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1
A NEW FOSSIL CONIFER, BELLARINEA RICHARDSII, FROM THE
EARLY CRETACEOUS STRZELECKI GROUP, SOUTHEASTERN
VICTORIA
NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN & STEPHEN MCLOUGHLIN
School of Botany, The University of Melbourne, Victoria, 3010, Australia
NAGALINGUM, NATHALIE S., DRINNAN, ANDREW N. & MCLOUGHLIN, STEPHEN 2005:07:01. A new fossil
conifer, Bellarinea richardsii, from the Early Cretaceous Strzelecki Group, southeastern
Victoria. Proceedings of the Royal Society of Victoria 117(1):1-12. ISSN 0035-9211.
A new species of fossil conifer foliage, Bellarinea richardsii, is described and illustrated
from Neocomian (Early Cretaceous) sediments of the Tyers River Subgroup in south-eastern Victo-
ria. The specimens consist of intact seasonal shoots and isolated leaves, and their depositional setting
in prominent leaf mats suggests a seasonal, deciduous habit. Individual leaves are spirally inserted
on the shoot axis but the leaf bases are twisted to lie on a common plane giving the shoots plagiotropic
symmetry. Although lacking attached reproductive structures the gross morphology and cuticular
details of the shoots and leaves suggests assignment to either Podocarpaceae or Taxodiaceae. This
species and a range of comparable forms represent a prominent component of Australian mid-Mesozoic
floras.
Key words: Palaeobotany, conifer, Early Cretacceous, Strzelecki Group
MESOZOIC FLORAS have been known from Vic-
toria since the pioneering work of McCoy (1874,
1875). Early reports assigned much of this material
to the Jurassic (McCoy 1860, 1875, Seward 1904),
but for the last 40 years it has been recognized that
this component is Early Cretaceous in age (Dettmann
1963, Douglas 1969, 1973). Victorian Cretaceous
floras are of interest because the forests that they rep-
resent grew at high southern latitudes, experienced
conditions unlike any that exist in the world today,
and were home to a diverse biota including polar di-
nosaurs (Rich et al. 1988).
Conifers represented an important component of
floras throughout the Victorian Early Cretaceous.
Assemblages assigned to Douglas’ (1969) Zone A
(=Ptilophyllum spinosum-P. castertonensis Zone:
latest Jurassic? to early Neocomian) are known only
from bore cores in the Otway Basin and their conifer
remains have not been studied in detail. Assemblages
referable to Douglas’ Zone B (=Phyllopteroides laevis
Zone of Cantrill & Webb 1987: Neocomian) are
recorded from the Boola Boola Forest of Gippsland,
small areas on the Mornington Peninsula and Philip
Island and from the subsurface of the western Otway
Basin. Rich plant assemblages from these beds have
recently been described by McLoughlin et al. (2002).
Assemblages assigned to Douglas’ Zone C (roughly
equivalent to the Phyllopteroides serrata Zone of
Cantrill & Webb 1987: Barremian to earliest Albian)
are widely represented in the Gippsland and Otway
basins. Several taxa of coniferous foliage have been
described from this zone including Bellarinea barklyi
Florin, Elatocladus mccoyi Florin, Elatocladus sp.,
Pod ozamites ellipticus McCoy, Brachyphyllum
gi pp sland ic um McC oy , Arauca ri a sp. cf. A.
heterophylla (Salisbury) Franco, and a range of cones,
cone scales, and seeds mostly with inferred araucarian
or podocarpacean affinities (McCoy 1874, Florin
1952, Drinnan & Chambers 1986). Douglas’ Zone D
(roughly equivalent to the Phyllopteroides dentata
Zone of Cantrill & Webb 1987: Albian) assemblages
are confined to the Otway Basin and are rich in
conifers. Cantrill & Douglas (1988) and Cantrill
(1991, 1992) documented the leaf morphology,
cuticular features and phylogenetic affinities of five
species ascribed to Araucaria, one to Agathis, and
several broad-leafed forms assigned to the form-genus
Podozamites. Cantrill (1991) suggested that the
Podozamites species were possibly representatives of
Podocarpaceae or Araucariaceae, and Pole (1995) later
transferred Podozamites taenioides to Aracarioides.
Cantrill & Douglas (1988) erected Geinitzia tetragona
for conifer foliage associated with roots bearing
mycorrhizal nodules and suggested a taxodiaceous
affinity for this species, but Pole (2000) considered
the cuticular micromorphology to be indicative of a
cheirolepidiaceous affinity and transferred this species
to the new genus Otwayia.
