantic Palaeoceanography:
polarity timescale for the Late Cretaceous and Cenozoic: Journal of
Geological Society of America, Special Publication v. 21, p. 67–78.
Geophysical Research, v. 100, p. 6093–6095.
HART, M.B., FEIST, S.E., HAKANSSON, E., HEINBERG, C., PRICE, G.D., LENG, M J.,
CARON, M., 1985, Cretaceous planktic foraminifera, in Bolli, H.M., Saunders,
AND WATKINSON, P.M., 2005, The Cretaceous–Palaeogene boundary succes-
J.B., and Perch-Nielsen, K., eds., Plankton Stratigraphy: Cambridge, U.K.,
sion at Stevns Klint, Denmark: Foraminifers and stable isotope stratigraphy:
Cambridge University Press, p. 17–86.
Palaeogeography, Palaeoclimatology, Palaeoecology, v. 224, p. 6–26.
COCCIONI, R., AND LUCIANI, V., 2004, Planktonic foraminifera and environmental
or
HAQ, B.U., HARDENBOL, J., AND VAIL, P.R., 1987, Chronology of fluctuating sea
changes across the Bonarelli Event (OAE2, latest Cenomanian) in its type
levels since the Triassic: Science v. 235, p. 1156–1167.
area: a high-resolution study from the tethyan reference Bottaccione section HEYMANN, D., YANCEY, T.E., WOLBACH, W.S., THIEMENS, M.H., JOHNSON, E.A.,
(Gubbio, central Italy): Journal of Foraminiferal Research, v. 34, p. 109–129. ROACH, D., AND MOECKER, S., 1998, Geochemical markers of the Cretaceous–
COCCIONI, R., AND LUCIANI, V., 2005, Planktonic foraminifers across the Tertiary Boundary Event at Brazos River, Texas, USA: Geochimica et
Bonarelli Event (OAE2, latest Cenomanian): The Italian record: Palae- Cosmochimica Acta, v. 62, p. 173–181.
ogeography, Palaeoclimatology, Palaeoecology, v. 224, p. 167–185. HUBER, B.T., MACLEOD, K.G., AND TUR, N.A., 2008, Chronostratigraphic
th
COCHRAN, J.K., LANDMAN, N.H., TUREKIAN, K.K., MICHARD, A., AND SCHRAG, framework for upper Campanian–Maastrichtian sediments on the Blake Nose
D.P., 2003, Paleoceanography of the Late Cretaceous (Maastrichtian) Western (Subtropical North Atlantic): Journal of Foraminiferal Research, v. 38, p.
Interior Seaway of North America: evidence from Sr and O isotopes: 162–182.
Palaeogeography, Palaeoclimatology, Palaeoecology v. 191, p. 45–64. KELLER, G., 1988, Extinctions, survivorship and evolution across the
DETTMAN, D.L., AND LOHMANN, K.C., 2000. Oxygen isotope evidence for high Cretaceous/Tertiary boundary at El Kef, Tunisia: Marine Micropaleontology,
altitude snow in the Laramide Rocky Mountains of North America during the v. 13, p. 239–263.
late Cretaceous and Paleogene: Geology v. 28, p. 243–246.
Au
KELLER, G., 1989a, Extended Cretaceous/Tertiary boundary extinctions and

