Cretaceous Research 25 (2004) 211–228
www.elsevier.com/locate/CretRes
Late Cretaceous plant mesofossils from
Table Nunatak, Antarctica
Helena Eklunda,), David J. Cantrillb, Jane E. Francisa
b
a
School of Earth Sciences, University of Leeds, Leeds LS2 9JT, UK
Department of Palaeobotany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden
Received 11 October 2002; revised 27 October 2003; accepted in revised form 19 November 2003
Abstract
Charred and structurally preserved plant remains have been recovered from Late Cretaceous strata from Table Nunatak,
Antarctica. The mesofossils, up to a few mm in length, represent structures from lycopods (megaspores), ferns (circinate rachides),
conifers (wood, leaves, leafy shoots, pollen cones, microsporophylls, ovulate cone scales, seeds), and angiosperms (leaves, flowers,
fruits, seeds). This work focuses on the megaspores, microsporophylls, ovulate cone scales and vegetative structures. The mesofossils
were extracted from unconsolidated sediments deposited in a shallow-marine environment during the late Santonian (ca. 83 Ma). The
Table Nunatak assemblage is the only known Cretaceous mesofossil assemblage with charred plant parts from Antarctica, and one of
only a few from the Southern Hemisphere as a whole. It represents the remains of forests that grew at palaeolatitudes of 65(S, during
a time in the Cretaceous in which high latitude environments experienced warm temperate climates in a global greenhouse world.
Ó 2004 Elsevier Ltd. All rights reserved.
Keywords: Angiosperms; Antarctica; Charcoal; Conifers; Ferns; Late Cretaceous; Lycopods; Plant mesofossils
1. Introduction
The preservation of plant parts as charcoal, which is
the result of wildfires, occurred on a large scale during
the Carboniferous, Jurassic and Cretaceous (Scott, 1989,
2000; Scott and Jones, 1991a). As shown experimentally
by Scott and Jones (1991a,b) plant parts can turn into
charcoal if they are subjected to sufficiently high
temperatures in an anoxic environment. Although the
charring process results in shrinkage of the organic
material and causes some morphological changes (Scott
and Jones, 1991b; Lupia, 1995; Scott, 2000), charcoalified plant fossils often show excellent preservation of
anatomical and morphological details. The reason for
this remarkable fossilisation potential is that charcoal,
which is almost pure carbon, is chemically highly inactive and remains three-dimensional unless subjected
to physical compression (Scott and Jones, 1991a).
Consequently charcoalified fossils often reveal pertinent
) Corresponding author.
E-mail address: [email protected] (H. Eklund).
0195-6671/$ - see front matter Ó 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cretres.2003.11.004
information for reconstructing the plant’s original morphology and anatomy.
Most of the plant parts preserved in this manner are
fragments of charcoalified secondary wood (Scott and
Jones, 1991a). However, records of charcoalified fossils
include a variety of plant parts of different texture and
delicacy, derived from a wide range of plant groups.
Charred structures from mosses, lycopods, ferns, conifers and angiosperms are known and include sporophytes, gametophytes, megaspores, microsporangia,
wood, leafy shoots, isolated leaves, cone fragments, inflorescence axes, isolated flowers, stamens, fruits, and
seeds (Friis and Skarby, 1981; Batten et al., 1984; Friis,
1984; Scott, 1989, 2000; Scott and Jones, 1991b; Crane
et al., 1994; Crepet and Nixon, 1994; Friis et al., 1994;
Herendeen et al., 1999; Takahashi et al., 1999a,b;
Gandolfo et al., 2001).
One field within botany/palaeobotany in which
information from charcoalified fossils has had a big
impact is the study of early angiosperms. Ever since the
novel and unexpected discovery of excellently preserved
flowers from the Late Cretaceous of Sweden early in the
212
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
1980s (Friis and Skarby, 1981; Friis, 1985), palaeobotanists have searched for additional Cretaceous
deposits of charcoal. As a consequence several Cretaceous formations containing charcoalified plant structures have been discovered, and a large diversity of small
charred fossils (so called mesofossils) have been extracted from these using bulk-sieving techniques. Today
mesofossil assemblages, containing charcoalified structures of angiosperms and other plant groups, are known
from widely separated localities in Europe, Asia and
North America. In Europe, mesofossil floras have been
reported from Sweden (Friis and Skarby, 1981; Eklund
et al., 1997; Schönenberger and Friis, 2001; Schönenberger et al., 2001a,b; and references therein), Portugal
(Friis et al., 2001; and references therein), and the Czech
Republic (Eklund and Kvaček, 1998); in Asia from
Kazakstan (Frumina et al., 1995; Frumin and Friis,
1996, 1999) and Japan (Takahashi et al., 1999a,b). These
types of floras are also widespread in North America
(Herendeen et al., 1999; Eklund, 2000; Gandolfo et al.,
2000, 2001; Magallón et al., 2001; and references
therein).
The majority of mesofossil sites occur in the Northern Hemisphere, strongly biasing our view of global
floristic composition. In the Southern Hemisphere Cretaceous mesofossils have been described from the Lower
Cretaceous in Australia (Tosolini, 2000). In this paper
we present descriptions of charcoalified mesofossils
from the Upper Cretaceous Table Nunatak Formation,
Antarctica (Fig. 1). To our knowledge, Table Nunatak
is the only formation in Antarctica that has yielded a
Cretaceous mesofossil flora. In general, the mesofossils
from Table Nunatak are less well-preserved than many
of the contemporary mesofossils from the Northern
Hemisphere. In many cases there is insufficient in-
formation available to enable systematic identification.
However, we agree with Herendeen et al. (1999) that
such fossils may, nevertheless, be important for systematic, palaeoecological and biogeographic comparisons
with other mesofossil assemblages. For these reasons we
provide an overview of the Table Nunatak assemblage,
which includes structures from lycopods (megaspores),
ferns (megaspores, circinate rachides), conifers (wood,
leaves, leafy shoots, pollen cones, microsporophylls,
ovulate cone scales, seeds), and angiosperms (leaves,
flowers, fruits, seeds). Here we present a selection of
these mesofossils, focusing on megaspores and vegetative structures. Although conifer microsporophylls and
ovulate cone scales are covered in this paper, the majority of reproductive structures (conifer seeds, angiosperm
flowers, fruits, seeds) will be described separately
(Eklund, 2003, work in progress).
2. Material and methods
The charcoalified plant fossils described here were
obtained by wet sieving of unconsolidated sediments
collected by D. I. M. Macdonald (1984–85; samples
R.2959.27, R.2959.8) and B. Hathway (1996; sample
DJ.755.1) at Table Nunatak, just east of Kenyon
Peninsula on the eastern side of the Antarctic Peninsula
(68(27#S, 62(43#W) (Fig. 1). Table Nunatak is an
isolated outcrop measuring 1 km long and 400 m wide,
with the long axis orientated WNW–ESE and the
relatively flat top tilted gently NE. The steep margins
of the nunatak are exposed from 10 to 30 m above the
level of the surrounding ice (Macdonald, 1985). The
formation has a total thickness of 62 m and the plantbearing strata are exposed on both the southwestern and
Fig. 1. Map showing the location of Table Nunatak at the end of Kenyon Peninsula on the eastern margin of the Antarctic Peninsula (redrawn from
Hathway et al., 1998 and Poole and Gottwald, 2001).
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
the northeastern sides of the nunatak (Hathway et al.,
1998). The material studied here was extracted from
sites on the southwestern side of the nunatak.
Three sedimentary facies have been recognised within
the geological sequence at Table Nunatak: Facies A consisting of fine- to very fine-grained, thick- to very thickbedded sandstone; Facies B consisting of plant-rich,
fine- to very fine-grained sandstone and siltstone; and
Facies C consisting of massive bioturbated mudstone
(Hathway et al., 1998). The plant-fossil-bearing Facies B
is poorly lithified and accounts for only 2–3% of the total
measured section. It consists of layers of charcoal in a silt
matrix and paler, cleaner, very fine sandstone. The layers
of plant debris are 0.1–0.5 cm thick and include wood
fragments that are up to 25 cm long. Small twigs, leaves,
fruits, seeds, and rare flowers also occur in these layers.