Assemblages representative of the Phyllopteroides
laevis Zone (Douglas’ Zone B) are best expressed in
exposures of the Tyers River Subgroup in the Boola
Boola Forest north-northwest of Traralgon (Fig. 1A).
2 NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN AND STEPHEN MCLOUGHLIN
This zone is notable for its abundance of small-leaved
bennetti talea ns (Otozamites) and severa l other
pteridosperm taxa [Taeniopteris daintreei McCoy,
Rintoul ia vari ab ilis (Douglas) McLoughlin &
Nagalingum in McLoughlin et al. (2002), Komlopteris
in di ca (Feistmantel) Barbacka and
Pachydermophyllum austropapillosum (Douglas)
McLoughlin & Nagalingum in McLoughlin et al.
(2002)], which most likely represented the principal
mid-storey elements of the vegetation. Associated with
these pterid os perm leav es in the Bool a Boola
assemblages are abundant conifer leaves belonging
to plants that probably constituted the upper stratum
of these Early Cretaceous forests. Three principal
conifer species are represented : Brachyphyllum
tyersensis Tosolini & McLoughlin (in McLoughlin et
al. 2002), Otwayia hermata Tosolini & McLoughlin
(in McLoughlin et al. 2002) and a new species of
Bellarinea, that is the basis of this paper.
GEOLOGICAL SETTING
The Boola Boola Forest is located approximately 12
km north-northwest of Traralgon, Gippsland (Fig. 1A).
Th e ro ck s ex po sed in this area repr es en t th e
northernmost extent of Cretaceous sediments in the
Gippsland Basin and they rest on Lower Devonian
metasedimentary rocks with an angular unconformity
of considerable relief. They are separated from
Cretaceou s exposur es of the Sout h Gippsla nd
Highlands by Tertiary sediments in the Latrobe Valley
Depression. Lower Cretaceous rocks of the Gippsland
Basin have been assigned in their entirety to the
Strzelecki Group (Douglas 1988). The upper part of
the succession is dominated by feldspathic sandstones
and has been assigned to the ‘Wonthaggi Formation’
by Constantine & Holdgate (1993). The lower part of
the succession, assigned to the Tyers River Subgroup
and principally exposed in the Boola Boola Forest
area, is dominated by conglomerates and quartzose
or lithic sandstones. The Tyers River Subgroup
incorporates the Tyers Conglomerate (c. 120 m thick)
and Rintoul Creek Formation (c. 480 m thick) in
ascending order (Tosolini et al. 1999). The Rintoul
Creek Formation has been further subdivided into a
lower unit (Locmany Member) of mixed lithologies
and an upper unit (Exalt Member) dominated by thick
sandstone packages. The Tyers Conglomerate is
interpreted to represent alluvial fan and proximal
braided river deposits whereas the succeeding Rintoul
Creek Form ati on represents mixed braided and
meandering river deposits in alluvial valley settings
(Tosolini et al. 1999). The material used in this study
is from the lower part of the type section of the Rintoul
Creek Formation (Locmany Member), approximately
160 m above the base of the unit. Biostratigraphic
studies of the sampled beds have assigned these rocks
to the Foraminisporis wonthaggiensis palynozone
(Dettmann 1963, Helby et al. 1987), Phyllopteroides
laevis macrofloral zone (Cantrill & Webb 1987), and
the Trikonia locmaniensis megaspore zone (Tosolini
et al. 2002) of Neocomian age (Fig. 1B).
MATERIAL AND METHODS
Specimens used in this study were collected over a
period of 35 years from the mid-1960s by Dr. J.
Douglas (formerly of the Geological Survey of
Victoria) and the present authors. The material is
derived from several localities in the Boola Boola
State Forest, 12 km NNW of Traralgon, Victoria (Fig.
1A). Cuticles were prepared by oxidation in Schultze’s
solution (nitric acid with dissolved potassium chloride
crystals) for up to one hour to remove coalified
mesophyll tissues. Slight heating (up to 45°C) and
further treatment with 5% sodium hydroxide or 5%
ammonia for up to 15 minutes was undertaken in an
attempt to clean the cuticle. Despite varied chemical
treatments and heating, B. richardsii cuticle proved
difficult to recover. It is thin and readily fragmented.