DONOVAN, A.D., BAUM, G.R., BLECHSCHMIDT, G.L., LOUTIT, T.S., PFLUM, C.E., AND delayed population change in planktic foraminifera from Brazos River, Texas:
VAIL, P.R., 1988, Sequence stratigraphy setting of the Cretaceous–Tertiary Paleoceanography, v. 4, p. 287–332.
boundary in central Alabama, in Wilgus, C.K., Hastings, B.K, Posamentier, KELLER, G., 1989b, Extended period of extinctions across the Cretaceous–
H., Wagoner., J.V., Ross, C.A., and Kendall, C.G. eds., Sea Level Changes: Tertiary boundary in planktic foraminifera of continental-shelf Sections—
An Integrated Approach: SEPM Special Publication 42, p. 299–307. implications for impact and volcanism theories: Geological Society of
FAIRBANKS, R., SVERDLOVE, M., FREE, R., WIEBE, P.H., AND BÉ, A.W., 1982, America Bulletin, v. 101, p. 1408–1419.
Vertical distribution and isotopic fractionation of living planktonic KELLER, G., 1996, The K/T mass extinction in planktic foraminifera biotic
foraminifera from the Panama Basin: Nature, v. 298, p. 841–844. constraints for catastrophe theories, in MacLeod, N., and Keller, G., eds., The
FAURE, G., 1986, Principles of Isotope Geology, Second Edition: Wiley, New Cretaceous–Tertiary Mass Extinction: Biotic and Environmental Events: New
York, 589 pp. York, Norton Press, p. 49–84.
FRANK, T.D., THOMAS, D.J., LECKIE, R.M., ARTHUR, M.A., BOWN, P.R., JONES, K., KELLER, G., 2002, Guembelitria dominated late Maastrichtian planktic
AND LEES, J.A., 2005, The Maastrichtian record from Shatsky Rise (northwest foraminiferal assemblages mimic early Danian in the Eastern Desert of
Pacific): A tropical perspective on global ecological and oceanographic Egypt: Marine Micropaleontology, v. 47, p. 71–99.
changes: Paleoceanography, v. 20. PA1008. doi:10.1029/2004PA001052. KELLER, G., 2003, Biotic effects of impacts and volcanism: Earth and Planetary
GALE, A.S., 2006, The Cretaceous–Palaeogene boundary on the Brazos River, Science Letters, v. 215, p. 249–264.
Falls County, Texas: Is there evidence for impact-induced tsunami KELLER, G., 2004, Paleoecology of Late Maastrichtian–early Danian planktic
sedimentation?: Proceedings of the Geologists’ Association, v. 117, p. 173– foraminifera in the eastern Tethys: Journal of Foraminiferal Research, v. 34, p.
185. 49–73.
KELLER, G., 2005, Biotic effects of late Maastrichtian mantle plume volcanism: LI, L., AND KELLER, G., 1999, Variability in Late Cretaceous climate and deep
implications for impacts and mass extinctions: Lithos, v. 79, p. 317–341. waters: evidence from stable isotopes: Marine Micropaleontology, v. 161,
KELLER, G., AND ABRAMOVICH, S., 2009, Lilliput effect in late Maastrichtian p.171–190.
planktic foraminifera: Response to environmental stress: Palaeogeography, LUCIANI, V., AND COBIANCHI, M., l999, The Bonarelli level and other black shales
Palaeoclimatology, Palaeoecology, v. 271, p. 52–68, doi:10.1016/j.palaeo. in the Cenomanian–Turonian of the northeastern Dolomites (Italy):
2008.09.007. Calcareous nannofossil and foraminiferal data: Cretaceous Research, v. 20,
KELLER, G., AND PARDO, A., 2004, Disaster opportunists Guembelitrinidae— p. 135–167.
Index for environmental catastrophes: Marine Micropaleontology, v. 53, p. MACLEOD, N., ORTIZ, N., FEFFERMAN, N., CLYDE, W., SCHULTER, C., AND
83–116. MACLEAN, J., 2000, Phenotypic response of foraminifera to episodes of global
KELLER, G., BARRERA, E., SCHMITZ, B., AND MATTSON, E., 1993, Gradual mass environmental change, in Culver, S.J., and Rawson, P., eds., Biotic Response
extinction, species survivorship, and long-term environmental changes across to Global Environmental Change: The Last 145 Million Years: Cambridge,