The succession is interpreted as representing shallowmarine deposition at the mouth of an estuary or a deltaic
distributary channel, and has been dated, using dinoflagellate cysts, as late Santonian (Hathway et al., 1998).
The palaeolatitude for Table Nunatak is considered to
have been approximately 65( South at this time (Lawver
et al., 1992).
The plant fossils are preserved as three-dimensional,
sometimes slightly flattened, charcoalifications. They
were extracted from the sediment matrix by soaking the
sediments in tap water and washing them through
a series of sieves. The fossils were subsequently cleaned
of adhering minerals by treatment with hydrochloric
and hydrofluoric acids, rinsed in tap water and air-dried.
The material was picked under the dissecting microscope, and selected fossils were documented with line
drawings and mounted on 0.5$ aluminium specimen
stubs using transparent nail varnish. They were subsequently coated with gold for 8 min using an Emscope
SC500 and examined and photographed using a SEM
CamScan-4 (Image Slave). All specimens are stored in
the collections of the British Antarctic Survey (BAS),
Cambridge, UK. Sample and specimen numbers refer to
the BAS catalogue system.
3. Systematic palaeobotany
213
polar view (Fig. 2A) and equatorial view (Fig. 2C). In
both specimens the trilete mark has split into a distinct
triradiate suture with arms extending beyond the
equator (Fig. 2A, C). The surface of the megaspores
shows numerous gemmate bodies of variable sizes; these
are most abundant in distal regions (Fig. 2A–C).
Remarks. This type is represented by two specimens
(R.2959.8.1, DJ.755.1.85; Fig. 2A–C) that are referred
to Cabochonicus Batten and Ferguson on the basis of the
glossy resinous gemmate elements on an otherwise
smooth surface (Batten and Ferguson, 1987; Batten,
1988). There has been some discussion whether the
globular bodies represent sculptural elements of an unusual appearance or are products of fungal attack (e.g.,
Marcinkiewicz, 1979; Batten et al., 1984; Batten and
Ferguson, 1987). The systematic affinity of Cabochonicus is not clear, but it is possible that these spores were
produced by plants related to Isoetales (A. R. Hemsley,
pers. comm. 2001) or Selaginellales (Batten, 1988).
Megaspore type 2
Description. Megaspores ranging from 860 to 940 mm
in diameter. All specimens are more or less globular.
The surface shows a rugose sculpturing formed by short
and undulating to irregularly arranged muri (Fig. 2D).
The trilete ridges are indistinct and the orientation of the
poles is therefore unclear. The spores have an abraded
appearance and the surface ornamentation may have
been more elaborate originally.
Remarks. Megaspore type 2 is represented by nine
specimens (e.g., DJ.755.1.45; Fig. 2D) that are similar to
Rugotriletes van der Hammen and possibly Erlansonisporites Potonié (1956), both of which are thought to be
related to extant Selaginellaceae (Collinson et al., 1985;
Batten, 1988). Both Rugotriletes and Erlansonisporites
have rugulate to imperfectly reticulate surface sculpturing formed by robust and irregularly distributed muri
(Potonié, 1956; Batten, 1988) similar to the material
described here. Although both genera have similar wall
structure the ornamentation differs but overlaps. Both
genera can be separated on size classes.
3.1. Lycophytes
Erlansonisporites sp. 1
Remarks. Lycophytes are relatively rare in the Table
Nunatak assemblage and are represented by four types
of isolated megaspores (Fig. 2A–F). Some megaspores,
such as specimens of type 4 below, tend to be lighter in
colour than the other megaspores. Other megaspores are
darker and may be reworked from older deposits.
Cabochonicus sp.1
Description. Megaspores approximately 390 to
440 mm in diameter, and with a more or less circular
outline in proximal polar view (Fig. 2E). The surface
ornamentation consists of relatively high and undulating
muri. At the proximal pole the trilete mark is not very
prominent, but can be discernible as three narrow and
slightly raised arms (Fig. 2E).
Description. Megaspores ranging from 750 mm to
1 mm in diameter, more or less circular in proximal
Remarks. Erlansonisporites sp. 1 is represented by a
single specimen (DJ755.1.131; Fig. 2E), which conforms
214
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Fig. 2. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): lycopod megaspores. A–C, Cabochonicus sp. 1. A,
proximal polar view showing triradiate suture and numerous globose bodies in distal regions (R.2959.8.1). B, close-up of globose bodies of
Cabochonicus sp. 1 (R.2959.8.1). C, equatorial view; note that the trilete mark has split into a distinct triradiate suture with arms extending beyond
the equator (DJ.755.1.85). D, megaspore type 2, probable selaginellaceous megaspore similar to fossil Rugotriletes van der Hammen and
Erlansonisporites Potonié (DJ.755.1.45). E, Erlansonisporites sp. 1, proximal polar view showing indistinct trilete mark discernible as three narrow
and slightly raised arms (DJ755.1.131). F, Megaspore type 4; note the three broad and rounded ridges that indicate the position of the trilete mark
(R.2959.27.26). G, H, Hughesisporites sp. 1. G, proximal view (R.2959.8.52). H, close-up of Hughesisporites sp. 1 showing radially arranged sinuous
muri (R.2959.8.52). Scale bars in A, C–G represent 200 mm; in B, H, 50 mm.
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
to this genus as it has an inconspicuous trilete mark that
is obscured by the muri. It is somewhat similar to
megaspore type 2, and it cannot be excluded that
Erlansonisporites sp. 1 represents less abraded specimens
of megaspore type 2. Erlansonisporites sp. 1 is, however,
about half the size of those of type 2 and for this reason
these megaspores are treated as separate types here.
Nevertheless, considering the low number of specimens
observed of each type it is possible that both types are
actually within the natural size range of a single taxon.
Another reason for separating the two megaspore types is
based on an observation by Takahashi et al. (1999b) that
the trilete mark is clearly visible in abraded specimens
but unclear in unabraded specimens of their megaspore
type 1, which probably has affinity to Erlansonisporites
erlansonii (Miner) Potonié. In the case of megaspore type
2 and Erlansonisporites sp. 1 from Table Nunatak the
opposite would be the case; i.e., the trilete mark is more
prominent in the less abraded specimens (Erlansonisporites sp. 1) than in the abraded (type 2).
Megaspore type 4
Description. In equatorial view the megaspores are
rounded triangular to subglobose and about 1.5 mm
wide and 814 mm long. All megaspores are strongly
abraded and the surface is glabrous and more or less
smooth to slightly rugulose (Fig. 2F). Three broad and
rounded ridges indicate the position of the trilete mark
(see arrows, Fig. 2F).
Remarks. Megaspore type 4 is represented by about
20 specimens (e.g., R.2959.27.26; Fig. 2F). It is possible
that megaspore type 4 represents strongly abraded
specimens of megaspore type 2, which is similar in size,
or the smaller megaspores of Erlansonisporites sp. 1.
However, megaspore type 4 differs from type 2 and
Erlansonisporites sp. 1 in the general shape and in the
trilete mark whose arms are broad with a rounded
profile.
Hughesisporites sp. 1
Description. Megaspore 580–620 mm in diameter and
with a more or less circular outline in proximal polar
view (Fig. 2G). The surface ornamentation at the proximal face consists of radially arranged sinuous muri
(Fig. 2H). The surface is smooth equatorially and
distally. The trilete mark is inconspicuous.
Remarks. Hughesisporites sp. 1 is represented by
a single specimen (R.2959.8.52; Fig. 2G, H). In gross
morphology and ornamentation this spore is best placed
in Hughesisporites Potonié, a Triassic to Early Cretaceous megaspore genus of unknown affinity (Kovach
and Batten, 1989; Koppelhus and Batten, 1992).
