Palynoassemblages associated with a theropod dinosaur from the Snow Hill Island Formation (Lower Maastrichtian) at The Naze, James Ross Island, Antarctica

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Review paper
Palynoassemblages associated with a theropod dinosaur from the
Snow Hill Island Formation (lower Maastrichtian) at the Naze,
James Ross Island, Antarctica
Mercedes di Pasquo
a
,
*
, James E. Martin
b
,
c
a
Consejo de Investigaciones Cientificas y Tecnológicas (CONICET), Laboratorio de Palinoestratigrafía y Paleobotánica, CICyTTP-CONICET,
Dr. Matteri y España s/n, Diamante CP E3105BWA, Entre Ríos, Argentina
b
J.E. Martin Geoscientific Consultation, 21051 Doral Ct., Sturgis, SD 57785, USA
c
School of Geosciences, University of Louisiana, Lafayette, LA 70504, USA
article info
Article history:
Received 3 February 2013
Accepted in revised form 27 July 2013
Available online
Keywords:
Palynostratigraphy
Paleoenvironment
Snow Hill Island Formation
Early Maastrichtian
James Ross Island
Antarctica
abstract
The Cape Lamb Member of the Snow Hill Island Formation at The Naze on the northern margin of James
Ross Island, east of the Antarctic Peninsula, yielded a theropod dinosaur recovered near the middle of a
90 m thick section that begins at sea level, ends below a basalt sill, and is composed of interbedded green
egray massive and laminated fine-grained sandstones and mudstones. Sixteen palynoassemblages were
recovered from this section, which yielded moderately diverse assemblages with a total of 100 relatively
well-preserved species. The principal terrestrial groups (32%) are represented by lycophytes (8 species),
pteridophytes (15 species), gymnosperms (13 species), angiosperms (21 species) and freshwater chlor-
ococcaleans (3 species). Marine palynomorphs (68%) belong to dinoflagellates (61 species), chlor-
ococcaleans (6 species), and one acritarch. The vertical distribution of selected species allows the
distinction of two informal assemblages, the lower Odontochitina porifera assemblage from the base to its
disappearance in the lower part of the section, and the remaining section characterized by the Batia-
casphaera grandis assemblage. The global stratigraphic ranges of selected palynomorphs suggest an early
Maastrichtian age for this section and the entombed dinosaur that is also supported by the presence of
the ammonoid Kitchinites darwinii. These assemblages share many species with latest Campanianeearly
Maastrichtian palynofloras from Vega and Humps Islands, New Zealand, and elsewhere in the Southern
Ocean, establishing a good correlation among them. The dominance or frequent presence of di-
noflagellates throughout the section supports the general interpretation of a shelf marine depocenter.
The consistent presence of terrestrial palynomorphs suggests contributions from littoral/inland
environments.
Ó2013 Elsevier Ltd. All rights reserved.
1. Introduction
The Late Cretaceous James Ross Basin (JRB) in the Antarctic
Peninsula (Fig. 1) has a long history of scientific studies and today it
is well known for extensive paleontological records. Determination
of the age of these fossils derived from the The Naze in the JRB is a
major goal of this contribution. Currently, the base of the Maas-
trichtian Stage in Antarctica is formally defined on the basis of a
mean
87
Sr/
86
Sr value (0.7077359) for the six best-preserved sam-
ples from a bivalveenautiloid assemblage within the Gunnarites
antarcticus assemblage from Cape Lamb, Vega Island (Crame et al.,
1999). An absolute age of 71.0 0.2 Ma was obtained for a strati-
graphic level 81.5e96.5 m above the base of the G. antarcticus
assemblage corresponding to the CampanianeMaastrichtian
boundary (Crame et al., 2004). This boundary figures significantly
in this contribution where the palynostratigraphic analysis of
sixteen productive samples from the Cape Lamb Member of the
Snow Hill Island Formation, were utilized to provide the age of a
small theropod dinosaur found in this section (Case et al., 2003,
2007), located on the western side of Comb Ridge at The Naze,
northern margin of James Ross Island, east of the Antarctic Penin-
sula (Figs. 1 and 2). Previous palynological studies from The Naze
were provided by Askin (1988a), who documented a short list of
palynomorphs from two samples derived from the basal López de
Bertodano succession. In conjunction with other assemblages from
*Corresponding author.
E-mail addresses: medipa@cicyttp.org.ar (M. di Pasquo), JEMartinGeoscientific@
gmail.com (J.E. Martin).
Contents lists available at ScienceDirect
Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes
0195-6671/$ esee front matter Ó2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.cretres.2013.07.008
Cretaceous Research 45 (2013) 135e154

Vega and Seymour Islands, she proposed an informal zonation
based on first appearances of several species (including Manu-
miella). Because The Naze assemblage was considered to lie below
the Vega Island assemblage (Fig. 2), she considered The Naze
assemblage as midelate Campanian. This zonation from the López
de Bertodano succession on Seymour Island was partially reviewed
by Bowman et al. (2012) who proposed a zonation based principally
on the first appearance of several species of Manumiella (Fig. 2). The
age assignment herein for the palynoassemblages is proposed
based upon the stratigraphic ranges of diagnostic palynomorph
species coupled with biostratigraphic ranges of ammonoid species
from The Naze section.
2. Materials and methods
The Upper Cretaceous deposits from which the Antarctic fossils
(mega and micro) were collected are referred to the upper Cam-
panian to lower Maastrichtian Cape Lamb Member of the Snow Hill
Island Formation at The Naze of the James Ross Island (Figs. 1e3). As
can be observed on Fig. 3, the Cape Lamb Member on the western
portion of Comb Ridge (The Naze) is represented by a rather
monotonous 90-m-thick section that begins at sea level and ends
below a Miocene basalt sill. This Cape Lamb section is composed of
interbedded greenegray massive and laminated fine-grained
quartz sandstones and greenish yellow argillaceous mudstones
and siltstones, interbedded with concretions and bentonite layers.
The bentonites are found principally in the lower portion of the
section, below the dinosaur level, but some layers occur above. The
calcareous concretions are either scattered or concentrated in
layers of gray claystone matrix. Interestingly, the theropod dinosaur
from the middle of the section was found with several vertebrates
(Martin et al., 2007; Case et al., 2003, 2007), invertebrates (am-
monites, pelecypods, decapods), and palynomorphs deposited in a
shelf environment. Both Diplomoceras lambi and Kitchinites darwini
were found in the upper part of the section (Figs. 2 and 3). For
biostratigraphical analysis, sixteen samples of argillaceous mud-
stones and siltstones were selected from the base and top of the
section, as well as from close to significant megafossils (theropod,
ammonoids, bivalves; see Fig. 3). Samples were processed using
standard palynological methods (HCl, HF) at the Laboratory of
Palynostratigraphy and Paleobotany (Department of Geology, Nat-
ural and Pure Sciences Faculty, University of Buenos Aires) in 2006.
All were productive, and several slides were prepared with sieved
residues (þ25 and þ10
m
m) mounted with glycerin jelly. Slides and
residues are housed at the Palynostratigraphy and Palaeobotany
Laboratory of the CICYTTPeCONICET in Diamante (Entre Ríos,
Argentina).
Identification of palynomorphs was undertaken using trans-
mitted light microscopes and digital video camera Nikon Eclipse 80i
(with DIC objectives) with a Pax-it (3.1 Mp) at the Laboratory of
Palynostratigraphy and Paleobotany (Department of Geology, Natu-
ral and Pure Sciences Faculty, University of Buenos Aires), and a Leica
DM500 with a fluorescence LED lamp (cold light) attached with a
fluorescein (ca. 450 nm) filterblock and a 3.0 Mp videocamera (Leica
EC3) at the CICYTTP. Different colors of the microphotographs are the
result of differing equipment. A brief survey of the autofluorescence
of palynomorphs wasconducted, and some species of dinoflagellates
and few species of spores showed light to dark orange auto-
fluorescence. Many others did not present autofluorescence (i.e.
extinguished fluorescence gray to black). The position of illustrated
specimens in the respective figures, quoted with the CICYTTP-Pl
acronym, are based on EnglandeFinder coordinates.
Fig. 1. Antarctic map with location of the outcrop bearing the theropod fossil remains and palynological samples here studied at the The Naze (modified from Crame et al., 2004). To
the right, photograph of The Naze, a peninsula on the northern coast of James Ross Island (photo by J.E. Martin).
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e154136

3. Geological setting
For the Upper Cretaceous James Ross Basin (JRB) in the
Antarctica, Crame et al. (1991) and Crame (1992) presented a
CampanianeMaastrichtian stratigraphic summary of the James
Ross Island (JRI) combining stratigraphic distribution of di-
noflagellates and ammonoids with that of other invertebrate fossils.
The lowereupper Maastrichtian boundary was isotopically reas-
sessed by Crame et al. (1999) based on strontium isotope dates of
the base and top of the stage on Vega, Snow Hill, and Seymour
Islands, James Ross Island region (northeastern Antarctic Penin-
sula). A comprehensive stratigraphical revision integrating lithol-
ogy, biostratigraphy (e.g., ammonoids, other invertebrates,
palynomorphs) and chronostratigraphy of Antarctica was pre-
sented by Crame et al. (2004). Ammonoids, in particular, were
studied by several authors (see Olivero and Medina, 2000 and
references therein). Olivero and Medina (2000) defined three
stratigraphic sequences for the SantonianeDanian succession of
the James Ross Basin; the N, Santonianelower Campanian; the NG,
upper Campanianelower Maastrichtian; and the MG, lower
MaastrichtianeDanian, sequences. The names of these sequences
were derived from the most common kossmaticeratid ammonites
that characterize each of them: N for Natalites; NG for Neo-
grahamites and Gunnarites; and MG for Maorites and Grossouvrites
(Fig. 2). Recently, Olivero (2012) addressed the sedimentary cycles,
ammonite diversity, and paleoenvironmental changes in the Upper
Cretaceous Marambio Group in Antarctica. He indicated that the
radiation and extinction patterns in the NG and MG sequences,
which are dominated by the relatively endemic kassmaticeratids,
do not reflect the enlargement and reduction of the shelf during
transgressions and regressions.
Other paleontological records include wood fossils (e.g., Francis,
1986, 1991) and paleovertebrates, especially marine reptiles (e.g.,
disarticulated plesiosaur and mosasaur skeletons, sharks teeth and
vertebrae) were also recorded and studied from several places in
the JRB (Martin et al., 2002, 2007; Martin and Crame, 2006; Case
et al., 2007, and their references). Several palynological studies
were also carried out in this basin by Baldoni (1992), Baldoni and
Barreda (1986), Dettmann (1986), Dettmann and Thomson
(1987), Baldoni and Medina (1989), Askin (1988a, 1988b, 1990a,
1990b, 1994, 1999), Pirrie et al. (1991, 1997), Riding et al. (1992),
Smith (1992), Dolding (1992), Wood and Askin (1992), Thorn
et al. (2009), Bowman et al. (2012). Some studies were conducted
in different areas on James Ross Island (and closer islands) and were
published together in 1992 (e.g., Pirrie et al., 1992). Of these studies,
those of Smith (1992) from Cape Lamb on Vega Island and Dolding
(1992) from Humps Island compared most favorably (Fig. 2) with
those analyzed herein.
4. Palynological results
The marine and terrestrial assemblages recovered from the
Cape Lamb Member are diverse with numerous specimens per
sample level and fairly well-preserved with minimal thermal
maturation. The overall composition of the assemblage (sixteen
sample levels), is represented by spores (5.3%), pollen grains (18%),
chlorophytes (8.7%), dinoflagellates and acritarch (66.6%), and
remaining groups (2%). The principal terrestrial groups (32%) are
represented by lycophytes (8 species), pteridophytes (15 species),
gymnosperms (13 species), angiosperms (21 species) and fresh-
water chlorococcaleans (3 species). Marine palynomorphs (68%)
belong to dinoflagellates (61 species), chlorococcaleans (6 species),
and one acritarch. Complementary groups such as bryophytes,
sphenophytes, organic foraminiferal linings, microthyriaceae fungi
and copepod eggs are also intermittently present (Figs. 3 and 4).
Fig. 2. Stratigraphy after Crame et al. (2004) and Olivero (2012), and absolute date after Crame et al. (1999). Sequence stratigraphy after Olivero and Medina (2000) and Olivero (2012):Natalites (N), Santonianelower Campanian;
Neograhamites and Gunnarites (NG), upper Campanianelower Maastrichtian; and the Maorites and Grossouvrites (MG), lower MaastrichtianeDanian. Species of palynomorphs with an asterisk are here considered under synonymy.
Species of ammonoids with an asterisk are recorded in this study. Other biostratigraphic schemes from southern Gondwana are as follows: 5-Australia (Helby et al., 1987), 6-Kerguelen Plateau (Mao and Mohr, 1992), 7-Maud Rise,
Antarctica (Mohr and Mao, 1997), 8-New Zealand (Roncaglia et al., 1999), 9-James Ross Basin, Peninsula Antarctica (Vega Island: Smith, 1992, 25 species in common of 55, including the lower Maastrichtian sample D3122.3 in Dettmann
and Thomson, 1987, with 40 species in common of 63; Humps Island: Dolding, 1992, 40 species in common of 140; Wood and Askin, 1992, 9 species in common of 43). Abbreviations: N-The Naze, V-Vega.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e154 137