the Cretaceous–Tertiary boundary in high-latitudes: Geological Society of U.K., Cambridge University Press, pp. 51–78.
America Bulletin, v. 105, p. 979–997. MASTERS, B.A., 1993, Re-evaluation of the species and subspecies of the genus
t
KELLER, G., LI, L., AND MACLEOD, N., 1995, The Cretaceous/Tertiary boundary Plummerita Brönnimann and a new species of Rugoglobigerina Brönnimann
stratotype section at El Kef, Tunisia: How catastrophic was the mass (Foraminiferida): Journal of foraminiferal research v. 23, p. 267–274.
in
extinction?: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 119, p. MÉDUS, J., 1992, Spores et Pollens du passage Cretace/Tertiaire d’une coupe de
221–254. Brazos River (Texas, U.S.A.): Archives de Science, Geneve, v. 45, p. 127–
KELLER, G., ADATTE, T., STINNESBECK, W., STÜBEN, D., KRAMAR, U., BERNER, Z., 133.
LI, L., AND VON SALIS PERCH-NIELSEN, K., 1998, The Cretaceous–Tertiary NEDERBRAGT, A.J., 1991, Late Cretaceous biostratigraphy and development of
transition on the shallow Saharan Platform in southern Tunisia: Geobios, v. Heterohelicidae (planktic foraminifera): Micropaleontology, v. 37, p. 329–
30, p. 951–975. 372.
Pr
KELLER, G., HAN, Q., ADATTE, T., AND BURNS, S.J., 2001, Paleoenvironment of the NEDERBRAGT, A., AND FIORENTINO, A., l999, Stratigraphy and paleoceanography
Cenomanian–Turonian transition at Eastbourne, England: Cretaceous of the Cenomanian–Turonian boundary event in Oued Mellegue, northwest-
Research, v. 22, p. 391–422. ern Tunisia: Cretaceous Research, v. 20, p. 47–62.
KELLER, G., ADATTE, T., BERNER, Z., HARTING, M., BAUM, G., PRAUSS, M. NEDERBRAGT, A.J., AND KONING, J.A., 1994, Morphologic variation in Turonian to
TANTAWY, A.A.M., AND STÜBEN, D., 2007a, Chicxulub Impact predates K–T Maastrichtian Heterohelix globulosa (Ehrenberg): Koninklijke Nederlandse
boundary: New evidence from Brazos, Texas: Earth and Planetary Science Akademie van Wetenschappen, Proceedings, v. 97, p. 429–444.
Letters, v. 255, p. 339–356. NEDERBRAGT, A.J., ERLICH, R.N., FOUKE, B.W., AND GANSSEN, G.M., 1998,
Palaeoecology of the biserial planktonic Foraminifer Heterohelix moremani
E-
KELLER, G., ADATTE, T., TANTAWY, A.A., BERNER, Z., STINNESBECK, W., AND
STUEBEN, D., 2007b, High stress late Maastrichtian–early Danian palae-
oenvironment in the Neuquen Basin, Argentina: Cretaceous Research, v. 28,
p. 939–960.
(Cushman) in the late Albian to middle Turonian circum-North Atlantic:
Palaeogeography, Palaeoclimatology, Palaeoecology, v. 144, p. 115–133.
NORDT, L., ATCHLEY, S., AND DWORKIN, S., 2003, Terrestrial evidence for two
greenhouse Events in the latest Cretaceous: Geological Society of America
KELLER, G., ADATTE, T., BAUM, G., AND BERNER, Z., 2008, Reply to ‘Chicxulub
Today, v. 13, p. 4–9.
Impact predates K–T Boundary: New evidence from Brazos, Texas’
OLSSON, R.K., WRIGHT, J.D., AND MILLER, K.G., 2001, Paleobiogeography of
Comment by Schulte et al.: Earth and Planetary Science Letters, v. 269, p.
Pseudotextularia elegans during the latest Maastrichtian global warming
620–628, doi:10.1016/j.epsl.2007.12.025.
event: Journal of Foraminiferal Research, v. 31, p. 275–282.
KELLER, G., ABRAMOVICH, S., BERNER, Z., AND ADATTE, T., 2009a, Biotic effects
PARDO, A., AND KELLER, G., 2008, Biotic effects of environmental catastrophes at
of the Chicxulub impact, K–T catastrophe and sea-level change in Texas:
or
the end of the Cretaceous: Guembelitria and Heterohelix blooms: Cretaceous
Palaeogeography, Palaeoclimatology, Palaeoecology, v. 271, p. 52–68, doi:10.
Research, v. 29, p. 1058–1073, doi:10.1016/j.cretres.2008.05.031.
1016/j.palaeo.2008.09.007.
PARDO, A., ORTIZ, N., AND KELLER, G., 1996, Latest Maastrichtian and K/T
KELLER, G., ADATTE, T., BERNER, Z., PARDO, A., AND LOPEZ-OLIVA, L., 2009b, Age boundary foraminiferal turnover and environmental changes at Agost, Spain,