215
3.2. Pteridophytes
Remarks. The presence of ferns in the Table Nunatak
flora is evidenced by the occurrence of Arcellites sp.
megaspores (Hathway et al., 1998; Fig. 3A–D this work)
and small fragments of circinate fronds (Fig. 3E, F). Of
the fronds, one specimen shows distinct pinnules not
seen in the other specimens. It is not known whether this
is due to the preservation or if it is a taxonomic feature.
Arcellites cf. A. reticulatus (Cookson and Dettmann)
Potter
Description. A single intact megaspore that is 572 mm
long by 317 mm wide. Spore differentiated into a distinct
spore body (241 mm long) and a well-developed apical
extension (331 mm long) (Fig. 3A). The spore body is
more or less circular in outline and the surface is covered
by short tubular appendages that are basally joined to
form a compact reticulum (Fig. 3C, D). The apical
extension, which is differentiated into a proximal neck
and a distal head, consists of an unknown number of
leaf-like folded appendages (Fig. 3A, B). The laesurae
are not clearly visible.
Remarks. Arcellites Miner (1935), which is a form
genus for Cretaceous fern megaspores, is represented
here by a few specimens, of which a single is completely
preserved (R.2959.8.51; Fig. 3A–D; see also Hathway
et al., 1998). Approximately 29 species of Arcellites are
known from the Cretaceous but many are characterised
by well-developed tubular appendages and lack a basal
reticulum. A well-developed reticulum with short
tubular appendages is seen in a few species [e.g., Arcellites
plicatus Li and Batten, 1986, A. reticulatus (Cookson
and Dettmann) Potter]. The material described here is
more like A. reticulatus as it lacks transversely plicate
ornamentation on the neck (Li and Batten, 1986).
Megaspores of Arcellites are known only from the
Cretaceous, and are thought to have been produced by
ferns growing in waterlogged or aquatic habitats with
affinities to extant Marsileaceae (e.g., Ellis and Tschudy,
1964; Li and Batten, 1986; Batten et al., 1996; Tosolini,
2000). Extant Marsileaceae includes the three genera
Marsilea L., Regnellidium Lindman and Pilularia L.
Interestingly, the leaves of Pilularia lack a lamina,
consisting only of a filiform petiole which is circinate in
bud (Kramer, 1990) and it is possible that the rachids of
?fern type 1 described below were produced by the
Arcellites plant.
Circinate fern rachis type 1
Description. The rachis is about 1.3 mm long. It has
a circinate vernation and lacks lateral pinnules.
Remarks. Circinate fern rachis type 1 is represented
by five specimens (e.g., R.2959.8.1; Fig. 3E). The
216
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Fig. 3. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): megaspore (A–D) and circinate rachids (E, F) of ferns. A,
probable marsileaceous megaspore, similar to fossil Arcellites Miner; equatorial view showing differentiation into distinct spore body and welldeveloped apical extension (R.2959.8.51). B, close-up of apical extension of marsileaceous megaspore in A; note leaf-like folded appendages
(R.2959.8.51). C, close-up of spore body of marsileaceous megaspore in A showing compact reticulum formed by short tubular appendages that are
joined basally (R.2959.8.51). D, close-up of spore body in C showing tubular appendages at higher magnification (R.2959.8.51). E, circinate fern
rachis ?type 1 (R.2959.8.1). F, circinate fern rachis ?type 2; note 12 or 13 pairs of lateral pinnules (DJ.755.1.62). Scale bars in A represent 200 mm; in
B, C, 50 mm; in D, 10 mm; in E, F, 0.5 mm.
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
systematic affinity of circinate fern rachis ?type 1 is uncertain. However, the lack of lateral pinnules is a
feature of ferns such as Pilularia and coupled with the
presence of spores allied to the Marsileaceae (Arcellites)
it is possible that that this material belongs to the
Marsileaceae.
217
the axis (Fig. 4A, B). The protuberances extend about
0.15–0.17 mm from the axis (Fig. 4B, C).
Remarks. This taxon is represented by a single specimen (DJ.755.1.62; Fig. 3F). The more precise systematic affinity of circinate fern rachis ?type 2 is unknown,
but it clearly differs from type described above.
Remarks. This single specimen is interpreted as an
axis of a pollen cone (DJ.755.1.22; Fig. 4A–C) based on
the wood nature of the axis and spiral insertion of the
organs. It is similar to material illustrated by Cantrill
et al. (1995) where a few microsporophylls remained
intact on the otherwise denuded axis. Although the
specimen can easily be interpreted as a pollen cone
axis, the lack of complete microsporophylls precludes
accurate taxonomic placement. However, based on
the number of microsporophylls, their spacing on the
axis it is more likely that this material is allied to
the Podocarpaceae or Taxodicaceae rather than the
Araucariaceae.
3.3. Conifers
Conifer microsporophyll
Circinate fern rachis ?type 2
Description. The rachis is 1.1 mm long. It has a
circinate vernation and 12 or 13 pairs of lateral pinnules.
Remarks. Conifers are abundant in the Table
Nunatak mesofossil assemblage and are represented by
vegetative as well as reproductive structures including
foliage shoots, isolated leaves, wood, pollen cones, isolated microsporophylls and seeds. Shoots are relatively
poorly preserved and it is not possible to link these with
the isolated leaves, which in general are better preserved.
In the following section all types of structures except
seeds are described; the seeds will be documented separately together with angiosperm fruits and seeds
(Eklund, 2003, work in progress). Conifer reproductive
structures include a single specimen each of an axis of
a pollen cone (Fig. 4A–C), a microsporophyll (Fig. 4D–
H) and an ovuliferous bract-scale complex (Fig. 4I).
Conifer shoots are represented by six different types of
conifer foliage shoots (Fig. 5A–F). A larger number of
conifer shoots have been recovered, but most are too
poorly preserved to be referred to the six types described
below. Therefore, each type of foliage shoot is described
here based on a single specimen. Two types of isolated
conifer leaves have been recovered (Fig. 6A–H). Conifer
wood is also present, including two types that are
documented here (Fig. 7A–E). The wood fragments are
dominantly 1.16–2.40 by 0.23–0.47 mm in size and
consist of fragments of xylem. Preservation of tracheids
and rays is generally good, and details such as bordered
pits on tracheids and crossfield pitting on the ray
crossfields are often preserved. Although the small size
of the specimens does not permit full taxonomic
description or full identification of wood types, in many
specimens diagnostic features are preserved (Fig. 7A–E).
Pollen cone axis
Description. 2.8 mm long by 0.32–0.52 mm wide.
Approximately 22 bases of microsporophylls are preserved as small protuberances arranged helically along
Description. A single conifer microsporophyll with
two pollen sacs has been recovered (DJ.755.1.115;
Fig. 4D–H). The microsporophyll is differentiated into
a filamentous stalk and a bilaterally flattened, leaf-like
head on which the pollen sacs are borne close to the
joint between the stalk and the head (Fig. 4D, E). One of
the pollen sacs is dehisced (Fig. 4E, G) whereas the other
one is intact (Fig. 4D, F). The latter is about 0.39 mm
long and 0.15 mm wide at the widest point (Fig. 4D, F),
and the dehisced pollen sac is about 0.53 mm long and
0.36 mm wide at the widest point (Fig. 4E, G). Both
pollen sacs have a narrowly elliptical to lens-shaped
outline (Fig. 4F, G). The basal part of the leaf-like head
of the microsporophyll is differentiated into a semicircular flange (Fig. 4H).
Remarks. Among extant conifers, microsporophylls
with two pollen sacs occur in the Cephalotaxaceae (2–3
or more pollen sacs per microsporophyll), Cupressaceae
(2–6), Pinaceae (2), Podocarpaceae (2), Sciadopityaceae
(2) and Taxodiaceae (2). Today the Cephalotaxaceae,
Pinaceae and Sciadopityaceae are restricted to the
Northern Hemisphere, whereas the Cupressaceae has a
wide distribution in both Hemispheres, Podocarpaceae
is mostly restricted to the Southern Hemisphere, and
Taxodiaceae is represented in the Southern Hemisphere
by one genus (Athrotaxis D. Don).