In most cases, the best detail of the external surface
was obtained by scanning electron microscopy of
unoxidized leaves (Fig. 2H, I). Recovery of cuticle
after oxidation of the coalified mesophyll did not yield
fragments large enough to provide significant details
of the cuticle’s inner surface morphology. Material
for scanning electron microscopy was air-dried,
attached to stubs using double sided carbon stickers,
and sputter coated with gold. All measurements of
epidermal fea tu res are fro m dried spe ci mens.
Specimens with the prefix MVP are registered with
Museum Victoria, Melbourne.
SYSTEMATIC PALAEOBOTANY
Phylum CONIFEROPHYTA
Class CONIFEROPSIDA
Order CONIFERALES
Genus Bellarinea Florin 1952 emend.
3
BELLARINEA RICHARDSII FROM SOUTHEASTERN VICTORIA
Fig. 1. Geographic and stratigraphic position of the fossil beds. A. Map of Victoria showing the distribution of Cretaceous sedimentary
rocks and the location of Boola Boola Forest; B. Cretaceous stratigraphy and biozones of the Gippsland Basin. Adapted from Douglas
(1969), Cantrill & Webb (1987), Helby et al. (1987), Smith (1988), Tosolini et al. (1999) and Tosolini et al. (2002).
4NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN AND STEPHEN MCLOUGHLIN
Fig. 2. Bellarinea richardsii. A. Shoot displaying typical leaf arrangement, MVP209950B, Loc. RC20A; B. Shoot with incurved, narrow,
falcate leaves, MVP209943, Loc. RC17D; C. Immature or under-developed shoot with leaves reducing in size towards the shoot apex,
MVP209949, Loc. RC17D; D. Shoot displaying typical pseudodistichous leaf arrangement, MVP209942 (holotype), Loc. L14; E. shoot
displaying typical leaf arrangement and form, NMVP209950A, Loc. RC20A; F. Enlargement of typical shoot axis showing pseudodistichous,
spirally inserted, leaf bases, MVP209942, Loc. L14; G. Shoot with narrow coalified leaves, MVP209948, Loc. RC20A; H. Scanning electron
micrograph of MVP209948 showing regular epidermal cells with bulging periclinal walls and stomates with papillae; I. Scanning electron
micrograph of MVP209948 showing stomates in longitudinal bands either side of the midvein. Scale bar = 10 mm for A-G; 100 !m for H; 1
mm for I.
Type species: Bell arinea barklyi Florin 1952;
Eumeralla Formation; Aptian; Bellarine Peninsula
near Geelong, Victoria, Australia.
Emended diagnosis. As per Florin (1952), but decidu-
ous determinate shoots without branches or terminal
resting buds.
Florin (1952) erected Bellarinea for shoots with
essentially identical gross architecture to those of
Elatocladus but where cuticular features revealed that
the leaves were hypostomatic with haplocheilic sto-
mata arranged in a band on each side of the midvein.
Florin’s specimens came from Aptian sediments in
southern Victoria, and his two species remain the only
ones attrib ut ed to this genus until this study.
Elatocladus was erected by Halle (1913) to encom-
pass sterile coniferous shoots of uncertain affinity with
spiral phyllotaxy, including those with leaf bases
twisted to give a plagiotropic orientation. Halle in-
cluded in the genus three species from the mid-
Mesozoic of India, which he considered identical to
the specimens he was describing from the Jurassic
Hope Bay locality in west Antarctica. Harris (1979)
later emended the diagnosis of Elatocladus to incor-
porate only coniferous shoots tha t bear linear,
univeined leaves that diverge from the stem and are
flattened to lie in the same plane. Although this re-
stricted to some extent the morphological scope of
the form-genus, it is still sufficiently broad to encom-
pass foliage as different as the rhythmically dimor-
phic shoots of Sequoia (Taxodiaceae), Cephalotaxus
(Cephalotaxaceae) and Prumnopitys (Podocarpaceae),
and the seasonally deciduous shoots of Taxodium and
Metasequoia (Taxodiaceae). Unfortunately, Halle’s
type species, Elatocladus heterophyllus, is quite dif-
ferent to most other species of the genus, which have
long, narrow, bifacial leaves that are twisted to give
the shoot a plagiotropic symmetry. Harris’ incorpo-
ration of a suite of species from the Jurassic of York-
shire also substantially expanded the geographical
extent of the genus, which was originally Gondwanan.