and biotic effects of the Chicxulub impact in Mexico: Geological Society of in McLeod, N., and Keller, G., eds., Biotic and Environmental Events across
London, Journal, v. 166, p. 393–411, doi:10.1144/0016-76492008-116. the Cretaceous/Tertiary Boundary: New York, W.W. Norton & Company, p.
KENNEDY, W.J., GALE, A.S., AND HANSEN, T.A., 2001, The last Maastrichtian 139–171.
th
ammonites from the Brazos River sections in Falls County, Texas: Cretea- PIERRE, CH., 1999, The carbon and oxygen isotope distribution in the
ceous Research, v. 22, p. 163–171. Mediterranean water masses: Marine Geology, v. 153, p. 41–55.
KHIM, B.K., AND PARK, B.K., 2000, Recent change of hydrographic features in PREMOLI SILVA, I., AND SLITER W.V., 1999, Cretaceous paleoceanography:
the Cheju Strait (southeastern Yellow Sea), Korea: contrast between Evidence from planktonic foraminiferal evolution, in Barrera E., and
planktonic and benthic foraminifera isotope composition: Marine Geology Johnson, C.C., eds., Evolution of the Cretaceous Ocean Climate System:
v. 169, p. 411–419. Geological Society of America, Special Paper 332, p. 301–328.
KROON, D., AND NEDERBRAGT, A.J., l990, Ecology and paleoecology of triserial
Au
ROHLING, E.J., AND DE RIJK, S., 1999. Holocene Climate Optimum and Last
planktic foraminifera: Marine Micropaleontology, v. 16, p. 25–38. Glacial Maximum in the Mediterranean: the marine oxygen isotope record:
LECKIE, R.M., 1987, Paleoecology of mid-Cretaceous planktonic foraminifera: Marine Geology, v. 153, p. 57–75.
A comparison of open ocean and epicontinental sea assemblages: SCHULTE, P., SPEIJER, R., MAI, H., AND KONTNY, A., 2006, The Cretaceous–
Micropaleontology, v. 33, p. 264–276. Paleogene (K–P) boundary at Brazos, Texas: Sequence stratigraphy,
LECKIE, R.M., YURETICH, R., WEST, O.L.O., FINKELSTEIN, D., AND SCHMIDT, M.G., depositional events and the Chicxulub impact: Sedimentary Geology, v.
1998, Paleoceanography of the southwestern western interior sea during the 184, p. 77–109.
time of the Cenomanian–Turonian boundary (Late Cretaceous), in Dean, W., SCHULTE, P., SPEIJER, R.P., BRINKHUIS, H., KONTNY, A., CALEYS, P., GALEOTTI, S.,
and Arthur, M.A., eds., Stratigraphy and Paleoenvironments of the AND SMIT, J., 2008, Comment on the paper; ‘Chicxulub impact predates K–T
Cretaceous Western Interior Seaway, USA: SEPM, Concepts in Sedimentol- Boundary: New evidence from Brazos, Texas’ by Keller et al. (2007): Earth
ogy and Paleontology, no. 6, p. 101–126. and Planetary Science Letters, doi:10.1016/j.epsl.2007.11.066.
LI, L., AND KELLER, G., 1998a, Maastrichtian climate, productivity and faunal SCHULTE, P., AND 40 OTHERS, 2010, The Chicxulub asteroid impact and mass
turnovers in planktic foraminifera of South Atlantic DSDP sites 525A and 21: extinction at the Cretaceous–Paleogene boundary: Science, v. 327, p. 1214–
Marine Micropaleontology, v. 33, p. 55–86. 1218.
LI, L., AND KELLER, G., 1998b, Diversification and extinction in Campanian– SMIT, J., ROEP, T.B., ALVAREZ, W., MONTANARI, A., CLAEYS, P., GRAJALES-
Maastrichtian planktic foraminifera of northwestern Tunisia: Eclogae Geo- NISHIMURA, J.M., AND BERMUDEZ, J., 1996, Coarse-grained clastic sandstone
logica Helvetica, v. 91, p. 75–102. complex at the K–T boundary around the Gulf of Mexico: Deposition by
LI, L., AND KELLER, G., l998c, Abrupt deep-sea warming at the end of the tsunami waves induced by the Chicxulub impact, in Ryder, G., Fastovsky, D.,
Cretaceous: Geology, v. 26, p. 995–998. and Gartner, S. eds., The Cretaceous–Tertiary Event and Other Catastrophes
in Earth History: Geological Society of America, Special Paper, 307, p.151–