?Conifer ovulate bract-scale complex
Description. Bract 2.5 mm long by 0.9 mm wide, with
a narrowly ovate outline. The main body of the
structure is interpreted as a bract (arrowed b in Fig. 4I)
which bears an obtriangular ovule, on the adaxial side
close to the base (arrowed o in Fig. 4I). The ovule is
about 0.75 mm long and about 0.71 mm wide at the
widest point. Above the ovule there is a triangular
structure (arrowed ts in Fig. 4I) that is here interpreted
218
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
as the free apical part of the ovuliferous scale that is
otherwise fused with the ovule.
Remarks. The single specimen recovered (DJ.755.1.
128; Fig. 4I) is interpreted as a conifer ovulate bractscale complex. Ovulate bract-scale complexes with the
same structure as the fossil occur within extant Araucariaceae, and similar fossils have been referred to
Araucarites Presl (e.g., Hill and Brodribb, 1999; Cantrill,
2000; Cantrill and Falcon-Lang, 2001). It should be
noted that this specimen is substantially smaller than
others described as araucarian, but it may be immature.
Conifer foliage shoot type 1
Description. Shoot 3.7 mm long by 0.7 mm wide
(DJ755.1.25; Fig. 5A). The leaves are scale-like and
arranged helically along the axis to which they are appressed. Individual leaves are generally incompletely preserved, but one complete leaf is 1.7 mm long by 0.7 mm
wide. It is dorsoventrally flattened and elliptic with an
acute apex. A pronounced longitudinal keel is not visible.
Remarks. The helical phyllotaxis and general morphology of the leaves are widely distributed among
extant gymnosperms, occurring for example in extant
Dacrycarpus de Laubenf. and Dacrydium Sol. ex
Lambert (Podocarpaceae), Athrotaxis D. Don (Taxodiaceae), and juvenile foliage of Araucaria Jussieu
(Araucariaceae). However, due to the lack of diagnostic
features the detailed systematic affinities are uncertain.
Conifer foliage shoot type 2
Description. Shoot 3.5 mm long by 0.9 mm wide
(DJ.755.1.47; Fig. 5B). The leaves are dorsoventrally
flattened and arranged helically along the shoot axis
from which they diverge at an angle of about 30–45(.
Leaves are narrow linear to oblong, and about 1 mm
long by 0.2 mm wide.
Remarks. Fossil conifer shoots with a similar phyllotaxis and leaf morphology are usually referred to the
form genus Pagiophyllum Heer (Stewart and Rothwell,
1993). Among extant conifers similar foliage shoots
occur in the Podocarpaceae (e.g., Dacrycarpus, Dacrydium), Araucariaceae (e.g., Araucaria), and Taxodiaceae
(e.g., Athrotaxis, Sequoiadendron Buchholz, Taiwania
219
Hayata). Although this form is similar to conifer shoot
type 1 the leaves are more spreading. However, this may
not be indicative of separate taxonomic affinity as many
conifers have a broad range of morphological variation
from arrested growth form in the winter through spring
and summer growth. Nevertheless we have preferred to
describe this foliage separately as it provides a picture of
the range of morphologies present in the flora.
Conifer foliage shoot type 3
Description. Shoot 3.6 mm long by 2.4 mm wide
(DJ.755.1.35; Fig. 5C). The leaves are arranged helically
along the shoot axis from which the apices of the leaves
diverge minutely. The leaves have a succulent appearance and a rounded triangular outline.
Remarks. Fossil conifer shoots with a similar phyllotaxis and leaf morphology are usually referred to the
form genus Brachyphyllum Brongniart (Stewart and
Rothwell, 1993). Among extant conifers, foliage shoots
of similar morphology are found in the Podocarpaceae,
Taxodiaceae and Araucariaceae (Stewart and Rothwell,
1993). However, due to the lack of diagnostic features
the detailed systematic affinities are uncertain.
Conifer foliage shoot type 4
Description. Shoot 2.6 mm long by 1.1 mm wide
(DJ.755.1.34; Fig. 5D). The leaves are arranged helically
along the shoot axis. They are about 1.4 mm long and
0.8 mm wide at the widest point, and narrowly triangular with a strongly recurved apex.
Remarks. The systematic affinities of conifer foliage
shoot type 4 are unknown.
Conifer foliage shoot type 5
Description. Shoot 3 mm long and 0.8 mm wide
(R.2959.27.52; Fig. 5E). The leaves are arranged helically
along the shoot axis to which they are appressed. They
are about 1.3 mm long and 0.8 mm wide, with an ovate
to broadly elliptical outline. The apex of the leaf is
weakly acuminate and there is a pronounced longitudinal keel running along the length of the leaf.
Remarks. Conifer foliage shoot type 5 may represent
a better preserved specimen of the same taxon as conifer
Fig. 4. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): reproductive structures of conifers. A, axis of pollen cone
showing bases of approximately 22 microsporophylls in a helical arrangement along the axis (DJ.755.1.22). B, close-up of the pollen cone axis in A
(DJ.755.1.22). C, close-up of one of the microsporophyll bases in B (DJ.755.1.22). D, E, conifer microsporophyll in two different views; note the two
pollen sacs borne close to the joint between the filamentous stalk and the leaf-like head (DJ.755.1.115). F, close-up of pollen sacs in D (DJ.755.1.115).
G, close-up of pollen sac in E (DJ.755.1.115). H, close-up of the basal part of the leaf-like head in E (DJ.755.1.115). I, probable araucariaceous
conifer ovulate bract-scale complex; the main body of the structure is interpreted as a bract (arrow b) which bears an obtriangular structure, here
interpreted as an ovule, on the adaxial side close to the base (arrow o) and a triangular structure (arrow ts) here interpreted as a free apical part of the
ovuliferous scale that is otherwise fused with the ovule (DJ.755.1.128). Scale bars in A, D, E, I represent 1 mm; in B, 100 mm; in C, 50 mm; in F–H,
10 mm.
220
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Fig. 5. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): conifer foliage shoots. A, conifer foliage shoot type 1,
showing scale-like and helically arranged leaves that are appressed to the shoot axis (DJ755.1.25). B, conifer foliage shoot type 2, showing narrow
and linear to oblong, dorsoventrally flattened and helically arranged leaves that diverge from the shoot axis at an angle of about 30–45(
(DJ.755.1.47). C, conifer foliage shoot type 3, showing leaves with a succulent appearance and a rounded triangular outline; the leaves are helically
arranged and diverge minutely from the shoot axis (DJ.755.1.35). D, conifer foliage shoot type 4, showing narrowly triangular leaves with a strongly
recurved apex; the leaves are arranged helically along the shoot axis (DJ.755.1.34). E, conifer foliage shoot type 5, showing ovate to broadly elliptical
leaves with a pronounced longitudinal keel; the leaves are arranged helically along the shoot axis to which they are appressed (R.2959.27.52). F,
conifer foliage shoot type 6, showing a shoot that bifurcates apically into two branches; the leaves have a long basal decurrent portion and a short,
more or less triangular and succulent-looking free blade (R.2959.27.38). Scale bars represent 1 mm.
foliage shoot type 1. However, the two specimens differ
in the shape of the leaf (more ovate in type 5) and in
the absence of a pronounced longitudinal keel in the
leaves of type 1; the two shoots are, therefore, tentatively treated as different types. Among extant conifers
the closest affinities are probably with the Podocarpaceae (Dacrycarpus, Dacrydium) or Taxodiaceae (e.g.,
Sequoiadendron), in which shoots of the same general
morphology occur.
Conifer foliage shoot type 6
Description. Shoot 2.3 mm long and 0.9–1 mm wide
(R.2959.27.38; Fig. 5F). At the apex the shoot bifurcates
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
into two branches. Below the bifurcating apex the leaves
are arranged helically along the shoot axis. The leaves
have a long basal decurrent portion and a short, more or
less triangular and succulent-looking free blade. The
basal leaves appear to be intact and scale-like but the
more apical leaves are broken so that the exact size
of the free laminar portion cannot be determined. Epidermal cell outlines are barely expressed, stomata are
rare but present basally on either side of the midvein.
Stomata consist of a well-developed Florin ring that is
slightly sunken and the stomatal aperture is apparently
longitudinally oriented.