Florin and Harris clearly had different approaches to
the use of generic names. Florin preferred smaller
genera that were restricted in morphology, and he in-
stituted new genera whenever features were available;
Harris preferred broadly circumscribed form-genera
and in fact synonomized several of Florin’s genera
into Elatocladus (including one Florin had named for
Harris (“Tomharrisia”). The reason for our choice of
an emended Bellarinea in preference to Elatocladus
is to make the distinction between deciduous shoots
(Bellarinea) and persistent shoots with rhythmic
growth (most Elatocladus).
Some Gondwanan Me sozoic conifers with
plagio tr op ic shoots similar to Bellarinea and
Elatocladus have been included in the genera Mataia
and Rissikia. Mataia was erected by Townrow (1967)
for Jurassic podocarpaceous remains from New Zea-
land and northeastern Australia. It incorporates shoots
bearing hypostomatic to weakly amphistomatic leaves
with contracted bases in pseudodistichous arrange-
ment but the genus is defined mainly on reproductive
characters that are unavailable for most species of
Bellarinea and Elatoc ladus. Rissikia leaves are
rhombic in section with a band of stomates located
on each flank. This genus is further differentiated from
Bellarinea and Elatocladus by the presence of small,
spirally arranged, scale-like leaves at the base of the
shoot proceeded by larger leaves generally in
pseudodistichous arrangement. Rissikia species are
mostly represented in the Triassic of Gondwana and
have probable podocarpaceous affinities (Townrow
1967, Anderson & Anderson 1985).
Bellarinea richardsii sp. nov.
Fig. 2A-I
?1958 Elatocladus sp. cf. E. confertus Halle - Philip,
p. 192.
?1958 Elatocladus mccoyi Florin - Philip, p. 192.
1969 Elatocladus sp. ‘a’ - Douglas, p. 265.
cf. 1969 Elatocladus sp. ‘b’ - Douglas, p. 90, pl.
9, fig. 2.
1986 Rissikia sp. - White, pp. 176, 185, figs 268, 284.
1994 Elatocladus sp. - Douglas, p. 178, fig. 9.6a.
Holotype. MVP209942.
Paratypes. MVP209943-209957.
Type locality. Loc. L14 of Douglas (1969), near Exalt
Creek, Boola Boola State Forest, central Gippsland,
Vic to ria (A us trali an Map Gr id ref er ence
DT045457,578132).
Type formation and age. Locmany Member, Rintoul
Creek Formation, Tyers River Subgroup, Strzelecki
Group; Phyllopteroides laevis macrofloral zone
(Cantrill & Webb 1987); Neocomian.
Etymology. After Dr Max Richards, former board
member of the CSIRO and chair of the University of
Melbourne, School of Botany Foundation.
5BELLARINEA RICHARDSII FROM SOUTHEASTERN VICTORIA
Diagnosis. Determinate axes bearing up to 45 spi-
rally inserted leaves that are twisted at the base into a
pseudo-distichous arrangement. Leaves are univeined,
linear to inflexed falcate. Leaf apices are obtuse, com-
monly possessing a mucronate tip; leaf bases are de-
current. Leaf density is 4-10 per 10 mm. Epidermal
cells are rectangular with straight walls. Stomates
surrounded by 4-6 papillate subsidiary cells.
DESCRIPTION
Gross morphology. Coniferous shoots up to 84 mm
long, consisting of an unbranched, determinate axis
bearing up to 45 leaves diverging at 40°-90° to the
axis. Leaves are spirally inserted on the axis, but the
leaf bases are twisted to give the appearance of a
distichous arrangement. The leaves are linear to
inflexed-falcate, <16 mm long (typically 5-10 mm)
and 0.5-2 mm wide (average 1 mm). Leaf apices are
obtuse and commonly mucronate, leaf bases are de-
current, and leaf margins entire. Adjacent margins of
leaves on the same side of a shoot are 0.5-4 mm apart,
leaf density is 4-10 leaves per 10 mm and the leaves
very rarely overlap. The leaves have a midvein that
extends into the mucronate tip.
Foliar micromorphology. Epidermal cells are more
or less rectangular, and orientated along the axis of
the leaf forming a brick-like pattern (Fig. 2H, I). The
leaf surface has an uneven or slightly verrucate tex-
ture due to bulging of the periclinal walls of epider-
mal cells. Stomates are arranged in two longitudinal
bands on the abaxial leaf surface. Stomatal pores are
26-34 !m long and are surrounded by 4-6 weakly
developed, roughly circular papillae or lobes that are
9 !m in diameter.