Fig. 3. Stratigraphic section at The Naze showing quantitative occurrences of main groups of palynomorphs after their biological affinities and in stratigraphical order. The database is presented in Fig. 4. Their relative frequencies (based
on counts of 250 to 400 specimens per sample) are represented as follows: R-rare (<1%), P-Present (<5%), C-Common (<10%), A-Abundant (10e30%) and VA-(>30e60%, considered as a “bloom”or peak of abundance). Location of other
fossils is also depicted. TeM-Terrestrial:Marine Ratio.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4138

Semi-quantitative stratigraphic distribution of selected paly-
nomorphs along the section is provided in Fig. 5. The complete list
of taxa (with full authority) documented at The Naze is presented
in Fig. 4, ordered by major groups and following a stratigraphical
order. Illustrations of selected species (Figs. 7e11) are in the
Appendix A.
The database of Raine et al. (2011) was used to classify most of
the terrestrial palynomorphs along with their botanical affinities,
whereas dinoflagellates and algal identification were chiefly based
on the Dinoflaj Database (Fensome et al., 2008), although later
taxonomical contributions were also used (e.g., Thorn et al., 2009;
Sluijs et al., 2009). Taxonomical remarks are addressed in the list
of taxa, along with synonymies of several species illustrated,
especially by Askin (1988a), among other authors (see Fig. 2 and
Chart 1, supplementary online material). All synonymies were
carefully based on identical morphologies; hence ancillary remarks
are included only when deemed necessary. The known global
stratigraphic ranges of selected palynomorphs are depicted in Fig. 6
based on information provided in Chart 1 (Supplementary online
information).
SPECIES / Repository numb er (CICYTTP-Pl) 286 287 288 289 51 52 290 291 292 293 29 4 8 9 295 10 296
AB Spores / field sample num be 62-CA12-CA02-CA51-CA41-CA31-CA1-21-CA01-CA5-CA2-CAr AC-30 AC-35 AC-40 AC-43 AC-46 AC-48
LY Perot rilites maju s (Cookson and Dettmann 1958) Evans 1970 1,90% 1,71% 0,53% 0,92% 1,65% 1,32% 0,85% 1,51% 1,65% 0,29% 1,28%
LY
Retitriletes austroclavatidites (Cookson 1953) Doring et al. in
Krutzsch 1963 0,95% 1,37% 1,06% 0,92% 1,89% 1,91% 1,65% 2,19% 1,41% 1,33% 3, 52% 0,89% 0,38% 1,65% 2,00% 0,85%
FBaculatisporites comaumensis (Cookson 1953) Potonié 1956 0,24% 0,34% 0,27% 0,69% 1, 26% 0,27% 0,55% 1,32% 0,28% 1,01% 0,30% 1,53% 0,83% 1,28%
FCyathidites australis /C. minor Couper 1953 %68,0%95,0%58,0%44,0%55,0%23,0%
96,0%72,0%43,0%42,0
FBirretisporites potoniaei Delcourt and Sprumont 1955 %23,0%42,0 0,29%
FTodisporites major /T. minor Couper 1958 %34,0%92,0%95,0%33,1%55,0%72,0%96,0%42,0
LY Ceratosporites equalis Cookson and Dettmann 1958 %64,0%74,0 1,13%
BTriporoletes radiatus (Dettm ann 1963) Playford 1971 0,47% 2,74% 0,23% 1,10% 0,28% 0,29%
F
Laevigatosporites ovatus Wilson and Webster 1946/L. major
(Cookson) Krutzsch 1959 %5
5,0%23,0%43,0 0,50% 0, 89% 0,38% 0,29%
B
Stereisporites antiquasporite
s
(Wilson and Webster) Dettm ann
1963 0,34% 0,30%
FIschyosporites volkheimeri Filatoff 1975 %82,0%55,0%72,0 0,43%
LY Herkosporites sp. 0,23%
FPolypodiisporites sp. 0,23%
LY Densoisporites velatus Weyland and Kri eger 1953 0,23% 0,38%
BStereisporites regium (Drozhastichich) Drugg 1967 0,63% 0,30% 0,38% 0,57%
F
Gleicheniidites senonicu
s
Ross 1949 (=Gleiche nidites cir cinidites
(Cookson) Brenner) 0,32% 0,43%
FCyatheacidites annulatus Cookson 1947 0,55% 3,33%
LY Lycopodiumsporites eminulus Dettmann 1963 0,44% 0,83%
LY
Camarozonosporites ohaiensis (Couper 1953) Dettmann and
Playford 1968 0,28%
SCalamospora sp. 0,67%
FTrilites parvallatus Krutzsch 1959 0,67%
FCyatheacidites archangelskii Dettmann 1986 6,00% 1,01%
LY Dictyotosporites speciosus Cookson and Dettmann 1958 0,50%
FTuberculatosporites parvus Archangelsky 1972 0,50%
FPolypodiisporites favus (Potonié) Potonié 1934 0,38%
FPeromonolites bowenii Couper 1953 0,38% 0,29%
F
cf. Klukisporites scaberis (Cookson and Dettmann 1958)
Dettmann 1963 0,29%
Gymnosperm pollen grains
AAraucariacites australis Cookson 1947 0,47% 0,68% 1,61% 1,26% 0,44% 1,69% 0,59% 0,38% 1,65% 1, 43%
PPhyllocladidites mawsonii Cookson 1947 ex Couper 1953 0,71% 1,37% 1,06% 2,30% 1,26% 2,46% 1,65% 1,32% 1,13% 0,67% 4,73% 3,44% 0, 83% 4,00% 0,43%
PPodocarpidites rugulatus Pocknall and Mil denhall 1984 0,47% 0,68% 0,23% 1,58% 3,01% 1,32% 0,67% 0,50% 0,30% 0,76% 1,14% 0,85%
PPodocarpidites spp. 4,03% 2,40% 5,31% 4, 61% 0, 32% 3,83% 5,49% 4,39% 12,96% 3,33% 3,02% 1,78% 4,58% 6,61% 7,43% 2,13%
ADilwynites granulatus Harris 1965 0,95% 0,34% 0,80% 0, 46% 0, 63% %92,2%81,1
PPodocarpidites marwickii Couper 1953 %58,0%92,0%95,0
%88,0%72,0%59,0%43,0%42,0
PPodocarpidites major Couper 1953 %23,0%32,0%42,0
PTrichotomosulcites subgranulatus Couper 1953 0,24% 1,09% 0,28% 4,14% 1,15% 1,65% 0,57% 0, 43%
PPodocarpidites otagoensis Couper 1953 %34,0%95,0%05,0%76,0%82,0%36,0%35,0
PMicrocachrydites antarcticus Cookson 1947 ex Couper 1953 %41,1%03,0%76
,0%82,0%36,2%28,0%59,0%32,0%35,0
PDacrycarpites australiensis Cookson and Pike 1953 0,27% 0,88% 0,56% 0,67%
EEquisetosporites sp. 0,95%
PPodocarpidites verrucosus Volkheim er 1972 0,57%
Angiosperm pollen grains
NNothofagidites dorotensis Romero 1973 0,24% %34,0%03,0%82,0
NNothofagidites spp 0, 71% 3,42% 0,27% 1,84% 13,88% 11,20% 1,10% 0,28% 0,67% 16,27% 5,73% 1,65% 12,00%
Pr Penins ulapollis gillii (Cookson 1957) Dettmann and Jarzen 1988 0,34% 0,53% 0,46% 2,52% 1,91% 1,65% 0,44% 1,13% 0,67% 0, 50% 1,48% 1,65% 3,43% 0,85%
Pr Penins ulapollis askin iae Dettm ann and Jarzen 1988 0,34% 0,53% 0, 46% 0, 32% %92,0%95,0
C-T
Periporopollenites polyoratus (Couper 1960) Stover in Stover and
Partridge 1973 %76,0%82,0%
23,0%43,0 0,29%
Pr Proteacidites spp. %73,1%43,0 1,18% 0,38%
Er Ericipit es scabra tus Harris 1965 0,27% 0,28% 1,01% 0,83%
Pr Proteacidites tenuiexinus Stover in Stover and Partridge 1973 0, 27% 0,27%
M-L Liliacid ites spp. 0,92% 1,26% 0,55% 0,59%
NNothofagidites americanus Zamaloa 1992 %95,0%65,0%36,0%64,0
DTriorites orbiculatus McIntyre 1965 0,23% 0,83% 0,29%
MMonosulcites palisadus Couper 1953 0,63%
MMonosulcites/Arecipites spp. 1,58%
NNothofagidites tehuelchesii Zamaloa and Barreda 1992 0,32%
DTricolpites confessus Stover in Stover and Partridge 1973 %03,0%10,1%82,0%23,0
C-M
Myricipites harrisii (Couper 1953) Dutt a and Sah 1970
(=Haloragacidites trioratus Couper 1953 0,63% %34,0%92,0%03,0
DBatte nipollis sect ilis (Stover) Jarzen and Dettmann 1991 0,28% 0,29%
NNothofagidites saraensis Menéndez and Caccavari de Fílice 1975 0,30%
Pr Proteacidites scaboratus Couper 1960 0,38%
Pr Proteacidites parvus Cookson 1950 0,38% 0,86%
M-L Longapertites sp. in Povilauskas et al. 2008 0,85%
A
Fig. 4. Percentages and stratigraphic distribution of taxa documented along the Comb Range section at The Naze. Biological affinities are also included in the row on the left.
Abbreviations: B-Bryophyte, LY-Lycophyte, S-Sphenophyte, F-FilicophyteePterophyte (Fern), A-Araucariaceae, P-Podocarpaceae, E-Ephedraceae, N-Nothofagaceae, Pr-Proteaceae,
Er-Ericaceae, CeM-Casuarinaceae, possibly also Myricaceae, CeT-Caryophyllaceae, Trimeniaceae, M-Monocotyledonae, MeL-Monocotyledonae, D-Dicotyledonae, Per-
Peridiniaceae, Gon-Gonyaulacaceae, Cer-Ceratiacea, Clo-Chlorophyceae, Ac-Acritarch, Mf-Linings of forams, Fg-Fungi eMicrothyriaceae (Asterothyrites Cookson), Co-Copepods eggs.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 139