182.
SMITH, C.A., AND PESSAGNO, E.A., JR., 1973, Planktonic foraminifera and
stratigraphy of the Corsicana Formation (Maastrichtian) north-central Texas:
Cushman Foundation for Foraminiferal Research, Special Publication 12, 68 p.
TANTAWY, A.A., 2003, Calcareous nannofossil biostratigraphy and paleoecology
of the Cretaceous–Tertiary transition in the western desert of Egypt: Marine
Micropaloentology, v. 47, p. 323–356.
WEISS, W., 1983, Heterohelicidae (seriale planktonische Foraminiferen) der
tethyalen Oberkreide (Santon bis Maastricht): Geologisches Jahrbuch (A), v.
72, p. 1–96.
WOLFF, T., GRIEGER, B., HALE, W., DÜSRKOOP, A., MULITZA, S., PÄSTZOLD, J., AND
WEFER, G., 1999, On the reconstruction of paleosalinities, in Fischer, G., and
t
Wefer, G., eds., Use of Proxies in Paleoceanography: Examples from the
South Atlantic: Springer, Berlin, p. 207–228.
in
Pr
E-
or
th
Au
PLATE 1.—Guembelitria, Globigerinelloides, and Globotruncanella in the late Maastrichtian zones CF1 to CF3, Brazos River Wells Mullinax-1
and Mullinax-3. Scale bar 1–7 ¼ 10 lm, scale bar 8–15 ¼ 50 lm.
1–5. Guembelitria cretacea Cushman. Mullinax-1, sample 383, 26–2, 42.865 m.
6. Guembelitria(?) irregularis Morozova. Note the atypical presence of costae on the surface of the chambers, and absence of pore mounds
suggests mutational morphology of a biserial test.
7. Globigerinelloides multispinus (Lalicker). Specimens with two lateral ultimate chambers. Mullinax-3, sample 61, 5–3, 8.38 m.
8. Globigerinelloides praevolutus (Petters). Mullinax-1, sample 359, 24–3, 40.475 m.
9, 10. Globigerinelloides asper (Ehrenberg). Mullinax-1, sample 359, 24–3, 40.475 m.
(9) Side view of a specimen showing one of the lateral apertures.
11. Globigerinelloides messinae (Broennimann). Mullinax-3, sample 66, 5–1, 8.69 m.
t
12. Globigerinelloides prairiehillensis Pessagno. Mullinax-3, sample 61, 5–3, 8.38 m. 13, Globigerinelloides subcarinatus Brönnimann.
Mullinax-1, sample 379, 26–1, 42.475 m.
in
14, 15. Globotruncanella petaloidea Gandolfi. Mullinax-1, sample 383, 26–2, 42.865 m.
16. Globotruncanella havanensis Voorwijk. Mullinax-3, sample 71, 6–1, 9.19 m.
PLATE 2.—Heterohelix species from late Maastrichtian zones CF1 to CF3 in the Brazos River wells Mullinax-1 and Mullinax-3. Scale bar ¼ 50
Pr
lm.
1–6. Heterohelix globulosa (Ehrenberg). (1, 2) Specimens with strongly reduced costae show incipient appearance; Mullinax-1, sample 101,
5–1, 10.6 m. (3–6) Specimens with discontinuous fine costae; Mullinax-1, Sample 88, 4–4, 9.73 m.
7–11. Heterohelix striata (Ehrenberg). Note the coarse costae and variation in chambers size and numbers. (7–9) Specimens from Mullinax-1,
sample 88, 4–3, 9.73 m; (10) Mullinax-3 sample 34, 4–2, 6.55 m.; (11) transitional form between H. striata and H. labellosa. Mullinax-1,
sample 303, 18–3, 30.975 m. E-
12, 13. Heterohelix sp. Note the reniform shape of the ultimate chamber and the backward oriented periapertural flanges. Mullinax-1, sample
153, 8–1, 15.155 m.
14. Heterohelix ultimatumida White; note that the last-formed chamber or pair of chambers is/are much more inflated than the others.
15–17. Heterohelix planata (Cushman). (15), Mullinax-1, sample 88, 4–3, 9.73 m (16, 17) Mullinax-1, sample 108, 5–2, 10.97 m.
PLATE 3.—Biserial morphologies (Paraspiroplecta, Spiroplecta, Heterohelix, Fleisherites and Pseudogumbelina) from late Maastrichtian zones
or
CF1 to CF3 in Brazos River Wells Mullinax-1 and Mullinax-3. Scale bar: 1–5, 9, 11–14 ¼ 10 lm, 6–8, 15–19 ¼ 50 lm).
1–4. Paraspiroplecta (Heterohelix) navarroensis (Loeblich), Mullinax-3, sample 77, 6–2, 9.79 m. Tests with early planispiral coil originally