Distribution. Localities L1, L4, L8, L14, L20, L23,
L24, L28, L30, LC1, LC3, RC17B, RC17C (see
Douglas 1969 for details); Localities RC17D, RC20A
(see McLoughlin et al. 2002 for details); Tyers River
Subgroup (Neocomian), Gippsland Basin, Victoria.
COMPARISON AND REMARKS
Morphology. There is considerable variation in leaf
dimensions and orientation among the specimens of
B. richardsii, however, all have a broadly similar ar-
chitecture consisting of shoots bearing bifacial,
univeined leaves with decurrent, twisted, but
unconstricted bases. In a few impressions the leaf
bases appear to be contracted but this is due to twist-
ing. Most leaves of Bellarinea richardsii are 1 mm
wide, linear or very slightly falcate and 2 mm apart.
This ‘typical’ condition was found in over 50 speci-
mens in the Boola Boola fossil flora (Fig. 2A, D, E,
F, G). There are several specimens with smaller (1.5-
4 mm long) and more closely spaced leaves (10 leaves
per 10 mm) but the shoots exhibit the same gross
morphology as the ‘typical’ specimens and are possi-
bly immature or under-developed shoots. Two other
unusual specimens have inflexed falcate leaves that
are spaced at 4 per 10 mm along the shoot (Fig. 2B),
compared to 6-8 leaves per 10 mm in the ‘typical’
shoots. The range of leaf arching may vary consider-
ably along a single shoot. In many cases, strongly
curved leaves are also relatively narrow (Fig. 2B).
The margins of extant Metasequoia glyptostroboides
leaves become recurved with desiccation and this
process may have been responsible for the arching,
twisting and enrollment of some B. richardsii leaves.
Complete shoots of Bellarinea richardsii are pre-
served in densely matted accumulations, which is
suggestive of a deciduous habit. The slender axes bear
leaves of nearly uniform length supporting the hy-
pothesis that the fossils represent detached short shoots
of a single season’s growth. The leaves along some
shoots of B. richardsii decrease in length distally (Fig.
2C), but the general condition is with leaves of roughly
the same length along the entire shoot. They are simi-
lar in appearance to the seasonally abscised shoots of
extant Taxodium distichum (L.) Richard. However,
detached Bellarinea-like leaves are also very com-
mon in the Boola Boola sediments suggesting that at
least some were shed before the shoots abscised. This
latter style of foliar detachment is similar to that of
extant Metasequoia glyptostroboides Miki ex Hu &
W.C. Cheng, which often sheds many of its leaves
before the shoots. There is no evidence of branching
on any of the shoots and none of the specimens pos-
sesses axillary or terminal buds. This favours the in-
terpretation that the shoots were seasonally shed units.
Affinities. The systematic affinities of Bellarinea
richardsii are unclear due to the lack of attached re-
productive organs and the difficulty in obtaining de-
finitive cuticular characters. Pollen from several coni-
fer families occurs in the Tyers River Subgroup
(Dettmann 1963). Araucariaceae is represented by the
pollen Araucariacites australis Cookson ex Couper.
However, this pollen is likely to be associated with
araucarian cone scales and small, slender twigs bear-
6NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN AND STEPHEN MCLOUGHLIN
7
ing appressed, scale-like leaves assigned to
Brachyphyllum tyersensis Tosolini & Nagalingum (in
McLoughlin et al. 2002). Brachyphyllum tyersensis
has wax-filled, obliquely orientated, cyclocytic
stomates with four to six subsidiary cells typical of
araucariacean leaves. Cheirolepidiacean pollen is rep-
resented by Corollina sp. cf. C. torosa (Reisinger)
Klaus. However, it is unlikely that B. richardsii is
cheirolepidiaceous as the leaves of that family are
typically scale-like with strongly sunken stomates
protected by prominent papillae, and borne in spi-
rals, whorls or opposite-decussate arrangement (Alvin
1982). Specimens attributed to Otwayia hermata
Tosolini & McLoughlin (in McLoughlin et al. 2002)
from the Boola Boola assemblage show these foliar
features and are the likely affiliates of the Corollina
pollen. Podocarpaceae is represented by pollen refer-
able to Podocarpidites sp. cf. P. ellipticus Cookson
an d Mi croc ac hyrid it es antarcticus Cookson.