Chlorophyceae
Clo Botryococcus brauni Kützing 1849 4,74% 2,05% 0,80% 2,07% 3,15% 4,64% 2,19% 3,94% 5,03% 5,92% 1,15% 4,29% 2,55%
Clo Nummus monoculatus Morgan 1975 3,08% 2,05% 0,27% 1,61% 2,21% 0,27% 2,75% 3,07% 3,66% 1,33% 1, 51% 1,78% 1,15% 1,14% 2,13%
Clo Palambages spp. 1,42% 0,34% 0,80% 0,92% 0,55% 1,10% 3,07% 0,56% 0,67% 1,51% 0 %41,1%03,
Clo Leiosphaeridia sp. %82,1%67,0%95,0%10,2%28,2%36,2%94,02%36,0%00,3%74,0
Clo Tetraedron cf. minimum (A. Braun) Hansgirg 1988 0,32% %34,0%83,0
Clo Pterospermella spp. 0,27% 0,30%
Clo Dictyotidium sp. 0,82% 0,59%
Clo
Paralecaniella indentata (Deflandre and Cookson) Cookson and
Eisenack 1970 emend Elsik 1977 0,24% 0,34% 0,53% 1,32% 0,28% 0, 67% 1,51% 0,76% 0,83% 1,28%
Clo
Pterospermella autraliensis (Deflandre and Cookson) Eisenack
1972 0,44%
Acritarchs
Ac Michrystridium piliferum Deflandre 1937 14,22% 27,40% 1,86% 1,84% 5,36% 12,30% 5,49% 4,23% 6,67% 5,03% 0,89% 0,76%
Mf Linnings of forams %41,1%98,0%82,0%55,0%55,0%36,0%08,0
Fg Fungi – Microthyriacea: Asterothyr ites Cookson 1947 0,23% 0,29% 0,85%
Co Copepods eggs 0,68% 0,27% 0,46% 11,04%
Dinoflagellate
Per
Chatangiella victoriensis (Cookson and Manum) Lentin and
Williams 1977 1,42% 0,68%
Gon Cyclonephelium compactum Deflandre and Cookson 1955 0,47%
Per
Diconodinium sp. cf. multispinum (Deflandre and Cookson)
Eisenak and Cookson emend Morgan 1977 0,24% 0,27% 0,50%
Gon Operculodinium flucturum Davey 1969 1,90%
Per Xenikoon australis Cookson and Eisenack 1960 0, 95% 0,56%
Per
Amphidiadema nucula (Cookson and Eisenack) Lentin and
Williams 1976 1,66% 1,37% 0,80% 1,84% %31,2%00,2%51,1
%65,0
Gon Valensiella reticulata (Davey) Courti nat 1989 0,24% 0,28%
Gon
Trichodinium castanea (Deflandre) Clarke and Verdier 1967 - T.
chilens is Troncoso and Doubinger 1980 0,24% 0,27% 1,15% 15,77% 0,27% 4,40% 21,93% 1,41% 40,00% 4,02% 11,24% 17,18% 4,96% 4,57% 8,51%
Cer Odontochitina porifera Cookson 1956 1,90% 4,11% 1, 86% 1,84%
Per Andalusiella mauthei R iegel 1974 %82,0%57,1%55,0%32,0%42,0
Per Manumiella druggii (Stover 1974) Bujak and Davies 1983 0,24% 1,01%
Per Manumiella seymourensis Askin 1999 0,95% 1,03% 1,06% 1,61% 0,55% 0,30% 5,79% 0,57%
Per Maduradinium pentagonum Cookson and Eisenack 1970 0,24% 0,68% 1,06% 1,15% 0,32% 0,55% 0,88% 0,56% 1,01% 0,76% 1,65% 1,28%
Per Manumiella spp. 0,24% 1,03% 0,85% 0,30%
Gon Batiacasphaera grandis Roncaglia et al. 1999 1, 66% 1,37% 2,39% 0,92% 0,32% 0,27% 1,65% 2,19% 1,69% 1,33% 7,54% 2,96% 8,02% 3,31% 2,00% 4,26%
Gon Canningia sp. 1 in Schiol er and Wilson 1998 4,27% 0,34% 0,53% 1,84% 7,69% 1,13% 2,01% 0,43%
Per
Chatangiella granulifera/verrucosa (Manum 1963) Lentin and
Williams 1976 7,11% 6,85% 0,44%
Per
Isabelidinium cretaceum (Cookson 1956) Lentin and Williams
1977 11,85% 7,88% 18,57% 12, 67% 16,72% 15,30% 5,49% 1,75% 14,93% 2,00% 5, 03% 12,72% 16,79% 12,40% 15,43% 4,26%
Per I. cretaceum gravidum Mao and Mohr 1992 2,37% 3,77% 2,65% 2,07% 0,32% 0,27% 5,49% 1,75% 1,97% 2,00% 5,03% 0,89% 8,26% 4,26%
Per
Isabelidinium pellucidum (Deflandre and Cookson 1955) Lentin
and Williams 1977 4,74% 0,34% 1,59% 0,92% 0,63% 0,27% 2,20% 1,13% 0,67% 0,59% 0,76% 1,65% 0,57% 2,13%
Gon Spongodinium reticulatus Hultberg 1985 1%92,0
%03,0%30,5%65,0%88,0%23,0%29,0%72,0%43,0%42,0 ,28%
Per Nelsoniella aceras Cookson and Eisenack 1960 0,71% 0,68% 0,27% 0,23% 0,32% 0,27% 1,65% 0,28% 0,50% 1,65%
Gon Operculodinium radiculatum Smith 1992 10,66% 2,05% 2,65% 7,60% 1,89% 2,73% 2,75% 2,63% 3,38% 13,33% 5,03% 9, 76% 2,29% 4,13% 4,86% 8,51%
Per Isabellidinium sp. 1,03% 1,59% 0,92% 0,32% 0,55% 5,49% 2,63% 1,33% 3,02% 0,59% 4,13% 0,57% 4,26%
Per
Palaeocystodinium granulatum (Wilson 1967) Lentin and W illiams
1976 0,24% 0,68% 0,27% 0,88% 0,50%
Gon
S. ramosus (ramosus-multibrevis-granosus) complex (Sp iniferite s
ramosus (Ehrenberg 1838) Mantell 1854 4,74% 4,45% 21,22% 12, 21% 1, 58% 3,01% 10,99% 13,16% 9,86% 2,00% 7,54% 4,73% 7,25% 4,96% 6,29% 8,51%
Cer Xenascus plotei Below 1981 0,47% 0,68% 14,59% 6,22% 4,92% 10,99% 3,51% 2,25% 1,51% 4,13% 3,43% 8, 51%
Per Paleocystodinium pilosum Guler et al. 2005 0,47% 0,34% 1,86% 1,84% 0,27% 0,28% 0,67% 2,51% 0,89% 8,78% 3,31% 2,57% 1,28%
Per Saeptodinium gravattensis Harris 1973 %07,1%83,0%03,0%57,1
%72,0%83,1%72,0%81,1
Per Eurydinium ellipticum Mao and Mohr 1992 %64,0%17,0 2,55%
Per Canninginopsis ordospinosa Smith 1992 0,24% 6,85% 1,06% 0,32% 1,37% 3,30% 4,39% 3,94%
Gon Impagidinium cristatum (May) Lentin and Williams 1981 %34,0%38,0%03,0%05,0%82,0%55,0%72,0%42,0
Cer Odontochitina indigena Marshall 1988 0,34% 0,43%
Per Isabellidinium cretaceum oviforme Mao and Mohr 1992 1,71% 1,38% 0,32% 0,55% 2, 75% 1,69% 3,52% 4,13% 2,29% 4,26%
Gon Batiacasphaera? reticulata (Davey) Davey 1979 0%56,1%6
7,0%95,0%15,2%76,2%65,0%91,2%56,1%23,0%43,0 ,29% 1,28%
Per Phelodinium exilicornutum Smi th 1992 0,34% 0,23% 1,65% 1,75% 0,56% 1,33% 2,01% 0,30% 4,13% 1,28%
Gon Membranilarnacia spp. 0,27% 0,23% %34,0%83,0
Per Isabellidinium papillum Sumner 1992 0,27% 0,32% 0,85%
Per Alterbidinium acutulum (W ilson) Lentin and Williams 1976 %82,0%55,0%72,0
Per Canninginopsis bretonica Marshall 1990 0,53% 1,32% 1,13%
Gon
Operculodinium centrocarpum (Deflandre and Cookson) Wall
1967 4,51% 8,06% 4,79% 0,50% 0,59% 0,83% 1,71% 1,28%
Per Paleocystodinium golzowense Alberti 1961 0,53% 2,29% 0,57% 0, 85%
Gon Batiacasphaera rifensis Slimani et al. 2008 0,27% 0,56% 0,83%
Per Manumiella conorata (Stover) Bujak and Davies 1983 %67,0%20,3%88,0%64,0
Per Manumiella seelandica (Lange) Bujak and Davies 1983 0,23% 0,32% 0,55% 0, 67% 0,30% 5,73% 0,29%
Gon Kallosphaeridium? ringnesorium in Mohr and Mao 1997 1,15%
Cer Xenascus ceratioides (Deflandre) Davey and Verdier 1971 0,69% 0,85% 1,65% 1,14% 2,13%
Gon
Circulodinium distinctum (Deflandre and Cookson) Jansonius
1986 0,32% 0,89% 0,76% 0,57%
Gon Batiacasphaera rugulata Schiøl er and Wilson 1998 1,65% 0,28% 1,51% 0,43%
Per Spinidium e ssoi Cookson and Eisenack 1967 0,88% 0,28%
Gon Spiniferites pseudofurcatus (Klump) Sarjeant 1970 1,32% 0,43%
Per
Diconodinium cristatum Cookson and Eisenack 1974 emend
Morgan 1977 0,44%
Per Cerodinium sp. in Mehrotra and Sarjeant 1987 0,28%
Gon Kallosphaeridium parvum Jan du Chêne 1988 0,56%
Per Indeterminate Dinoflagellate 0,28%
Gon Pterodinium cretaceum Slim ani et al. 2008 0,67% 0,43%
Per Isabellidinium cooksoniae (Alberti) Lentin and W illiams 1977 0,50%
Per Manumiella sp. cf. M. bertodano Thorn et al. 2009 2,01% 0,30% 4,13%
Per Senegalinium bicavatum Jain and Millepied 1973 0,50%
Per Nelsoniella tuberculata Cookson and Eisenack 1960 0,85%
822281663
713434773292224detnuocsnemicepsforebmuN 355 150 199 338 262 121 350 235
SPECIES / Repository number (CI CYTTP-Pl) 286 287 288 289 51 52 290 291 292 293 294 8 9 295 10 296
B
Fig. 4. (continued).
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4140