Remarks. The variation in leaf size suggests a conifer
with an interrupted ( perhaps seasonal) growth habit.
Conifer shoots with similar leaf morphology are found
in the Podocarpaceae (e.g., Dacrycarpus) and Taxodiaceae (e.g., Metasequoia Miki, Taxodium L.C. Rich). Of
these families, the well-developed Florin ring and
longitudinal orientation of the stomata suggests that
this taxon is closest to the Podocarpaceae.
Conifer leaf type 1
Description. The leaf fragments range from 1.3 to
2.0 mm long by 0.39 to 0.5 mm wide. Leaves range from
being linear to narrowly elliptic or narrowly ovate in
outline (Fig. 6A, B, E). The leaf base is slightly
constricted just above the base (Fig. 6A, E), and the
leaf apex is acute to acuminate. The midvein is raised on
the abaxial surface (Fig. 6A, B) but lacks expression on
the adaxial surface (Fig. 6E).
Leaves are hypostomatic. On the abaxial surface
numerous sunken stomata are distributed in two bands
on either side of the midvein (Fig. 6B–D). Stomata are
apparently arranged irregularly and are not in obvious
rows. The stomatal apparatus, comprises subsidiary
cells forming a raised Florin ring that is oval and about
45 mm long by 37 mm wide (Fig. 6D). Guard cells are
sunken and the stomatal aperture is oriented longitudinally. The upper surface is smooth and devoid of
stomates. Erosion of specimen DJ.755.1.49 reveals
a longitudinal structure within the leaf and lying above
the midvein (Fig. 6E).
Remarks. This leaf type is described based on three
specimens (DJ.755.1.29, Fig. 6A; DJ.755.1.129, Fig. 6B–
D; DJ.755.1.49, Fig. 6E). One specimen (DJ.755.1.29;
Fig. 6A) consists of three leaves surrounding a shoot
apex. Conifer leaf type 1 rarely has the leaf base preserved and it is apparent in DJ.755.1.49 that the base is
decurrent on the shoot (Fig. 6E). This leaf is eroded to
reveal the internal anatomy, although not well preserved, the smooth, longitudinally oriented structure
above the midvein is located where resin canals or resinfilled cells are found in modern podocarpaceous leaves.
It is likely that this leaf morphotype goes with conifer
221
shoot axis 6 (Fig. 5F) and represents the spreading
foliage phase. Dimorphic foliage is characteristic of
a number of genera within the Podocarpaceae. The
longitudinally orientated stomata and raised, smooth
Florin ring are also consistent with placement in the
Podocarpaceae.
Conifer leaf type 2
Description. The leaf is 1.7 mm long and 0.6 mm
wide, narrowly ovate in outline, with an acute apex
and a broad and raised longitudinal midvein (Fig. 6).
Numerous sunken stomata are very closely spaced on
both sides of the midvein (Fig. 6G, H). They have
prominent circular Florin rings that are about 39 mm in
diameter (Fig. 6H). The Florin rings do not appear
smooth and have surface expression of the underlying
subsidiary cells resulting in constrictions around the
circumference of the ring (Fig. 6H). The epidermal cells
appear papillate (Fig. 6G).
Remarks. This leaf type is known from a single
specimen (DJ.755.1.48; Fig. 6F–H). The well-developed
Florin rings, with constrictions marking the position of
subsidiary cells, verge on being papillate. This type of
micromorphology is commonly encountered in extant
Cupressaceae and Taxodiaceae, but is also present in the
Cephalotaxaceae and Taxaceae. Fossil Cheirolepidiaceae also have a similar micromorphology in some
species, but generally these have well-developed papillae
or lappets unlike the material described here. Given the
present-day distribution of conifer families it seems most
likely that this leaf type belongs to either the Taxodiaceae or Cupressaceae or the extinct Cheirolepidiaceae.
Podocarpoxylon sp.
Description. A single specimen R.2959.8.35 (Fig. 7A–
C) that has single rows of separate bordered pits in the
tracheids (Fig. 7B, C), and numerous small simple pits
on each ray cell (Fig. 7A). In other sections of this
sample these appear as 1–2 pits per crossfield, some with
thin borders.
Remarks. This wood can be most closely compared
with Podocarpoxylon Kräusel as it has bordered pits
with large, steeply inclined pores and small pits in the
crossfields. Conifer wood type 1 probably has affinities
with extant Podocarpaceae.
Phyllocladoxylon sp.
Description. A single specimen R.2959.8.48 (Fig. 7D,
E) that is characterised by vertical tracheids with single
and separate bordered pits on the walls of the tracheids
(Fig. 7E). The crossfield pits in the ray cells consist of
one or two large open pits with no borders (Fig. 7D).
222
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Remarks. The single large open pit in the crossfield is
characteristic of the form genus Phyllocladoxylon
Kräusel. This wood type is similar to those found in
some types of living Podocarpaceae.
Fossil wood with features similar to wood of modern
Podocarpaceae are commonly found in Cretaceous and
Cenozoic Antarctic fossil wood assemblages, including
those from James Ross and Seymour islands (Francis,
1991; Torres et al., 1994), Alexander Island (FalconLang and Cantrill, 2000) and the South Shetland Islands
(Falcon-Lang and Cantrill, 2001). Fossil wood of Araucarioxylon is also a common component of Antarctic
fossil floras but has yet to be described from the Table
Nunatak mesofossil assemblage. Furthermore, angiosperm wood has not been detected either.
3.4. Angiosperms
Remarks. Angiosperms are abundant in the Table
Nunatak mesofossil assemblage. They are mainly
represented by reproductive structures ( flowers, fruits
and seeds), but also by a few vegetative structures
( fragments of leaves). In the following section the leaf
fragments are described (Fig. 8A–H); the flowers, fruits
and seeds will be documented separately together with
conifer seeds (Eklund, 2003 and work in progress).
None of the leaf fragments shows stomata or other
taxonomic features, and their detailed systematic affinities within the angiosperms are unknown. Even the
suggested angiosperm affinities, based on the general
shapes, may in some cases be ambiguous. Nevertheless,
the variation seen points to a range in size, including
microphyllous forms, and a number of different morphologies (e.g., bilobed, entire, ?palmate).
Angiosperm leaf type 1
Description. The leaf is more or less complete, about
2.6 mm long by 1 mm wide (R.2959.27.35; Fig. 8A). It
has a short and wedge-shaped petiole that is about
0.5 mm long and 0.6 mm wide at the widest point at the
base. The leaf blade is ovate in outline. Veins and
stomata are not visible.
223
Angiosperm leaf type 2
Description. Leaf fragment that is about 2.3 mm long
by 1.8 mm wide (R.2959.27.36; Fig. 8B). The original
shape is unclear due to the fragmented nature of the
fossil. On the supposedly lower leaf surface there is
a pronounced vein that is about 0.25 mm wide and
unbranched. Stomata are not visible.
Angiosperm leaf type 3
Description. Leaf fragment that is about 2.4 mm long
by 1.4 mm wide (R.2959.27.37; Fig. 8C). The original
shape is unclear due to the fragmented nature of the
fossil. On the supposedly lower leaf surface there is
a pronounced vein that is about 0.36 mm wide and
which diverges into a secondary vein on the right-hand
side. Stomata are not visible.
?Angiosperm leaf type 4
Description. The leaf is complete, about 1.6 mm long
by 0.8 mm wide at the widest point at the apex
(DJ.755.1.40; Fig. 8D). It is obovate with a succulent
appearance. The apex appears to be less succulent than
the rest of the leaf and is bent almost perpendicular to
the main axis of the leaf. Veins and stomata are not
visible.
Angiosperm leaf type 5
Description. Leaf fragment, about 3.2 mm long by
1.8 mm wide (DJ.755.1.46; Fig. 8E). The original shape
is unclear due to the fragmented nature of the fossil, but
the general appearance indicates that it may have been
palmately lobed. The midvein and a basal secondary vein can be discerned as two diffuse ridges on
the supposedly upper leaf surface. Stomata are not
visible.