described as Heterohelix navarroensis. New genus Paraspiroplecta erected by Georgescu and Abramovich (2008) to accommodate forms with
early planispiral coil, coarse costae, and weakly acute periphery of the early part of the test.
5. Spiroplecta americana Ehrenberg, Mullinax-1, Note the early planispiral stage, the globular chambers in adult specimens, fine costate
ornamentation, and the absence of weakly acute periphery. The definition of this species was emended by Georgescu and Abramovich (2009).
th
Spiroplecta differs from Paraspiroplecta mainly by lacking the weekly acute appearance of the peripheral test and being finely striate rather
than coarse costae.
6–9. Heterohelix labellosa Nederbragt. Note the reniform shape of the chambers that are covered by thick costae. (6) Mullinax-1, sample 323,
21–1, 34.075 m. (7–9) Mullinax-1, sample 98, 5–1, 10.4 m. (9) close up of chamber surface shows large pores between costae.
10–12. Fleisherites (Laeviheterohelix) glabrans (Cushman). Note the reniform shape of the chambers. This species was formerly identified as
Au
Laeviheterohelix glabrans Nederbragt, 1991. Georgescu (2009) has emended the genus Laeviheterohelix based on its polyphyletic nature. (10–
12) Mullinax-3, sample 80, 4–2, 9.28 m, 13, Fleisherites (Laheviheterohelix) pulchra? Mullinax-1, sample 325, 21–2, 34.285 m. Note the tube
shape ultimate chamber and the thickening of the costae in earlier chambers. Although this morphology is here included in F. pulchra, this
specimen differs by the unique shape of its ultimate chambers and by the presence of costae rather than a smooth surface; it may be a new
species.
14, 15. Pseudoguembelina costulata (Cushman). Small specimens with broken last chambers. Note the absence of supplementary apertures
and the relatively large size of the pores. Mullinax-1, sample 325, 34.285 m.
16–19. Pseudoguembelina excolata (Cushman). Note strong costae and supplementary apertures. (15–17, 19) Mullinax-1, sample 98, 5–1,
10.4 m; (18) Mullinax-1, sample 325, 34.285 m.
PLATE 4.—Specimens of Pseudogumbelina, Racemiguembelina, and Planoglobulina from the late Maastrichtian zones CF1 to CF3 in Brazos
River wells Mullinax-1 and Mullinax-3. Scale bar ¼ 50 lm.
1–3. Pseudoguembelina palpebra Bronnimann and Brown. Note the relatively large sized pores. (3) Specimen with broken last chamber and
no supplementary apertures. Mullinax-1, sample 90, 4–3, 9.89 m.
4. Pseudoguembelina kempensis Esker. Mullinax-3, Sample 77, 6–2, 9.79 m, small-sized adult specimen.
5, 6. Pseudoguembelina hariaensis Nederbragt. (5) Intermediate form between P. hariaensis and P. palpebra as suggested by the large-sized
pores. Mullinax-3, sample 34, 4–2, 6.55 m. 7, 8. Pseudotextularia elegans Rzehak. Mullinax-1, sample 325, 21–2, 34.285 m. Note that this
morphology is frequently identified as P. deformis in earlier studies by Keller, Abramovich and others following Weiss (1983). However, this
morphology was defined as P. elegans based on the holotype of Rzehak, 1891 (Nederbragt, 1990; Li and Keller, 1998a, 1998b).
9, 10. Racemiguembelina powelli Smith and Pessango. Mullinax-1, sample 91, 4–3, 9.95 m.
11, 12. Planoglobulina brazoensis (Martin). (11) Mullinax-1, sample 325, 21–2, 34.28 m. (12) Mullinax-1, sample 108, 5–2, 10.97 m.
t
13–16, Planoglobulina carseyae (Plummer). (13) Intermediate form between P. carseyae and P. brazoensis. Mullinax-3, sample 53, 2–2, 7.9
in
m. (14) Mullinax-1, sample 107, 10.9 m. (15) Mullinax-1, sample 99, 5–1, 10.42 m. (16) Mullinax-1, sample 385, 26–2, 43.06 m.
PLATE 5.—Rugoglobigerina and Trinitella species from late Maastrichtian zones CF1 to CF3, Brazos River wells Mullinax-1 and Mullinax-3.
Scale bar ¼ 50 lm.
Pr