Fig. 5. Relative frequencies (based on counts of 250e400 specimens per sample) of selected taxa along the section are represented as follows: R-rare (<1%), P-Present (<5%), C-Common (<10%), A-Abundant (10e30%) and VA-(>30e
60%, considered as a “bloom”or peak of abundance). The database is presented in Fig. 4.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 141

Spores
Biretisporites potoniaei Delcourt and Sprumont 1955 (Fig. 7C)
Birretisporites sp. Barreda et al., 1999; Pl. 1, fig. 1.
Birretisporites sp. A Dettmann and Thomson, 1987; Fig. 3, b.
Camarozonosporites ohioensis (Couper) Dettmann and Playford
196 8
Camarozonosporites ambigens (Fradkina) Playford,1971 (auct.
non); Barreda et al. (1999), Pl. 1, Fig. 7.
Triporoletes radiatus (Dettmann) Playford 1971 (Fig. 7N)
Gen and sp. Askin, 1988a, Fig. 8.10.
Gen and sp. Askin, 1988a, Smith (1992), Fig. 11, mep.
Gen and sp. Askin, 1988a, Dolding (1992), Fig. 6, n.
Pollen
Gymnosperm
Podocarpidites otagoensis Couper 1953 (Fig. 7M)
Podocarpidites elegans Romero, 1977; Pl. 3, Figs. 1e11.
Dinoflagellate
Batiacasphaera grandis Roncaglia et al. 1999 (Fig. 10K, L)
Canningia? sp.1 Ioannides and McIntyre,1980; Pl. 31.2, Fig.1, 3.
Batiacasphaera rifensis Slimani et al. 2008 (Fig. 9A)
Batiacasphaera cf. kekerengensisMa renssi et al., 2004; Fig. 5N,O.
Batiacasphaera? reticulata (Davey) Davey 1979 (Fig. 9C, D)
Batiacasphaera sp. Pirrie et al., 1997; Fig. 10i.
Canninginopsis ordospinosa Smith 1992 (Fig. 9BB)
cf. Canninginopsis sp. Askin, 1988a; Fig. 7.6.
Impagidinium cristatum (May) Lentin and Williams 1981
(Fig. 10E, F)
Pterodinium sp. Schiøler and Wilson, 1998; Pl. 6, Fig. 8.
Isabelidinium cretaceum gravidum Mao and Mohr 1992 (Fig. 9B)
Isabellidinium cf. bakeri Smith, 1992; Fig. 7f.
Odontochitina indigena Marshall 1988 (Fig. 10O)
Xenascus sp. Wood and Askin, 1992; Fig. 5d.
Operculodinium radiculatum Smith 1992 (Fig. 11F)
Operculodinium sp. Askin, 1988a; Fig. 7.7.
Operculodinium sp. Wood and Askin, 1992; Fig. 4e.
Phelodinium exilicornutum Smith 1992 (Fig. 9U)
Octodinium askiniae Wrenn and Hart 1988 (auct. non), Wood
and Askin, 1992; Fig. 4j.
Phelodinium sp. Askin, 1988a; Fig. 7.8.
Spongodinium reticulatum Hultberg 1985 (Fig. 10B)
Cyclonephelium cf. clathromarginatum Smith, 1992; Fig. 8d.
Leberidocysta sp. A Mohr and Mao, 1997; Pl. 1, Figs. 10, 14.
Trichodinium castaneum (Deflandre) Clarke and Verdier 1967 e
T. chilensis Troncoso and Doubinger 1980, plexus (Fig. 10M, P, Q)
Cribroperidinium sp. Askin, 1988a; Fig. 7.4.
Cribroperidinium sp. A Dettmann and Thomson, 1987; Fig. 8b.
Cribroperidinium muderongense (Cookson and Eisenack)
Davey 1969 (auct. non), Wrenn and Hart, 1988; Fig. 18, 1.
Cribroperidinium edwarsii (Cookson and Eisenack) Davey
1969 (auct. non), Wrenn and Hart, 1988, Fig. 18, 3e4.
Remarks: Specimens attributable to Trichodinium castanea
associated with those of Trichodinium chilensis (from the
upper Maastrichtian of southern Chile) in the same levels
supports our proposal of a taxonomic group (plexus) among
these species. T. chilensis is distinguished from T. castanea
mainly in having a less spheroidal cyst with a longer apical
horn and a more prominent and dense ornamentation of
spines overall except for the equatorial cingulum where the
spines are shorter. Askin (1988a, Fig. 7, 4) illustrated Cri-
broperidinium sp. from Zone 1 at The Naze on JRI and Cape
Lamb on Vega Island that she considered equivalent to Cri-
broperidinium sp. A Dettmann and Thomson (1987, Fig. 8b).
The specimens illustrated by Wrenn and Hart (1988) as Cri-
broperidinium muderongense (Fig. 18, 1) and C. edwarsii
(Fig. 18, 3e4), considered reworked from the Cretaceous in
the Paleogene of JRI in Antarctica, and are so similar to this
group that they are here reassigned to this complex.
Xenascus plotei Below 1981 (Fig. 11H, J)
Xenascus sp. Barreda et al. 1999; Pl. 6, Fig. 13.
Chlorophyceae
Nummus monoculatus Morgan 1975 (Fig. 8R, U)
Cyclopsiella sp. Askin 1988a; Fig. 8.9.
Cyclopsiella sp. Dolding 1992; Fig. 6i, m.
Remarks: This species is smaller (less than 100
m
m) than
N. similis (Cookson and Eisenack) Burger.
5. Biostratigraphy and correlation
The stratigraphic distribution of most of the species from the
sample assemblages exhibit some major shifts with local bio-
stratigraphical importance (Figs. 3e5), for example, at ca. 20 m
from the base, the disappearance of Odontochitina porifera and the
appearance of Manumiella coronata and M. seelandica occur. This
event is located ca. 25 m below the level where the theropod and
decapods occur. The appearance of Penninsulapollis gillii,Proteaci-
dites spp., Alterbidinium acutulum,Batiacasphaera?reticulata,
Manumiella seymourensis,M. druggii,Phelodinium exilicornutum,
Fig. 6. Global stratigraphic range of selected species recorded at The Naze section
(based on information provided in Chart 1 of the supplementary online information).
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4142