Angiosperm leaf type 6
Description. Leaf fragment, about 4 mm long by
1.4 mm wide at the widest point (R.2959.27.50; Fig. 8F).
It has a long and elongate petiole that is about 1.4 mm
long and 0.4 mm wide. The original shape of the leaf
Fig. 6. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): isolated conifer leaves. A–E, conifer leaf type 1; F–H,
conifer leaf type 2. A, conifer leaf type 1; three leaves surrounding a shoot apex (DJ.755.1.29). B, conifer leaf type 1 in abaxial view; leaf with a linear
to narrowly elliptic to narrowly ovate outline and a midvein that is raised on the abaxial surface (DJ.755.1.129). C, close-up of leaf in B showing
numerous sunken stomata distributed in two bands on either side of the midvein on the abaxial surface (DJ.755.1.129). D, close-up of the stomatal
apparatus in specimen DJ.755.1.129 (B, C) showing subsidiary cells forming an oval and raised Florin ring and sunken guard cells. E, conifer leaf
type 1 in adaxial view, showing that the leaf base is decurrent on the shoot; note probable longitudinal resin canal above the midvein (DJ.755.1.49).
F, conifer leaf type 2 in adaxial view, showing narrowly ovate outline, with an acute apex and a broad and raised longitudinal midvein (DJ.755.1.48).
G, close-up of margin of leaf in F, showing numerous sunken and very closely spaced stomata (DJ.755.1.48). H, close-up of stomata in G, showing
prominent circular Florin rings which appear papillate (DJ.755.1.48). Scale bars in A, B, E, F represent 1 mm; in C, 100 mm; in D, 10 mm; in G, H,
50 mm.
224
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Fig. 7. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): fragments of charcoalified conifer wood. A, radial
longitudinal section (RLS) of conifer wood type 1, showing horizontal ray cells containing several small simple pits (R.2959.8.35). B, RLS of conifer
wood type 1, showing vertical tracheids with single and separate bordered pits in the tracheid walls (R.2959.8.35). C, RLS of conifer wood type 1
showing detail of the bordered pits in the tracheid walls (R.2959.8.35). D, RLS of conifer wood type 2 showing horizontal ray cells with large oval
pits (R.2959.8.48). E, RLS of conifer wood type 2 showing vertical tracheids with single and separate bordered tracheid pits (R.2959.8.48). Scale bars
in A, B, D, E represent 50 mm; in C, 5 mm.
blade is unclear due to the fragmented preservation of
the fossil. Veins and stomata are not visible.
?Angiosperm leaf type 7
Description. Leaf bilobed with a succulent appearance (R.2959.27.45; Fig. 8G, H). One of the lobes is
completely preserved whereas the other lobe is fragmented. The complete lobe is about 2.9 mm long from
apex of lobe to base of leaf, and 0.9 mm wide. The basal
unlobed part of the leaf is about 1.1 mm long whereas
the apical free part is about 1.8 mm long. At the base of
the leaf there is a ring-like cushion that is about 0.3 mm
long and 0.7 mm wide. At the apical part of this
structure a longitudinal ridge is present. At a distance of
about 0.4 mm from the apex of the ring-like cushion, the
ridge bifurcates and continues along each of the two leaf
lobes respectively. The ridges are interpreted as primary
and secondary veins. Stomata are not visible.
4. Discussion
The application of Quaternary sieving techniques to
Mesozoic and Cenozoic strata has revealed an untapped
source of information on the plant fossil record. This
has resulted in a revolution of our knowledge of the
early radiation of flowering plants (Crane et al., 1995)
but these mesofossil floras are also revealing information on vegetational composition that augments the
more traditional macrofloral (leaves and wood) and
microfloral (spores and pollen) record (e.g., Herendeen
et al., 1999; Takahashi et al., 1999b). To date the
discovery of mesofossil floras has been centred in the
Northern Hemisphere (e.g., Europe, Asia and North
America). In particular, these floras have demonstrated
the presence of many families of flowering plants and
contributed to our understanding of the pattern of
radiation in this group. Although many floras are
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
225
Fig. 8. Mesofossils from the Table Nunatak Formation (late Santonian; Late Cretaceous): isolated angiosperm leaves. A, angiosperm leaf type 1,
almost complete leaf showing a short and wedge-shaped petiole (R.2959.27.35). B, angiosperm leaf type 2, fragment of leaf showing the supposedly
lower leaf surface; note the pronounced vein (R.2959.27.36). C, angiosperm leaf type 3, fragment of leaf showing the supposedly lower leaf surface;
note the pronounced vein that diverges into a secondary vein on the right side (R.2959.27.37). D, ?angiosperm leaf type 4, supposedly lower surface of
an almost complete leaf (DJ.755.1.40). E, angiosperm leaf type 5, fragment of leaf showing the supposedly upper leaf surface (DJ.755.1.46). F,
angiosperm leaf type 6, fragment of leaf with an elongate petiole (R.2959.27.50). G, ?angiosperm leaf type 7, incompletely preserved bilobed leaf
showing a bifurcating longitudinal ridge that may represent primary and secondary veins (R.2959.27.45). H, ?angiosperm leaf type 7, close-up of the
bifurcating leaf in G (R.2959.27.45). Scale bars in A–G represent 1 mm; in H, 0.5 mm.
226
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
known from the Northern Hemisphere, few have been
described from the Southern Hemisphere and none from
the high latitude regions of Gondwana. In view of the
marked differences in taxonomic composition of the
Southern Hemisphere floras today, and the long history
of separate floristic development, mesofossil floras from
this part of the world are critical for understanding the
radiation of Southern Hemisphere groups of flowering
plants. Consequently the discovery of the first mesofossil
flora from Antarctica provides an important perspective
of the vegetation that is not apparent from examination
of the leaf, wood and pollen record.
The Table Nunatak flora is the first mesofossil flora
described from the Antarctic and one of the few from
the Southern Hemisphere. However, the flora is not as
well preserved as those described from the Northern
Hemisphere due to the depositional environment. The
flora accumulated in shallow marine sands (Hathway
et al., 1998) and represents the remains of vegetation
that were washed into the marine environment from the
land. Consequently the assemblage is abraded and
sorted so that some size classes are probably missing.
Floras are sparsely known from the Upper Cretaceous
of the Antarctic including leaf floras from the Coniacian
and Santonian (Hayes, 1999). Wood has been described
from Coniacian through Maastrichtian deposits (e.g.,
Poole and Francis, 1999, 2000; Poole and Gottwald,
2001) and spore/pollen floras occur throughout the
Upper Cretaceous (e.g., Dettmann and Thomson, 1987;
Askin, 1992). The Coniacian to early Campanian interval marked major changes in diversity (Cantrill and
Poole, 2002) and community structure.
The largely vegetative remains described here give an
incomplete picture of the vegetation. Coniferous foliage
dominates the assemblage but it is clear that angiosperms are important. The angiosperm leaf material is
very fragmentary and is broken into size classes that are
of a similar dimension to the coniferous foliage,
indicating a strong taphonomic bias. The conifer foliage
and wood indicates the importance of Podocarpaceae,
a feature that is supported by wood and pollen records
elsewhere in the Antarctic Peninsula. It is interesting to
note the abundance of Phyllocladoxylon, a wood type
that is relatively rare in Lower Cretaceous strata but
which becomes more important up-sequence.
The diversity of angiosperm leaves from Table
Nunatak is rather low but not surprising given the taphonomic biases. It is difficult to estimate the diversity and
abundance of angiosperms in the vegetation from this
deposit, but greater angiosperm diversity is suggested by
the seed and fruit flora (Eklund, work in progress). The
few flowers described from the deposits indicate the
presence of magnoliids and eudicots within the flora
(Eklund, 2003). This interpretation is supported by leaf
floras from the Larsen Basin to the north. Coniacian and
Santonian sediments that were quickly deposited and
less strongly transported, have a high angiosperm
diversity (60–70%) and abundance (Hayes, 1999).