1. Intermediate form between Rugoglobigeriana rugosa (Plummer) and Rugloglobigerina pennyi Brönnimann. Mullinax 3, sample 47, 5–2,
7.53 m.
2–4. Rugoglobigeriana rugosa (Plummer). Mullinax-1, sample 85, 4–3, 9.55 m.
5. Intermediate form between Rugoglobigeriana rugosa and Rugoglobigerina hexacamerata Brönnimann. Mullinax 1, sample 85, 4–3, 9.55 m.
6–10. Rugoglobigeriana macrocephalla Brönnimann. Mullinax-1, sample 85, 4–3, 9.55 m. 11, Rugoglobigerina hexacamerata Brönnimann.
12–16. Trinitella scotti Brönnimann. Mullinax-1, sample 105, 5–2, 10.78 m (13–14). Specimens with early chambers covered with spines.
E-
PLATE 6.—Plummerita, Rugoblobigerina, Archaeoglobigerina, Hedbergella and Globotruncanella in the late Maastrichtian zones CF1 to CF3,
Brazos River wells Mullinax-1. Scale bar ¼ 10 lm for 5, 12–13, and 50 lm for 1–4, 6, 7–11, 14–15.
1, 2. Plummerita hantkeninoides Brönnimann. Mullinax-1, sample 110, 5–2, 11.09 m.
3–5. Rugoblobigerina reicheli Brönnimann. Mullinax-1, sample 89, 4–3, 9.8 m.
6, 7. Intermediate forms between Rugoglobigerina rugosa and Archaeoglobigerina cretacea (d’Orbigny) Mullinax 1, sample 85, 4–3, 9.55 m.
Note the hispid surface of the test.
or
8, 9. Archaeoglobigerina cretacea (d’Orbigny). Mullinax-1, sample 85, 4–3, 9.55 m. 10, 11. Rugoblobigerina rotundata Brönnimann. Mullinax-
1, sample 80, 4–2, 9.28 m.
12, 13. Hedbergella monmouthensis (?). Note apertural lip appears broken. Mullinax-1, sample 303, 18–3, 30.97 m.
14, 15. Globotruncanella minuta Caron and Gonzalez Donoso. Mullinax-1, sample 365, 41.07 m.
th
PLATE 7.—Globotruncana species from late Maastrichtian zones CF1 to CF3, Brazos River wells Mullinax-1 and Mullinax-3. Scale bar ¼ 50 lm.
1–8. Globotruncana aegyptiaca Nakkady. (1) Note the atypical high density of pustules on surface of the chambers and incipient spinose
appearance of the two keels. Mullinax-1, sample 98, 5–1, 10.36 m.
9–12. Globotruncana duwi Nakkady. (9, 10) Note the high density of pustules on surface of the chambers. Spiral side characterized by three
Au
chambers, rapidly increasing in size with the ultimate chamber comprising one half of the test. Mullinax-1, sample 103, 5–2, 10.66 m.
13–16. Globotruncana arca (Cushman). (13) Umbilical view of juvenile specimens with 3 and a half chambers and a narrow double keel.
Mullinax-1, sample 86, 4–3, 9.61 m. (14) Spiral view of poorly developed adult with four and a half chambers. Mullinax-3, sample 53, 5–2, 7.9
m. (15, 16) Umbilical view of juvenile or poorly developed adult specimens with double keel that merge along the ultimate chamber. Mullinax-3,
sample 53, 5–2, 7.9 m.
PLATE 8.—Globotruncana and Globotruncanita species from late Maastrichtian zones CF1 to CF3, Brazos River wells Mullinax-1 and Mullinax-3.
Scale bar ¼ 50 lm.
1–5. Globotruncana arca (Cushman). Mullinax-3, sample 53, 5–2, 7.9 m. (1–3) Umbilical view of specimens with well-developed double keel.
(4, 5) Note that final whorl consist of only 5.5 chambers.
6, 7. Globotruncana orientalis El Naggar. Mullinax-1, sample 100, 5–1, 10.49 m.
8–10. Globotruncana esnehensis Nakkady. Mullinax-3, sample 74, 9.49 m. (8) Umbilical view of specimens with 4.5 chambers in the final whorl.
11. Globotruncana rosetta (Carsey). Mullinax-3, sample 60, 5–3, 8.31 m.
12–16. Globotruncanita stuartiformis (Dalbiez). Mullinax-3, sample 74, 6–2, 9.49 m. Note the characteristic circular outline of the test and the
atypical small number of chambers of the last whorl.
t
in
Pr
E-
or
th
Au
PLATE 1.
t
in
Pr
E-
or
th
Au
Plate 2.
t
in
Pr
E-
or
th
Au
Plate 3.
t
in
Pr
E-
or
th
Au
Plate 4.
t
in
Pr
E-
or
th
Au
Plate 5.
t
in
Pr
E-
or
th
Au
Plate 6.
t
in
Pr
E-
or
th
Au
Plate 7.
t
in
Pr
E-
or
th
Au
Plate 8.