M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 143

Canninginopsis bretonica,Batiacasphaera rifensis and a peak of
abundance of Canninginopsis ordospinosa occur in the basal part of
the section as well. Few other species appear from the middle part
of the section (e.g., Battenipollis sectilis,Spinidium essoi,Pterodinium
cretaceum). Instead, other species show variable frequencies
throughout the section (Fig. 5). For example, the most consistently
abundant dinoflagellate species are Trichodinium castaneumechi-
lensis complex, Isabellidinium cretaceum,Operculodinium radi-
culatum,Spiniferites ramosus complex, Batiacasphaera grandis, and
Xenascus plotei, as well as the acritarch Michrystridium piliferum, the
chlorophycean Botryococcus brauni,Nummus monoculatus and
Palambages morulosa, and some podocarpacean (Podocarpus,Phyl-
locladidites) and proteacean (Penninsulapollis gillii; Figs. 3,4).
Although leiosphaerids and arthropod eggs are abundant
throughout the section (Figs. 3,4), other species show shorter or
more restricted vertical distribution (e.g., Alterbidinium acutulum,
Canninginopsis ordospinosa,C. bretonica,Spinidium essoi,Sene-
galinium bicavatum,Battenipollis sectilis), and others appeared
intermittently (e.g., Paleocystodinium granulatum,P. golzowense,
Impagidinium cristatum,Manumiella spp., Ericipites scabratus,Pro-
teacidites). Many of these changes in abundances are interpreted as
variations of environmental conditions (i.e., sea-level changes,
nutrients, bloomings) rather than of biostratigraphic significance.
The global stratigraphic range of selected species depicted in
Fig. 6 (Chart 1, supplementary online information) reveals that
several long-ranging taxa mainly from CampanianeMaastrichtian
age occur throughout this section, notably some trilete spores
(Perotrilites majus), pollen grains (Podocarpidites,Nothofagidites,
Phyllocladidites), dinoflagellates (Isabelidinium cretaceum,
I. pellucidum,Operculodinium radiculatum,Spiniferites ramosus
complex) and chlorococcaleans (Nummus monoculatus and Paral-
ecaniella indentata). Some species appearing in the lowest level
show a close relationship with taxa of the Campanian stage (e.g.,
Cyclonephelium compactum,Diconodinium sp. cf. multispinum,
Xenikoon australis,Canningia sp. 1 in Schiøler and Wilson, Madur-
adinium pentagonum,Battenipollis sectilis,Odontochitina porifera
and Batiacasphaera grandis). A few species are known from the Late
Maastrichtian or Danian such as Saeptodinium gravattensis and
Polypodiisporites favus (Chart 1, supplementary online information).
Key taxa indicate that at least eight dinoflagellate species (Batia-
casphaera rifensis,Impagidinium cristatum,Isabellidinium cretaceum
gravidum,Manumiella seymourensis,M. seelandica,Paleo-
cystodinium pilosum,Spinidium essoi,Spongodinium reticulatum)
appeared in the early Maastrichtian, whereas Odontochitina porifera
and Batiacasphaera grandis disappeared at the end of this time
(Fig. 6). Hence, considering the vertical distribution of selected
species, and the biostratigraphic significance of those key species,
two informal assemblages are defined and named as lower
assemblage Odontochitina porifera and upper assemblage Batia-
casphaera grandis (Figs. 2 and 5). The range of the former is based
on the last occurrence (LO) of Odontochitina porifera, and the latter
is named after a well-represented taxon along the section. Both
assemblages suggest an early Maastrichtian age.
However, Xenascus plotei and X. ceratioides are two dinoflagel-
late cyst that occur mostly together in the same levels throughout
the vertical section (Fig. 4 and Chart 1, supplementary online in-
formation), whose age affiliation may be questioned. The former is
recorded originally in the mid-Cretaceous (post-Aptian) of the USA,
and the latter is widely recorded worldwide (Williams et al., 2004).
Mao and Mohr (1992, pl. 6, figs. 10, 12) illustrated X. ceratioides in
the Campanian of the southern Atlantic Ocean, is better assigned to
X. plotei. Other specimens of X. plotei are illustrated by Smith (1992,
fig. 5m) as occurring from the latest Campanian to earliest Maas-
trichtian of Antarctica. The last occurrence datum (LOD) of
X. ceratioides is documented up to the early Maastrichtian (Premaor
et al., 2010;Chart 1, supplementary online information) and was
illustrated by Williams et al. (2004, pl. 27, figs. 4 and 5) by Schiøler
and Wilson (1998, pl. 8, fig. 9) from the Coniacianeearly Campanian
of New Zealand, and by Premaor et al. (2010,fig. 4N) from the
Campanian of Brazil. Another SantonianeCampanian record of
X. ceratioides in England was mentioned but not illustrated by
Prince et al. (1999). These taxa are here considered indigenous due
to their frequency along the section (Fig. 4,5and Chart 1, supple-
mentary online information) and their fine preservation. In
contrast, scarce specimens of cosmopolitan dinoflagellate such as
Diconodinium species (Fig. 4 and Chart 1, supplementary online
information), and few miospores (e.g., Dictyotosporites speciosus)
are Early Cretaceous taxa that are likely reworked from older rocks
rather than holdovers in this assemblage. Askin (1990a) and
Baldoni (1992) have also documented Early Cretaceous paly-
nomorphs reworked into the CampanianeMaastrichtian of James
Ross and Seymour Islands. These assemblages from The Naze
(Fig. 2) were correlated to selected biostratigraphic schemes from
New Zealand (Roncaglia et al., 1999), Antarctica (Askin, 1988a;
Smith, 1992 and others related), Australia (Helby et al., 1987) and
microfloras from the southern South Atlantic Ocean (Mao and
Mohr, 1992; Mohr and Mao, 1997).
Of these areas, Crame et al. (1991) provided composite strati-
graphical sections for the James Ross Basin, integrating paleonto-
logical information derived from invertebrates and dinoflagellates,
which we utilized for correlation. Our dataset of dinoflagellates and
ammonites from The Naze allowed correlation to the lower Cape
Lamb Member in the Cape Lamb, Vega and northern James Ross
Island regions, based on the co-occurrences of several species, such
as the dinoflagellates Isabellidinium pellucidum,I. cretaceum,
Odontochitina porifera,Canninginopsis bretonica, and the ammonite
Diplomoceras lambi among others. Species of Manumiella were
recorded in the upper Cape Lamb Member of the López de Berto-
dano Formation to its top. The lower Cape Lamb Member was
attributed to the uppermost Campanian and lower Maastrichtian
(Fig. 2 and its references).
Crame et al. (2004) presented a chronostratigraphy for the
1150 m thick Maastrichtian succession in the James Ross Basin
integrating ammonite biostratigraphy and isotopic information.
They placed the lowereupper Maastrichtian boundary at a level of
91 m in section DJ.1042 corresponding to the absolute date of
71 0.2 Ma obtained from the middle portion of the Cape Lamb
Member (Crame et al., 1999) in the concurrent range portionsof the
ammonites Diplomoceras lambi,Kitchinites darwini, and Gunnarites
antarcticus (Pirrie et al., 1991; Crame et al., 1991; Olivero, 2012).
Fig. 7. Illustration of selected palynomorphs occurring within the lithostratigraphic section containing the theropod dinosaur following the order in Fig. 4. Spores and pollen. Scale
bar is 15
m
m. A, Perotrilites majus (Cookson and Dettmann) Evans, CICYTTP-Pl 292-1: Y32/0. B, Baculatisporites comaumensis (Cookson) Potonié, CICYTTP-Pl 919-1: V41/4. C, Bir-
retisporites potoniaei Delcourt and Sprumont, CICYTTP-Pl 51-3: X57/2. D, Retitriletes austroclavatidites (Cookson) Doring et al. in Krutzsch, CICYTTP-Pl 296-1:A30/0. E, Cyatheacidites
archangelskii Dettmann, CICYTTP-Pl 293-1: A34/3. F, Stereisporites regium (Drozhastichich) Drugg, CICYTTP-Pl 51-1: K34-3. G, Densoisporites velatus Weyland and Krieger, CICYTTP-Pl
289-2: W42/4. H, Ischyosporites volkheimeri Filatoff, CICYTTP-Pl 296-1: Q37/1. I, Polypodiisporites sp., CICYTTP-Pl 289-1:T53/0. J, Ceratosporites equalis Cookson and Dettmann,
CICYTTP-Pl 286-3: Z38/4. K, Laevigatosporites major (Cookson) Krutzsch, CICYTTP-Pl 9-2: F26/4. L, Cyatheacidites annulatus Cookson, CICYTTP-Pl 293-1:W57/3 (tetrad). M, Podo-
carpidites otagoensis Couper, CICYTTP-Pl 292-2: R32/0. N, Triporoletes radiatus (Dettmann) Playford, CICYTTP-Pl 287-1: E29/4. O, Podocarpidites rugulatus Pocknall and Mildenhall,
CICYTTP-Pl 296-3: S25/0. P, Dilwynites granulatus Harris, CICYTTP-Pl 286-1: V31/0. Q, Peromonolites bowenii Couper, CICYTTP-Pl 9-2: K30/1. R, Longapertites sp. in Povilauskas et al.,
CICYTTP-Pl 296-3: X28/3. S, Podocarpidites rugulatus Pocknall and Mildenhall, CICYTTP-Pl 293-1: U42/3. T, Podocarpidites major Cookson, CICYTTP-Pl 289-1: R44/4. U, Araucariacites
australis Cookson, CICYTTP-Pl 292-2: V21/3. V, Phyllocladidites mawsonii Cookson ex Couper, CICYTTP-Pl 52-3:C38/0. W, Equisetosporites sp., 51-2: X51/3.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4144

Fig. 8. Illustration of selected palynomorphs occurring within the lithostratigraphic section containing the theropod dinosaur following the order in Fig. 4. Pollen and other re-
mains. Scale bar is 15
m
m except for Figs. Q and X that is 20
m
m. A, Microcachrydites antarcticus Cookson ex Couper, CICYTTP-Pl 10-1: J29/0. B, Battenipollis sectilis (Stover) Jarzen and
Dettmann, CICYTTP-Pl 292-2: Q32/3. C, Proteacidites scaboratus Couper, CICYTTP-Pl 9-3: H27/0. D, Tricolpites confessus Stover in Stover and Partridge, CICYTTP-Pl 51-1: L43/0. E,
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 145

Although they showed that it extends diachronously across the
basin, as it would have been affected by the unconformity at the
base of the Sandwich Bluff Member recorded on Cape Lamb.
The co-occurrence of the first two ammonoid taxa allow the
attribution of this section to the early Maastrichtian Gunnarites
biozone after Crame et al. (2004) or Ammonite Assemblage 10
(Fig. 2) in the NG sequence after Olivero (2012). These authors
agreed that these sequence stratigraphic framework established for
the SantonianeDanian of the James Ross Basin probably represents
a low cyclicity frequency of second or third-order cycles. The time
involved is probably of the order of 7e8 Ma for the N Sequence, at
the beginning of the Santonian to the lowereupper Campanian
boundary; about 8e9 Ma for the NG Sequence, upper Campaniane
lower Maastrichtian; and about 5 Ma for the MG Sequence, lower
MaastrichtianeDanian.
Palynological comparisons with other assemblages of late
Campanianeearly Maastrichtian age from Vega and James Ross
Islands and surrounding areas (Askin, 1988a; Pirrie et al. 1991, 1997;
Riding et al., 1992; Smith, 1992; Dolding, 1992; Wood and Askin,
1992) showed a relatively high number of common species
mainly ranging from late Campanian to early Maastrichtian,
although most of them are extended up to late Maastrichtian (see
Figs. 2,6and Chart 1, supplementary online information).
The most similar palynofloras are from Vega Island, documented
by Smith (1992), who recorded 55 species and, of them, 25 are
shared with our assemblages. The early Maastrichtian age of the
sample D3122.3 described by Dettmann and Thomson (1987)
contained 40 species in common from 63 total species. In Humps
Island, the upper Campanian rocks studied by Dolding (1992)
yielded 140 species and 40 are here in common, whereas Wood
and Askin (1992) from the same place and age, mentioned 43
species and 9 are in common.
The correlation chart presented by Pirrie et al. (1991) for the
López de Bertodano Formation in Cape Lamb, Vega Island illus-
trated a concurrent appearance of Gunnarites antarcticus and Dip-
lomoceras lambieKitchinites darwini with the ranges of
dinoflagellates such as Isabellidinium cretaceum,I. pellucidum and
Canninginopsis bretonica in their B member. They attributed this
assemblage of long-ranging species to the late Campanian and early
Maastrichtian. The frequent occurrence of Manumiella seymourensis
is concurrent with the overlying Maorites assemblage in their C
member, although this dinoflagellate appears slightly earlier in
conjunction with the underlying assemblages as they showed in
their chart of occurrences. Other species of Manumiella (i.e.,
M. seelandica,M. bertodano,M. coronata,M. druggii) were recorded
by Pirrie et al. (1991) later in the late Maastrichtian.
Askin (1988a) studied two samples from a section at The Naze
(Fig. 2) that yielded quite similar assemblages to the ones here
studied, although few species were mentioned, including Odonto-
chitina operculata,O. spinosa,O. porifera,Trichodinium castanea,
Canninginopsis ordospinosa, and a peridinoid cysts plexus, as well as
rare specimens of Operculodinium radiculatum,Paleocystodinium
spp., Nummus similis,Nelsoniella aceras,Phelodinium exilicornutum
(Fig. 6), Triporoletes radiatus, and Spiniferites spp. (see synonymies
of species into item 4). On the basis of this assemblage in
conjunction with assemblages from Vega and Seymour Islands,
Askin (1988a) proposed an informal zonation scheme (Fig. 2).
Bowman et al. (2012) restudied the dinoflagellate assemblages
from the López de Bertodano Formation on Seymour Island, pre-
viously analyzed by Askin (1988a), and proposed a new biozonation
of the late Maastrichtian mainly based on the first appearances of
Manumiella species. The studied section crops out continuously
over approximately 70 km
2
with a thickness of ca. 1000 m and is
bounded by unconformities with the Haslum Crag Member (up-
permost Snow Hill Island Formation) beneath, and with the over-
lying Sobral Formation (Pirrie et al., 1997; Crame et al., 2004;
Olivero et al., 2008). Crame et al. (2004) discussed the Late Maas-
trichtian age given to the base of the López de Bertodano Formation
by Pirrie et al. (1997) based on Manumiella druggii and
M. seymouriensis, and Grapnelispora evansii, considered it in error.
Both Manumiella species were recorded from the late Campanian
Isabellidinium pellucidum Assemblage up to the late Maastrichtian
Manumiella druggii Interval Zone (Fig. 2) in New Zealand (Roncaglia
et al., 1999). This early appearance is in agreement with the record
of both species from the base of the section here studied, and the
introduction of other species successively (M. seelandica,
M. coronata,M. sp. cf. M. bertodano) is in disagreement with the
order of appearances of these species presented by Bowman et al.
(2012) (Figs. 2,5and Chart 1, supplementary online information).
The absence of typically late Maastrichtian marine species, such
as Bosedinia laevigata,Cassidium fragile,Cerodinium medcalfii,Eise-
nackia reticulata, and Palaeoperidinium pyrophorum (see Askin,
1988a; Bowman et al., 2012, and references therein), indicate an
early Maastrichtian age, older in comparison with the palynofloras
from the upper Maastrichtian López de Bertodano Formation pre-
sented by Bowman et al. (2012). In summary, the age of the section
at The Naze, and as a consequence, the age of the theropod, appears
more likely early Maastrichtian than late Maastrichtian; moreover,
the species occurrences are more likely biostratigraphically
controlled rather than paleoecological as indicated by the first
occurrence (FOs) of Batiacasphaera rifensis,Impagidinium cristatum,
Isabellidinium cretaceum gravidum,Manumiella seymourensis,
M. seelandica,Paleocystodinium pilosum,Spinidium essoi, and
Spongodinium reticulatum that are introduced in the early Maas-
trichtian. In addition, the last occurrence (LOs) of Odontochitina
porifera and Batiacasphaera grandis occur at the end of the early
Maastrichtian. Also, the palynofloral evidence is reinforced by the
stratigraphic ranges of the two ammonoid species mentioned
above (see Figs. 2 and 6).
6. Paleoenvironmental approach based on terrestrial and
marine palynomorphs
Marine palynomorphs
Overall, the dominance of marine dinoflagellates in all levels
through this section (Figs. 3e5) at The Naze supports the existence
of a permanent marine depocentre. The predominance of peri-
dinialean cysts with algal remains (e.g., Nummus monoculatus,
prasinophytes) and the acritarch Michrystridium pilosum within the
section, are evidences of more inner neritic settings, close to the
paleoshoreline (Chart 2, supplementary online material). Nummus
monoculatus exhibits an encrusting habit (bentonic), and indicates
Proteacidites tenuiexinus Stover in Stover and Partridge, CICYTTP-Pl 288-1: J26/0. F, Peninsulapollis gillii (Cookson) Dettmann and Jarzen, CICYTTP-Pl 291-1: V41/1. G, Myricipites
harrisii (Couper) Dutta and Sah, CICYTTP-Pl 51-2: O38/4. H, Peninsulapollis askiniae Dettmann and Jarzen, CICYTTP-Pl 51-1: M44/1. I, Liliacidites sp., CICYTTP-Pl 290-1: Q27/2. J,
Nothofagidites americanus Zamaloa, CICYTTP-Pl 8-1: X27/2. K, Ericipites scabratus Harris, CICYTTP-Pl 295-1: B52/1. L, Periporopollenites polyoratus (Couper) Stover in Stover and
Partridge, CICYTTP-Pl 51-2: U46/3. M, Triorites orbiculatus McIntyre, CICYTTP-Pl 10-2: Z28/2. N, Proteacidites parvus Cookson, CICYTTP-Pl 10-1: K30/0. O, Trichotomosulcites sub-
granulatus Couper, CICYTTP-Pl 295-1: S55/0. P, Nothofagidites saraensis Menéndez and Caccavari de Fílice, CICYTTP-Pl 292-2: Z46/3. Q, Botryococcus brauni Kützing, CICYTTP-Pl 289-
2: T49/1. R, Nummus monoculatus Morgan, CICYTTP-Pl 289-2: V51/1. S, Paralecaniella indentata (Deflandre and Cookson) Cookson and Eisenack emend. Elsik, CICYTTP-Pl 293-1:
W57/3. T, Tetraedron cf. minimum (A. Braun) Hansgirg, CICYTTP-Pl 296-1: W32/0. U, Nummus monoculatus Morgan, CICYTTP-Pl 291-1: Q20/0. V, Pterospermella aureolata (Deflandre
and Cookson) Eisenack, CICYTTP-Pl 52-3: C37/0. W, Michrystridium piliferum Deflandre, CICYTTP-Pl 292-2: P21/0. X, Palambages sp., CICYTTP-Pl 289-2: O58/3. Y, Asterothyrites
Cookson, CICYTTP-Pl 296-3: L30/0. Z, Foraminiferal lining, CICYTTP-Pl 51-3: V57/0. AA, Foraminiferal lining, CICYTTP-Pl 290-1: D52/1.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4146