One interesting feature of the Table Nunatak
assemblage is the preservation of the material by charcoalification. This indicates the importance of wildfires
within the ecosystem, a characteristic that has been
noted for other Cretaceous floras in the Antarctic
Peninsula (Falcon-Lang and Cantrill, 2002). Considering that Table Nunatak is located in the Larsen Basin,
a tectonically-active basin situated on the back-arc side
of the Antarctic Peninsula volcanic arc (Hathway,
2000), this may not be too surprising. Nevertheless it
indicates that wildfires probably played an important
role in the ecology of the vegetation.
5. Conclusions
The Table Nunatak assemblage is similar to Late
Cretaceous mesofossil assemblages from the Northern
Hemisphere (e.g., the late Santonian Allon flora and the
early Coniacian Kamikitaba flora) in the general
composition of vegetational elements (lycopods, ferns,
conifers, angiosperms). The more precise systematic affinities within these major groups are difficult to establish with certainty, largely because many of the fossils
from Table Nunatak have suffered considerable abrasion. The reason for this may be that they were deposited
in a marine environment (the more well-preserved fossils
in the Allon flora, for example, were deposited in a
floodplain pond). The mesofossils from Table Nunatak
nevertheless provide important complementary information to that provided by micro- and macrofossils. To
obtain as accurate a picture as possible of the living
vegetation in Antarctica during the Late Cretaceous, all
kinds of fossils have to be taken into consideration,
especially if morphological as well as taxonomic diversity of the ecosystems are to be understood.
This account of the Table Nunatak assemblage
shows that there is a considerable potential for finding
additional Cretaceous mesofossil floras in the Southern
Hemisphere. Hopefully, future studies will reveal new
outcrops with more complete fossils that can be reliably identified. Considering that, at present, the Table
Nunatak flora is one of few mesofossil assemblages from
the Southern Hemisphere with diverse angiosperm
remains, and the only one from Antarctica, the
discovery of new such floras is crucial for obtaining
a more complete understanding of the evolution of the
southern floras and of Cretaceous plants in general.
Acknowledgements
We thank Alan R. Hemsley and Anne-Marie Tosolini
for valuable discussions and comments on the fossils;
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
Eric Condliffe for assistance with the SEM; David Banks
for help with HF work; and The British Antarctic Survey
for letting us work on the material. This work was supported by grants to HE from STINT (Stiftelsen för
internationalisering av högre utbildning och forskning,
The Swedish Foundation for International Cooperation
in Research and Higher Education), Helge Ax:son
Johnsons Stiftelse and Stiftelsen P. E. Lindahls fond.
References
Askin, R.A., 1992. Late Cretaceous–early Tertiary Antarctic outcrop
evidence for past vegetation and climates. Antarctic Research
Series 56, 61–73.
Batten, D.J., 1988. Revision of S.J. Dijkstra’s Late Cretaceous
megaspores and other plant microfossils from Limburg, The
Netherlands. Mededelingen Rijks Geologische Dienst 41-3, 2–55.
Batten, D.J., Ferguson, D.J.P., 1987. Cabochonicus, a new genus for
species of gemmate megaspores previously referred to Verrutriletes.
Journal of Micropalaeontology 6, 65–75.
Batten, D.J., Creber, G.T., Zhou Zhiyan, 1984. Fossil plants and
other debris in Cretaceous sediments from Deep Sea Drilling
Project Leg 80: their paleoenvironmental significance and source
potential. Initial Reports of the Deep Sea Drilling Project 80,
pp. 629–641.
Batten, D.J., Dutta, R.J., Knobloch, E., 1996. Differentiation, affinities
and palaeoenvironmental significance of the megaspores Arcellites
and Bohemisporites in Wealden and other Cretaceous successions.
Cretaceous Research 17, 39–65.
Cantrill, D.J., 2000. A Cretaceous (Aptian) flora from President Head,
Snow Island, Antarctica. Palaeontographica B 253, 153–191.
Cantrill, D.J., Drinnan, A.N., Webb, J.A., 1995. Late Triassic plant
fossils from the Prince Charles Mountains, East Antarctica.
Antarctic Science 7, 51–62.
Cantrill, D.J., Falcon-Lang, H.J., 2001. Cretaceous (Late Albian)
Coniferales of Alexander Island, Antarctica. 2. Leaves, reproductive structures and roots. Review of Palaeobotany and Palynology
115, 119–145.
Cantrill, D.J., Poole, I., 2002. Cretaceous patterns of floristic change
in the Antarctic Peninsula. In: Crame, J.A., Owen, A.W. (Eds.),
Palaeobiogeography and Biodiversity Change: the Ordovician
and Mesozoic–Cenozoic Radiations. Geological Society, London,
Special Publication 194, pp. 141–152.
Collinson, M.E., Batten, D.J., Scott, A.C., Ayonghe, S.N., 1985.
Palaeozoic, Mesozoic and contemporaneous megaspores from the
Tertiary of southern England: indicators of sedimentary provenance and ancient vegetation. Journal of the Geological Society,
London 142, 375–395.
Crane, P.R., Friis, E.M., Pedersen, K.R., 1994. Palaeobotanical
evidence on the early radiation of magnoliid angiosperms. Plant
Systematics and Evolution 8 (Suppl.), 51–72.
Crane, P.R., Friis, E.M., Pedersen, K.R., 1995. The origin and early
diversification of angiosperms. Nature 374, 27–33.
Crepet, W.L., Nixon, K.C., 1994. Flowers of Turonian Magnoliidae
and their implications. Plant Systematics and Evolution 8 (Suppl.),
73–91.
Dettmann, M.E., Thomson, M.R.A., 1987. Cretaceous palynomorphs
from the James Ross Island area, Antarcticada pilot study. British
Antarctic Survey Bulletin 77, 13–59.
Eklund, H., 2000. Lauraceous flowers from the Late Cretaceous of
North Carolina, U.S.A. Botanical Journal of the Linnean Society
132, 397–428.
Eklund, H., 2003. First Cretaceous flowers from Antarctica. Review of
Palaeobotany and Palynology 127, 187–217.
227
Eklund, H., Kvaček, J., 1998. Lauraceous inflorescences and flowers
from the Cenomanian of Bohemia (Czech Republic, central
Europe). International Journal of Plant Sciences 159, 668–686.
Eklund, H., Friis, E.M., Pedersen, K.R., 1997. Chloranthaceous floral
structures from the Late Cretaceous of Sweden. Plant Systematics
and Evolution 207, 13–42.
Ellis, C.H., Tschudy, R.H., 1964. The Cretaceous megaspore genus
Arcellites Miner. Micropaleontology 10, 73–79.
Falcon-Lang, H.J., Cantrill, D.J., 2000. Cretaceous (late Albian)
Coniferales of Alexander Island, Antarctica. 1: Wood taxonomy,
a quantitative approach. Review of Palaeobotany and Palynology
111, 1–17.
Falcon-Lang, H.J., Cantrill, D.J., 2001. Gymnosperm woods from
the Cretaceous (mid-Aptian) Cerro Negro Formation, Byers
Peninsula, Livingston Island, Antarctica: the aborescent vegetation
of a volcanic arc. Cretaceous Research 22, 277–293.
Falcon-Lang, H.J., Cantrill, D.J., 2002. Terrestrial palaeoecology of
an Early Cretaceous volcanic archipelago, Byers Peninsula and
President Head, South Shetlands Islands, Antarctica. Palaios 17,
535–549.
Francis, J.E., 1991. Palaeoclimatic significance of Cretaceous–early
Tertiary fossil forests on the Antarctic Peninsula. In: Thomson,
M.R.A., Crame, J.A., Thomson, J.W. (Eds.), Geological Evolution of Antarctica. Cambridge University Press, Cambridge,
pp. 623–627.
Friis, E.M., 1984. Preliminary report of Upper Cretaceous angiosperm
reproductive organs from Sweden and their level of organization.
Annals of the Missouri Botanical Garden 71, 403–418.
Friis, E.M., 1985. Structure and function in Late Cretaceous
angiosperm flowers. Biologiske SkrifterdDet Kongelige Danske
Videnskabernes Selskab 25, 1–37.