TMP 4 F17

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

TMP 4 F17

Uploaded by

Copyright:

Available Formats

Author E-Print 1/3/2012

MAASTRICHTIAN PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY AND

MATAN CYMBALISTA AND CARMI RAK

The End-Cretaceous Mass Extinction and the Chicxulub Impact in Texas

For this study we chose well Mullinax-1, which contains a nearly

larger species. In Mullinax-3 the . 38 lm size fraction was also analyzed

terrigenous influx and transitions from dysoxic to oxic conditions on

elegans is very rare. In addition, P. elegans first appears in zone CF6 in

sharply at these episodes by as much as 2ø, relative to background

Ecological Opportunists Persistent and Sporadic Taxa

Guembelitria Blooms: Disaster Opportunists

significant in this section of the . 63 lm size fraction is the

interval consists of a sandy, glauconitic mudstone with clasts, shell

FIGURE 16.—Guembelitria and Heterohelix blooms during the late

Late Maastrichtian Stable Isotope Depth Ranking of Planktic Foraminifera

mixed layer Thermocine Subthermocline

Surface Subsurface intermediate Deep

Pseudoguembelina excolata Globotruncana aegyptiaca Globotruncana arca Planoglobulina multicamerata

Species also present in Brazos area (this study)

DISCUSSION diversity and morphologically smaller species (, 150 lm). In these

logging support from Schlumberger during two drilling phases. We

KELLER, G., 1989a, Extended Cretaceous/Tertiary boundary extinctions and

in Earth History: Geological Society of America, Special Paper, 307, p.151–

You might also like