Fig. 9. Illustration of selected palynomorphs occurring within the lithostratigraphic section containing the theropod dinosaur following the order in Fig. 4. Dinoflagellates, Peri-
diniales. Scale bar is 20
m
m for Figs. D, H, I, P, U, DD, and 40
m
m for the remaining figures. A, Isabelidinium cretaceum (Cookson) Lentin and Williams, CICYTTP-Pl 287-1: L34/4. B,
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 147

coastal marine settings; the acritarch is characteristic of the shal-
lowest, inner neritic settings, or partly restricted settings (Duane,
1996;Chart 2, supplementary online material). This occurs at
sample AC-14 where a low number of species and dinocysts suggest
a restricted environment, where only a limited number of taxa can
survive (Figs. 3e5). Conversely, the recovery of dinoflagellates
Spiniferites and Impagidinium (Mudie and Harland, 1996; Prebble
et al., 2006) within the section, together with the frequent pyriti-
zation of palynomorphs and other marine elements such as organic
foraminiferal linings and megafossils (ammonoids and sharks and
plesiosaurs), suggest more open water influences (Figs. 3,4and
Chart 2, supplementary online material). Nevertheless, terrestrial
components such as Podocarpaceae, Nothofagaceae, and Chlor-
ophyceae also occur and are abundant in many sample levels, thus
indicating the coastal line probably was nearby and a fresh-water
influx was permanent, reflecting the surrounding vegetation at
any instant in time (Figs. 3,4and Charts 1 and 2, supplementary
online material). The appearance of the terrestrial dinosaur at AC-
26 level (theropod horizon, Fig. 3) suggests the proximity to a
coastline; however, its poor preservation may be the result of
prolonged transport (although poor preservation is also the result
of extensive subaerial degradation).
Variations in abundances of different marine and terrestrial
palynomorphs (Fig. 3, MT ratio) reflect the influence of variable
terrigenous supplies from deltaic and other terrestrial environ-
ments established around the marine depocenter where variable
conditions (i.e., local sea-level changes, content of nutrients,
bloomings) also would have occurred during the time of deposi-
tion. A high percentage of Trichodinium castaneaechilensis in three
levels of The Naze section (Fig. 3) is interpreted as blooms occurring
in a shallow protected environment, probably the result of an in-
crease in nutrients (Edet and Nyong, 1993; Brinkhuis, 1994;
Matthiessen, 1995; Mudie and Harland, 1996;Chart 2, supple-
mentary online material). Blooms could be induced by upwelling
waters close to the coastline of the continents independently of the
seaward currents carrying land-derived materials to the shelf (e.g.,
Malloy, 1972).
Terrestrial palynomorphs
Cyatheacidites species (tree fern, Lophosoria spores) and species
of NothofagiditesePodocarpaceaeeProteaceae supports the nearby
development of cool-temperate moist rain forest during the Late
Cretaceous of Antarctica, similar to the current rain forest to
heathland habitats in Central to South America (e.g., Dettmann,
1986; Askin, 1990a;Chart 2 of the supplementary online mate-
rial). Moreover, the discovery of several tetrads (Fig. 7L) of this
species and considering its resistance to long transport owing to its
heavy nature (due to its cingulate amb), these cool-temperate for-
ests were likely developed close to the shoreline.
In these forests, other “primitive”endemic communities of an-
giosperms such as Gunneraceae, Ericaceae, Casuarinaceae, Epacri-
daceae, Winteraceae and Trimeniaceae occurred adjacent to
estuaries or coal forming swamps at high southern paleolatitudes
(e.g., Dettmann and Jarzen, 1990; Hill and Scriven, 1995; Wagstaff
et al., 2006). Some of these angiosperms are recorded herein in
low frequencies (Fig. 3), such as the pollen of Palmae (e.g., Lil-
iacidites,Longapertites) that suggests warm to temperate conditions
in frost-free lowland areas. Palmae pollen was transported by rivers
close to the marine environment, as they are not usually trans-
ported far from their habitat (e.g., Edet and Nyong, 1993;Chart 2
supplementary online information). The occurrence of isolated
and fragmented ascocarps of epiphyllous fungi of Microthyriaceae
affinity in three specific levels (Fig. 3) of this section indicates a
warm and humid climate in the surrounding terrestrial environ-
ments (e.g., Duane, 1996; Kalgutkar and Braman, 2008 ). Hence, cool
to temperate and mainly humid environments were developed
during the deposition of The Naze assemblages, which is also
supported by several species of miospores, particularly fern spores
together with bryophyte and lycophyte spores. The consistently
low number of Araucariaceae and other taxa (Equisetosporites,
Ericaceae) throughout the section would represent more xero-
phytic environments or minor elements into those forests (Figs. 3,4
and Chart 2 supplementary online information).
7. Summary and conclusions
Sixteen palynoassemblages were studied from the Cape Lamb
Member of the Snow Hill Island Formation at The Naze where a
theropod dinosaur was recovered near the middle portion of a 90 m
thick section. The assemblages presented moderate diversity and a
total of 100 relatively well-preserved species. The main terrestrial
groups (32%) are represented by lycophytes (8 species), pterido-
phytes (15 species), gymnosperms (13 species), angiosperms (21
species) and freshwater chlorococcaleans (3 species). Marine
palynomorphs (68%) belong to dinoflagellates (61 species), chlor-
ococcaleans (6 species), and one acritarch (Michrystridium pilife-
rum). The vertical distribution of selected species allows the
distinction of two informal assemblages, the lower Odontochitina
porifera assemblage from the base to where it disappears in the
lower portion of the section and the upper Batiacasphaera grandis
assemblage from the highest occurrence of O. porifera to the top of
the exposed Cretaceous section.
The global stratigraphic ranges of selected palynomorphs sug-
gest an early Maastrichtian age that is also supported by the
presence of the ammonoid Kitchinites darwinii. These assemblages
share many species with latest Campanianeearly Maastrichtian
palynofloras from Vega and Humps Islands, New Zealand, and
elsewhere in the Southern Ocean, establishing good lateral corre-
lation (Fig. 2).
The Cape Lamb, Sanctuary Cliffs and Karlsen Cliffs members of
the Snow Hill Island Formation consist of a gradational coarsening
and thickening-upward succession of mudstones and sandstones
within the James Ross Basin and represent near-shore facies with
the overall section shallowing upward (e.g., Crame et al., 1991;
Pirrie et al., 1991, 1997). Olivero (2012) interpreted a deltaic
wedge that prograded during the time intervals encompassed by
Ammonite Assemblages 8e10 for about 80 km to the east, from
Santa Marta CoveeVega Island to Snow HilleSeymour Islands.
During the final regressive phases, the forced regressive strata of
I. cretaceum gravidum Mao and Mohr, CICYTTP-Pl 296-1: E32/4. C, I. cretaceum oviforme Mao and Mohr, CICYTTP-Pl 290-1: X51/3. D, Spinidium essoi Cookson and Eisenack, CICYTTP-
Pl 291-1: Y47/1. E, Isabelidinium pellucidum (Deflandre and Cookson) Lentin and Williams, CICYTTP-Pl 10-2: D27/0. F, Xenikoon australis Cookson and Eisenack, CICYTTP-Pl 286-3:
Z40/3. G, Amphidiadema nucula (Cookson and Eisenack) Lentin and Williams, CICYTTP-Pl 286-3: X49/4. H, Senegalinium bicavatum Jain and Millepied, CICYTTP-Pl 294-1: O30/4. I,
Paleocystodinium pilosum Guler et al., CICYTTP-Pl 289-1: O47/0. J, Chatangiella granulifera (Manum) Lentin and Williams, CICYTTP-Pl 286-1: Q49/1. K, Maduradinium pentagonum
Cookson and Eisenack, CICYTTP-Pl 286-1: O49/3. L, Nelsoniella aceras Cookson and Eisenack, CICYTTP-Pl 286-3: X43/2. M, Eurydinium ellipticum Mao and Mohr, CICYTTP-Pl 286-3:
Z56/3. N, Manumiella druggii (Stover) Bujak and Davies, CICYTTP-Pl 286-1: P55/0. O, Manumiella coronata (Stover) Bujak and Davies, CICYTTP-Pl 9-2: B37/3. P, Alterbidinium acutulum
(Wilson) Lentin and Williams, CICYTTP-Pl 290-1: M31/3. Q, Paleocystodinium pilosum Guler et al., CICYTTP-Pl 289-2: O44/1. ReT, Manumiella seelandica (Lange) Bujak and Davies. R,
CICYTTP-Pl 293-1: Y41/0. S, CICYTTP-Pl 295-1: H44/1. T, CICYTTP-Pl 289-1: S32/0. U, Phelodinium exilicornutum Smith, CICYTTP-Pl 296-1: B37/3. V, Cerodinium sp., CICYTTP-Pl 9-2:
Z46/3. WeX, Manumiella seymourensis Askin. W, CICYTTP-Pl 290-1: X20/1. X, CICYTTP-Pl 10-2: D48/0. Y, Canninginopsis bretonica Marshall, CICYTTP-Pl 9-2: S35/4. Z-AA, Manumiella
sp. cf. M. bertodano Thorn et al. Z, CICYTTP-Pl 8-5: M54/2. AA, CICYTTP-Pl 294-1: K54/4. BB, Canninginopsis ordospinosa Smith, CICYTTP-Pl 287-1: D34/0. CC, Palaeocystodinium
granulatum (Wilson) Lentin and Williams, CICYTTP-Pl 296-1: B32/0. DD, Saeptodinium gravattensis Harris, CICYTTP-Pl 291-1: T51/0.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4148