Taxodiacean/cupressacean pollen is unknown from
these deposits and has only been regularly reported
in post-Cretaceous sediments in Australia (Macphail
et al. 1994). Nevertheless, taxodiacean macrofossils
are known from th e mid-Cretaceo us (P eters &
Christophel 1978) suggesting that the early pollen
record of this group has been overlooked on this con-
tinent. Small asulcate grains such as those referred
to Spheripollenites, although not recorded from Boola
Boola, are found elsewhere in the Victorian Early
Cretaceous, and these are not unlike the pollen of
extant Taxodiaceae. Florin (1963) argued that most
Elatocladus-type shoots from the Southern Hemi-
sphere, including Bellarinea, were probably
podocarpaceous but the dearth of consistently distinc-
tive architectural or cuticular characters separating
Taxodiaceae and Podocarpaceae foliage and the ab-
sence of reproductive remains associated with the
Boola Boola fossils prevents definitive familial as-
signment of B. richardsii.
Leaves of B. richardsii are basally twisted, which
results in a distichous appearance (Fig. 2F). This leaf
arrangement is common among some extant and fos-
sil Taxodiaceae (e.g., Metasequoia, Taxodium and
Sequoia) and Podocarpaceae (e.g., Falcatifolium,
Afrocarpus, Nageia, Retrophyllum, Prumnopitys,
Dacrycarpus, Acmopyle, Mataia, Smithtonia, and
Willungia). Those extant Taxodiaceae with linear
leaves in pseudodistichous arrangement are mostly
deciduous and have thin cuticle with either a smooth
surface (Taxodium: Alvin & Boulter 1974, Sung Soo
Whang & Hill 1999) or with a uneven surface caused
by lobing of the periclinal walls of epidermal cells
(Metasequoia: Qin Le ng et al . 2001). So me
Taxodiaceae lack Flo ri n rings (e.g., Sequoia,
Cunni nghamia) but others (e.g., Meta sequoia ,
Athrotaxus) have Florin rings that are typically lobed
(Oladele 1983). Stomata of linear-leafed Taxodiaceae,
such as Metasequoia, generally have apertures orien-
tated parallel to the leaf axis, strongly cutinized guard
cells, and around 4-6 subsidiary cells that are not
strongly differentiated from surrounding epidermal
cells. The cuticle of B. richardsii is thin, and very
diffi cult to pre pare compar ed to co- fossilize d
Bennettitales and other conifers; this thin cuticle is
further evidence for a deciduous habit. Its stomata
are axially aligned and the subsidiary cells are, at best,
only weakly raised. Some podocarp genera, such as
Falcatifolium and Retrophyllum, produce short shoots
with a pseudodistichous array of bifacial leaves
whereas others, such as Dacrycarpus and Acmopyle,
show similar twisting at the base of leaves but the
leaves are bilaterally flattened. Bellarinea richardsii,
with symmetrical bands of stomata restricted to the
abaxial surface appears to possess bifacial leaves.
Most Podocarpaceae have smooth cuticle but some
possess low papillae, irregular ridges, or the epider-
mal cells have an inflated appearance or verrucate
texture (Hill & Pole 1992). In many cases podocarps
have prominent Florin rings around the stomates but
in some cases (e.g., Smithtonia and Willungia) these
may be poorly developed or divided into irregular
lobes. All extant Podocarpaceae are evergreen, and
most have a relatively robust cuticle. Possession of a
distinct abaxial stomatal band on either side of the
midvein, longitudinally aligned, slightly inflated epi-
dermal cells, and weakly papillate subsidiary cells, is
co ns is tent with Flo ri n’s (19 52) place me nt of
Bel larinea in Podocarpaceae but an affinity to
Taxodiaceae can not be excluded.
Comparisons with other mid- to late Mesozoic coni-
fer fossils. Several conifer fossils with similar gross
morphology to B. richardsii have been recorded from
Australian Jurassic-Cretaceous strata (Table 1). These
have been variously assigned to the form-genus
Elatocladus, or to genera with implied cuticular or
reproductive affinities to modern conifer families. All
of these fossils differ to a greater or lesser degree to
the specimens here assigned to B. richardsii.
Although Douglas (1969) did not describe any
conifers from the Victorian Lower Cretaceous he listed
‘Elatocladus sp. a’ in the Boola Boola fossil flora.