Friis, E.M., Skarby, A., 1981. Structurally preserved angiosperm
flowers from the Upper Cretaceous of southern Sweden. Nature
291, 485–486.
Friis, E.M., Pedersen, K.R., Crane, P.R., 1994. Angiosperm floral
structures from the Early Cretaceous of Portugal. Plant Systematics and Evolution 8 (Suppl.), 31–49.
Friis, E.M., Pedersen, K.R., Crane, P.R., 2001. Fossil evidence of
water lilies (Nymphaeales) in the Early Cretaceous. Nature 410,
357–360.
Frumin, S., Friis, E.M., 1996. Liriodendroid seeds from the Late
Cretaceous of Kazakhstan and North Carolina, USA. Review of
Palaeobotany and Palynology 94, 39–55.
Frumin, S., Friis, E.M., 1999. Magnoliid reproductive organs from the
Cenomanian–Turonian of north-western Kazakhstan: Magnoliaceae and Illiciaceae. Plant Systematics and Evolution 216, 265–288.
Frumina, S.I., Zhilin, S.G., Korchagina, I.A., 1995. Alapaja (Taxodiaceae) seeds from the Cenomanian–Turonian of northern
Kazakhstan. Paleontological Journal 29, 194–202.
Gandolfo, M.A., Nixon, K.C., Crepet, W.L., 2001. Turonian Pinaceae
of the Raritan Formation, New Jersey. Plant Systematics and
Evolution 226, 187–203.
Gandolfo, M.A., Nixon, K.C., Crepet, W.L., Ratcliffe, G.E., 2000.
Sorophores of Lygodium Sw. (Schizaeaceae) from the Late Cretaceous of New Jersey. Plant Systematics and Evolution 221, 113–123.
Hathway, B., 2000. Continental rift to back-arc basin: Jurassic–
Cretaceous stratigraphical and structural evolution of the Larsen
Basin, Antarctic Peninsula. Journal of the Geological Society,
London 157, 417–432.
Hathway, B., Macdonald, D.I.M., Riding, J.B., Cantrill, D.J., 1998.
Table Nunatak: a key outcrop of Upper Cretaceous shallowmarine strata in the southern Larsen Basin, Antarctic Peninsula.
Geological Magazine 135, 519–535.
Hayes, P.A., 1999. Cretaceous angiosperm leaf floras from Antarctica.
PhD thesis, University of Leeds (unpublished).
Herendeen, P.S., Magallón-Puebla, S., Lupia, R., Crane, P.R.,
Kobylinska, J., 1999. A preliminary conspectus of the Allon flora
228
H. Eklund et al. / Cretaceous Research 25 (2004) 211–228
from the Late Cretaceous (late Santonian) of central Georgia,
U.S.A.. Annals of the Missouri Botanical Garden 86, 407–471.
Hill, R.S., Brodribb, T.J., 1999. Southern conifers in time and space.
Australian Journal of Botany 47, 639–696.
Kovach, W.L., Batten, D.J., 1989. Worldwide stratigraphic occurrences
of Mesozoic and Tertiary megaspores. Palynology 13, 247–277.
Koppelhus, E.B., Batten, D.J., 1992. Megaspore assemblages from
the Jurassic and lowermost Cretaceous of Bornholm, Denmark.
Geological Survey of Denmark DGU Series A 32, 1–81.
Kramer, K.U., 1990. Marsileaceae. In: Kubitzki, K. (Ed.), The Families
and Genera of Vascular Plants. Pteridophytes and Gymnosperms,
vol. 1. Springer, Berlin, pp. 180–183.
Lawver, L.A., Gahagan, L.M., Coffin, M.F., 1992. The development
of paleoseaways around Antarctica. In: Kennett, J.P., Warnke,
D.A. (Eds.), The Antarctic Paleoenvironment: a Perspective on
Global Change. Antarctic Research Series 56, pp. 7–30.
Li, W.-B., Batten, D.J., 1986. The Early Cretaceous megaspores
Arcellites and closely associated Crybelosporites microspores from
northeast Inner Mongolia, P.R. China. Review of Palaeobotany
and Palynology 46, 189–208.
Lupia, R., 1995. Paleobotanical data from fossil charcoal: an actualistic
study of seed plant reproductive structures. Palaios 10, 465–477.
Macdonald, D.I.M., 1985. Sedimentological investigations in the
Antarctic Peninsula area. Geological field report R/1984-85/G4,
British Antarctic Survey, Cambridge, UK.
Magallón, S., Herendeen, P.S., Crane, P.R., 2001. Androdecidua
endressii gen. et sp. nov., from the Late Cretaceous of Georgia
(United States): further floral diversity in Hamamelidoideae
(Hamamelidaceae). International Journal of Plant Sciences 162,
963–983.
Marcinkiewicz, T., 1979. Fungi-like forms on Jurassic megaspores.
Acta Paleobotanica 20, 123–128.
Miner, E.L., 1935. Paleobotanical examinations of Cretaceous and
Tertiary coals. The American Midland Naturalist 16, 585–625.
Poole, I., Francis, J.E., 1999. The first record of antherospermataceous
wood from the upper Cretaceous of Antarctica. Review of
Palaeobotany and Palynology 107, 97–107.
Poole, I., Francis, J.E., 2000. The first record of fossil wood of
Winteraceae from the Upper Cretaceous of Antarctica. Annals of
Botany 85, 307–315.
Poole, I., Gottwald, H., 2001. Monimiaceae sensu lato, an element of
Gondwanan polar forests: evidence from the Late Cretaceous–
early Tertiary wood flora of Antarctica. Australian Systematic
Botany 14, 207–230.
Potonié, R., 1956. Synopsis der Gattungen der Sporae dispersae.
Beihefte zum Geologischen Jahrbuch 23, 1–103.
Schönenberger, J., Friis, E.M., 2001. Fossil flowers of ericalean affinity
from the Late Cretaceous of southern Sweden. American Journal
of Botany 88, 467–480.
Schönenberger, J., Pedersen, K.R., Friis, E.M., 2001a. Normapolles
flowers of fagalean affinity from the Late Cretaceous of Portugal.
Plant Systematics and Evolution 226, 205–230.
Schönenberger, J., Friis, E.M., Matthews, M.L., Endress, P.K., 2001b.
Cunoniaceae in the Cretaceous of Europe: evidence from fossil
flowers. Annals of Botany 88, 423–437.
Scott, A.C., 1989. Observations on the nature and origin of fusain.
International Journal of Coal Geology 12, 443–475.
Scott, A.C., 2000. The pre-Quaternary history of fire. Palaeogeography, Palaeoclimatology, Palaeoecology 164, 281–329.
Scott, A.C., Jones, T.P., 1991a. Fossil charcoal: a plant-fossil record
preserved by fire. Geology Today 7, 214–216.
Scott, A.C., Jones, T.P., 1991b. Microscopical observations of recent
and fossil charcoal. Microscopy and Analysis 24, 13–15.
Stewart, W.N., Rothwell, G.W., 1993. Paleobotany and the Evolution
of Plants, second ed. Cambridge University Press, Cambridge,
x C 521 pp.
Takahashi, M., Crane, P.R., Ando, H., 1999a. Esgueiria futabenensis
sp. nov., a new angiosperm flower from the Upper Cretaceous
(lower Coniacian) of northeastern Honshu, Japan. Paleontological
Research 3, 81–87.
Takahashi, M., Crane, P.R., Ando, H., 1999b. Fossil flowers and
associated plant fossils from the Kamikitaba locality (Ashizawa
Formation, Futaba Group, lower Coniacian, Upper Cretaceous) of
northeast Japan. Journal of Plant Research 112, 187–206.
Torres, T., Marenssi, S., Santillana, S., 1994. Maderas fósiles de la isla
Seymour, Formación La Meseta, Antártica. Serie Cientı́fica del
INACH 44, 17–38.
Tosolini, A.-M.P., 2000. Early Cretaceous plant biofacies of the
Otway and Strzelecki groups, Victoria. PhD thesis, University of
Melbourne (unpublished).