Fig. 10. Illustration of selected palynomorphs occurringwithin the lithostratigraphic section containingthe theropod dinosaur following the order in Fig. 4.Dinoflagellates, Gonyaulacales.
Scale bar is 15
m
mforFigs.AeD, 10
m
mforFigs.EeF, IeJ., 20
m
mforFigs.GeH, NeO, R, and 40
m
mforFigs.KeM, PeQ. A, Batiacasphaera rifensis Slimani et al., CICYTTP-Pl 295-1: D56/0. B,
Spongodinium reticulatum Hultberg, CICYTTP-Pl 290-1: X58/0. CeD, Batiacasphaera? reticulata (Davey) Davey. C, CICYTTP-Pl 291-1: X58. D, CICYTTP-Pl 291-1: O33/4. EeF, Impagidinium
cristatum(May) Lentin and Williams, CI CYTTP-Pl290- 1:Z3 9/4.G ,C anningia sp.1 Schiøler and Wil son, CICYTTP-Pl 286-1: H58/1. H, Batiacasphaera rugulataSch iøler and Wilson, CICYTTP-Pl 294-
1: V26/0. IeJ, Pterodinium cretaceum Slimani et al., CICYTTP-Pl 293-1: J39/2. KeL, Batiacasphaera grandis Roncaglia et al. K, CICYTTP-Pl 292-2: X25/0. L, CICYTTP-Pl 9-2: Z22/0. M, P, Q, Tri-
chodinium castanea (Deflandre) Clarke and Verdier eT. chilensisTroncoso andDoubinger. M, CICYTTP-Pl8-4: C29/4. P, CICYTTP-Pl290-1: J20/4.Q, CICYTTP-Pl51-3: V40/3. N, R,Odontochitina
porifera Cookson. N, CICYTTP-Pl 286-1: U42/2. R, CICYTTP-Pl 289-2: B24/0. O, Odontochitina indigena Marshall, CICYTTP-Pl 287-1: J28/1.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 149

Fig. 11. Illustration ofselectedpalynomorphsoccurringwithinthe lithostratigraphicsectioncontainingthe theropoddinosaur followingthe orderin Fig. 4. DinoflagellateseGonyaulacalesand
others. Scale baris 20
m
mforFigs.AeB, EeH,J, L, OeQ, 15
m
mforFigs.CeD, I, K, and 40
m
mforFigs.M,N.AeB, L, O, Spiniferitesramosus (ramosusemultibrevisegranosus) complex(Ehrenberg)
Mantell.A, S. brevis, CICYTTP-Pl 296-1: Z37/0. B, S.ramosus,CICYTTP-Pl8-4: C42/2.L, S. ramosus, CICYTTP-Pl 289-2: T49 (under fluorescence light). O,same specimenin L undertransmittedlight.
C, Kallosphaeridiumparvum Jan du Chene in Sli mani et al., CICYTTP-Pl 292-2: L21/4. D, Kallosphaeridium? ringnesoriumin Mohr and Ma o, CICYTTP-Pl 289-2: K37/4. E, Spiniferites pseudofurcatus
(Klump) Sarjeant, CICYTTP-Pl 287-1: Z46/3. F, Operculodiniumradiculatum Smith, CICYTTP-Pl293-1:W32/3. G, Xenascus ceratioides(Deflandre) Davey and Verdier. G, CICYTTP-Pl 296-1: W39/3.
H, J, Xenascusplotei Below. H, CICYTTP-Pl 10-2:V 45/3.J, CI CYTTP-Pl296- 1: W28/0. I, Operculodinium centrocarpum (Deflandre and Cookson) Wall,CI CYTTP-Pl 289-3: Z30/3. M, Arthropod egg,
CI CYTTP-P l 51-3: T51. L, Membranilarnacia densaCookson and Eisenack,CICYTTP-Pl288-2: S55/4.N, Cuticle of Bennettitaleanaffinity,CICYTTP-Pl 296-1: K25/2. PeQ, Isabelidinium pellucidum
(Deflandreand Cookson) Lentin and Williams. P, CICYTTP-Pl 289-2: U28/0 (under fluorescencelight). Q, same specimen in P under transmitted light.
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4150

the Haslum Crag Sandstone suggest that the shoreline was at that
time located near Seymour Island (Fig. 2; see also Olivero et al.,
2008). Smith (1992) interpreted that the Cape Lamb Member at
Vega Island (lowest Maastrichtian) was deposited in a low-energy
shelf setting, below storm wave base. The inferred sea-level curve
presented by Olivero (2012) for this NG sequence shows for the late
Campanian Ammonite Assemblages (AA) 8 a progressive trans-
gressive trend and a regressive trend from the AA 9 to AA 10.
Following the sequence stratigraphic framework of Olivero
(2012), our palynoassemblage along the 90 m section of the Cape
Lamb Member (Fig. 3) equates to the 8e9 Ma NG Sequence (see also
Crame et al., 2004) and should correspond to its Ammonite
Assemblage 10 of earliest Maastrichtian age based on the concur-
rent ranges of ammonoids and palynomorphs documented in the
studied section (see Figs. 2, 3 and 6). The Naze section was probably
deposited during 1e2 Ma (4th or 5th order in Miall,1996), in a low-
energy shelf setting, based on the lithological composition, thick-
ness of the section, and the palynological information discussed
above. The dominance or frequent presence of dinoflagellates
throughout the section supports the general interpretation of a
shelf marine depocenter. The quali-quantitative variations of di-
noflagellates (Figs. 3e5)reflect environmental changes, mainly
shoreline shifts that influenced the proximaledistal trend of the
permanent marine depocenter at The Naze. The inferred sea level
curve based on palynological and other paleontological evidence
along the section, points to an important role of local sea level
changes in controlling the abundance and composition of vegeta-
tion probably largely due to the amount of coastal plain available
for colonization. These small-scale temporal changes could be
related to Milankovitch cycles in a regional and larger scale cooling
and regressive event that would have occurred from the end of the
Campanian through the early Maastrichtian.
Based on The Naze section (see Fig. 3), the terrestrial environ-
ments surrounding this depocenter were represented by podo-
carpeNothofagus rainforests established mainly in lowlands with
other angiosperm groups and ferns developed in understories,
under highly humid and temperate to cool-temperate conditions.
Lycophytes and bryophytes and subordinated elements (e.g., Pro-
teaceae, Battenipollis, Liliaceae, Palmae, Microthyriaceae, and other
herbeshrub dicotyledonous species), likely endemic (see Askin,
1989), were mostly related to the vegetation along riparian flood-
plains, swamps (with Stereisporites and Lycopodiales) and lakes
(with Chlorophytes and fresh water dinoflagellates Saeptodinium)
that were fringing estuaries with brackish waters (Michrystridium,
some Chlorophytes and few peridinalean species) to the shallow
marine shelf (the permanent depocenter). These communities
were developed under cool-temperate, frost-free and high-rainfall
conditions (Dettmann, 1986, 1989; Askin, 1990a, 1990b; Jarzen and
Dettmann, 1992; Crame, 1992; Hill and Scriven,1995). Well-defined
growth rings within fossil wood samples (Araucarioxylon Krausel)
recovered from Lachman Crags and The Naze (James Ross Island)
showed that this climate was markedly seasonal (Francis, 1986,
1991 ).
Bowman et al. (2012) presented a paleobiogeographical synthesis
including a proposal of a South Polar Province characteristic of cool
waters embracing northwestern Antarctica (James Ross Basin), the
East TasmanPlateau, New Zealand and southern South America and
Australia forthe late Maastrichtian to earliest Paleocene. The marine
endemic palynomorphs in our assemblages (e.g., Manumiella sey-
mourensis,Operculodinium exilicornutum,Batiacasphaera grandis)
confirm the inclusion of this area in this phytocore during the early
Maastrichtian. Some marine cosmopolitan dinoflagellate species
here recorded (e.g., Chatangiella granulifera,Isabellidinium pelluci-
dum,Isabellidinium cooksoniae,Xenascus plotei,Xenascus ceratioides,
Spiniferites ramosus,Operculodinium centrocarpum) indicate a
relatively free connection of the western regions along the Paleo-
pacific Ocean to other regions of the World. This is also reinforced by
the record of Odontochitina porifera in the Campanian of southern-
most Brazil (Premaor et al., 2010), and by Chatangiella granulifera
recorded in the basal part of the The Naze section (Fig. 4), which has
been suggested to be characteristic of cold waters of the North Polar
region (Canadian Arctic and Siberia) during the latest Cretaceous
(Harker et al., 1990).
Acknowledgments
The authors would like to sincerely thank the Instituto Antárc-
tico Argentino and the National Science Foundation, Office of Polar
Programs, who were responsible for the support of the expeditions
(NSF grants awarded to J.E. Martin (OPP#0087972) and J.A. Case
(OPP#0003844)). This research would not have been possible
without the collaboration from Dr. Judd A. Case, Eastern Wash-
ington University. Dr. J. Foster Sawyer (SD School of Mines and
Technology), collaborated in measuring and sampling the section in
Antarctica. M. Di Pasquo acknowledges the Fulbright Scholarship
Program and CONICET of Argentina for providing her the oppor-
tunity for cooperation with colleagues from USA and for continuing
collaboration. Support from home institutions is greatly appreci-
ated (e.g. ANPCyT Pict 07-36166). Captain Mike Terminal and crew
of the Lawrence M. Gould research vessel are thanked for their
congeniality and for transportation to James Ross Island. Mr. John
Evans and his staff in Raytheon Polar made every effort to make our
sojourn possible, safe, and comfortable. We thank all our field
companions, Dr. Marcelo Reguero, Dr. J. Foster Sawyer, Dr. Allen J.
Kihm, Dr. Jennifer Hargrave, Dr. Robert Meredith, Ms. Amanda
Person, Dr. Wayne Thompson, Ms. Melissa Rider, Dr. Kristin van
Konynenburg, Mr. Joe Pettit, Mr. Dan Martin, and Ms. Lucy Bledsoe,
whose dedication to science and congeniality during extremely
difficult field conditions are the cornerstone of this research. Dr.
Wayne Thompson aided further in the sketch of the outcrop for
which we are thankful. The authors thank to the anonimous re-
viewers and the Editor E.A.M. Koutsoukos for their suggestions that
allowed us to improve our manuscript.
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Appendix A. Supplementary data
Supplementary data related to this article can be found at http://dx.doi.org/10.
1016/j.cretres.2013.07.008.
Appendix B
List of taxa documented in the Snow Hill Island Formation at The Naze, arranged
by major groups in alphabetical order. The reference of illustrated specimens in
Figs. 7e11 (mostly following the order in Fig. 4) is included. See full authority in
Fig. 4.
Spores
Baculatisporites comaumensis (Cookson) Potonié (Fig. 7B)
Biretisporites potoniaei Delcourt and Sprumont (Fig. 7C)
Calamospora sp.
Camarozonosporites ohioensis (Couper) Dettmann and Playford
Ceratosporites equalis Cookson and Dettmann (Fig. 7J)
Cyatheacidites annulatus Cookson (Fig. 7L)
Cyatheacidites archangelskii Dettmann (Fig. 7E)
Cyathidites australis/C. minor Couper
Densoisporites velatus Weyland and Krieger (Fig. 7G)
Dictyotosporites speciosus Cookson and Dettmann
Gleicheniidites senonicus Ross
Herkosporites sp.
Ischyosporites volkheimeri Filatoff (Fig. 7H)
cf. Klukisporites scaberis (Cookson and Dettmann) Dettmann
Laevigatosporites ovatus Wilson and Webster/L. major (Cookson) Krutzsch (Fig. 7K)
Lycopodiumsporites eminulus Dettmann
Peromonolites bowenii Couper (Fig. 7Q)
Perotrilites majus (Cookson and Dettmann) Evans (Fig. 7A)
Polypodiisporites favus (Potonié) Potonié
Polypodiisporites sp. (Fig. 7I)
Retitriletes austroclavatidites (Cookson) Döring et al. in Krutzsch (Fig. 7D)
Stereisporites antiquasporites (Wilson and Webster) Dettmann
Stereisporites regium (Drozhastichich) Drugg (Fig. 7F)
Todisporites major/T. minor Couper
Trilites parvallatus Krutzsch
Triporoletes radiatus (Dettmann) Playford (Fig. 7N)
Tuberculatosporites parvus Archangelsky
Pollen
Gymnosperm
Araucariacites australis Cookson (Fig. 7U)
Dacrycarpites australiensis Cookson and Pike
Dilwynites granulatus Harris (Fig. 7P)
Equisetosporites sp. (Fig. 7W)
Microcachrydites antarcticus Cookson ex Couper (Fig. 8A)
Phyllocladidites mawsonii Cookson ex Couper (Fig. 7V)
Podocarpidites major Couper (Fig. 7T)
Podocarpidites marwickii Couper
Podocarpidites otagoensis Couper (Fig. 7M)
Podocarpidites rugulatus Pocknall and Mildenhall (Fig. 7O, S)
Podocarpidites verrucosus Volkheimer
Podocarpidites spp.
Trichotomosulcites subgranulatus Couper (Fig. 8O)
Angiosperm
Battenipollis sectilis (Stover) Jarzen and Dettmann (Fig. 8B)
Ericipites scabratus Harris (Fig. 8K)
Liliacidites spp. (Fig. 8I)
Longapertites sp. Povilauskas et al. (Fig. 7R)
Monosulcites palisadus Couper
Monosulcites/Arecipites spp.
Myricipites harrisii (Couper) Dutta and Sah (Fig. 8G)
Nothofagidites americanus Zamaloa (Fig. 8J)
Nothofagidites dorotensis Romero
Nothofagidites saraensis Menéndez and Caccavari de Fílice (Fig. 8P)
Nothofagidites tehuelchesii Zamaloa and Barreda
Nothofagidites spp.
Peninsulapollis askiniae Dettmann and Jarzen (Fig. 8H)
Peninsulapollis gillii (Cookson) Dettmann and Jarzen (Fig. 8F)
Periporopollenites polyoratus (Couper) Stover in Stover and Partridge (Fig. 8L)
Proteacidites parvus Cookson (Fig. 8N)
Proteacidites scaboratus Couper (Fig. 8C)
Proteacidites tenuiexinus Stover in Stover and Partridge (Fig. 8E)
Proteacidites spp.
Tricolpites confessus Stover in Stover and Partridge (Fig. 8D)
Triorites orbiculatus McIntyre (Fig. 8M)
Dinoflagellate
Alterbidinium acutulum (Wilson) Lentin and Williams emend. Khowaja-
Ateequzzaman, Garg and Jain (Fig. 9P)
Amphidiadema nucula (Cookson and Eisenack) Lentin and Williams (Fig. 9G)
Andalusiella mauthei Riëgel
Batiacasphaera grandis Roncaglia et al. (Fig. 10K, L)
Batiacasphaera rifensis Slimani et al. (Fig. 9A)
Batiacasphaera rugulata Schiøler and Wilson (Fig. 10H)
Batiacasphaera? reticulata (Davey) Davey (Fig. 9C, D)
Canningia sp. 1 in Schiøler and Wilson (Fig. 10G)
Canninginopsis bretonica Marshall (Fig. 9Y)
Canninginopsis ordospinosa Smith (Fig. 9BB)
Cerodinium (“Ceratiopsis”) sp. in Mehrotra and Sarjeant (1987)
Chatangiella granulifera/verrucosa (Manum) Lentin and Williams (Fig. 9J)
Chatangiella victoriensis (Cookson and Manum) Lentin and Williams
Circulodinium distinctum (Deflandre and Cookson) Jansonius
Cyclonephelium compactum Deflandre and Cookson
Diconodinium cristatum Cookson and Eisenack emend Morgan
Diconodinium sp. cf. multispinum (Deflandre and Cookson) Eisenak and Cookson
emend Morgan
Eurydinium ellipticum Mao and Mohr (Fig. 9M)
Impagidinium cristatum (May) Lentin and Williams (Fig. 10E, F)
Isabellidinium cooksoniae (Alberti) Lentin and Williams
Isabellidinium cretaceum (Cookson) Lentin and Williams (Fig. 9A)
Isabelidinium cretaceum gravidum Mao and Mohr (Fig. 9B)
Isabellidinium cretaceum oviforme Mao and Mohr (Fig. 9C)
Isabelidinium pellucidum (Deflandre and Cookson) Lentin and Williams (Fig. 9E,
Fig. 11P, Q)
Isabellidinium papillum Sumner
Isabellidinium sp.
Kallosphaeridium parvum Jan du Chêne (Fig. 11C)
Kallosphaeridium? ringnesorium in Mohr and Mao (Fig. 11D)
Maduradinium pentagonum Cookson and Eisenack (Fig. 9K)
Manumiella conorata (Stover) Bujak and Davies (Fig. 9O)
Manumiella druggii (Stover) Bujak and Davies (Fig. 9N)
Manumiella seelandica (Lange) Bujak and Davies (Fig. 9R, T)
Manumiella seymourensis Askin (Fig. 9W, X)
Manumiella sp. cf. M. bertodano Thorn et al. (Fig. 9Z, AA)
Manumiella spp.
Membranilarnacia densa Cookson and Eisenack (Fig. 11L)
Membranilarnacia spp.
Nelsoniella aceras Cookson and Eisenack (Fig. 9L)
Nelsoniella tuberculata Cookson and Eisenack
Odontochitina indigena Marshall (Fig. 10O)
Odontochitina porifera Cookson (Fig. 10N, R)
Operculodinium centrocarpum (Deflandre and Cookson) Wall (Fig. 11I)
Operculodinium flucturum Davey
Operculodinium radiculatum Smith (Fig. 11F)
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4 153

Palaeocystodinium granulatum (Wilson) Lentin and Williams (Fig. 9CC)
Paleocystodinium golzowense Alberti
Paleocystodinium pilosum Guler et al. (Fig. 9I, Q)
Phelodinium exilicornutum Smith (Fig. 9U)
Pterodinium cretaceum Slimani et al. (Fig. 10I, J)
Saeptodinium gravattensis Harris (Fig. 9DD)
Senegalinium bicavatum Jain and Millepied (Fig. 9H)
Spinidium essoi Cookson and Eisenack (Fig. 9D)
Spiniferites pseudofurcatus (Klump) Sarjeant (Fig. 11E)
Spiniferites ramosus (ramosusemultibrevisegranosus) complex (S. ramosus (Ehren-
berg) Mantell) (Fig. 11A, B, L, O)
Spongodinium reticulatum Hultberg (Fig. 10B)
Trichodinium castaneum (Deflandre) Clarke and Verdier eT. chilensis Troncoso and
Doubinger, complex (Fig. 10M, P, Q)
Valensiella reticulata (Davey) Courtinat
Xenascus ceratioides (Deflandre) Davey and Verdier (Fig. 11G)
Xenascus plotei Below (Fig. 11H, J)
Xenikoon australis Cookson and Eisenack (Fig. 9F)
Chlorophyceae
Botryococcus brauni Kützing (Fig. 8Q)
Dictyotidium sp.
Leiosphaeridia sp.
Nummus monoculatus Morgan (Fig. 8R, U)
Palambages spp. (Fig. 8X)
Paralecaniella indentata (Deflandre and Cookson) Cookson and Eisenack emend Elsik
(Fig. 8S)
Pterospermella australensis (Deflandre and Cookson) Eisenack
Pterospermella aureolata (Deflandre and Cookson) Eisenack (Fig. 8V)
Pterospermella spp.
Tetraedron cf. minimum (A. Braun) Hansgirg (Fig. 8T)
Acritarchs
Michrystridium piliferum Deflandre (Fig. 8W)
Fungi, Microthyriaceae
Asterothyrites Cookson (Fig. 8Y)
Organic foraminiferal linings (Fig. 8Z, AA)
Arthropod (Copepods) eggs (Fig. 11M)
M. di Pasquo, J.E. Martin / Cretaceous Research 45 (2013) 135e15 4154