These remains are probably conspecific with B.
richardsii given the appearance of Elatocladus re-
BELLARINEA RICHARDSII FROM SOUTHEASTERN VICTORIA
mains from this flora that he illustrated in later stud-
ies (Douglas 1994). Douglas (1969) also figured the
apex of a conifer leaf assigned to ‘Elatocladus sp. b’
from Boola Boola. However, this illustration does not
have sufficient detail to assess its affinity with E.
ric hardsii. As part of an investigation into the
sedimentology of the Tyers River Subgroup, Philip
(1958) listed the presence of Elatocladus sp. cf. E.
confertus and E. mccoyi, but these identifications were
not supported by illustrations or descriptions.
Stirling (1892, 1900) assigned Elatocladus-like
shoots from Aptian of the Gippsland Basin to Palissya
australis McCoy. These shoots exhibit rhythmic
growth of the leaves along their length, but differ from
B. richardsii by having either spirally arranged leaves
(Stirling 1892, Parris et al. 1995) or pseudodistichous
leaves borne on shoots with multiple branches (Stir-
ling 1900). At least some of these leaves also differ
from B. richardsii in having bilaterally flattened leaves
(see Parris et al. 1995, fig. 8a). The Palissya australis
specimens were later transferred to E. confertus by
Arber (1917) and considered synonymous with New
Zealand, Antarctic and Indian specimens (Arber 1917,
Sahni 1928). Medwell (1954) assigned several Vic-
torian specimens to E. confertus but they were not
figured or described and the specimens cannot be lo-
cated. Townrow (1967) later re-assigned Arber’s
specimens of E. confertus to Mataia podocarpoides
Townrow because they exhibited irregular branching
whereas the Antarctic type material has more or less
pinnate branching. Mataia podocarpoides is further
distinguished from B. richardsii by the possession of
attached ovuliferous cones (Townrow 1967).
Elatocladus mccoyi was described by Florin
(1952) based on an Aptian specimen from the Otway
Basin. The leaves of both B. richardsii and E. mccoyi
are borne spirally but twisted to give a distichous ap-
pearance. The lengths of some B. richardsii leaves
fall within the size range of E. mccoyi, however, the
two species are regarded as distinct because the leaves
of the former are typically linear to oblong and straight
to inflexed, whereas leaves of the latter are lanceo-
late to falcate and strongly reflexed. An additional
Aptian specimen from the Otway Basin was assigned
to Elatocladus sp. by Florin (1952) but this shoot has
spirally arranged, 2.5-4.5 mm long, triangular leaves
and should be transferred to Pagiophyllum or Otwayia.
Aptian twigs from the Gippsland and Otway ba-
si ns assig ne d to Bellarinea barklyi have
pseudodistichous leaves with decurrent bases (Florin
1952, Drinnan & Chambers 1986). This species has
bifacial leaves 14-32 mm long and 1.8-3 mm wide.
Most leaves of B. richardsii are slightly smaller than
those of B. barklyi. The stomates of both B. barklyi,
known only from the Aptian, and B. richardsii, re-
corded only from the Neocomian, are arranged in two
longitudinal bands but the former is distinguished by
having sinuous-walled epidermal cells in the non-
stomatiferous areas.
Elatocladus planus (Feistmantel) Seward has been
recorded from numerous Gondwanan Jurassic-Cre-
taceous localities (Townrow 1967). In Australia, E.
planus has been described from the Talbragar flora of
New South Wales (Walkom 1921), Algebuckina Sand-
stone, South Australia (Glaessner & Rao 1955) and
several Mesozoic formations of Queensland (Walkom
1917, 1919). It is not clear that all, or any, of the Aus-
tralian forms referred to this species are conspecific
with the Indian type material. Townrow (1967) noted
that this species ‘almost certainly will prove to be
composite’ upon further investigation. This sugges-
tion proved correct in the case of the ?Middle Jurassic
Talbragar specimens, which were later reassigned to
Rissikia talbragarensis White 1981. The Talbragar
species differs from both E. planus and B. richardsii
on the basis of its more closely spaced, longer leaves
with distinct transverse striations (White 1981).
Townrow (1967) excluded the South Australian speci-
me ns fr om hi s synon ym y lis t for E. planus .
McLoughlin (1996) suggested that the South Austral-
ian E. planus (Glaessner & Rao 1955) specimens may
be synonym ou s wi th Elatocladus ginginensis
McLoughlin but could not confirm their identity as
the former are ill-preserved. Bellarinea richardsii is
broadly similar in morphology and size to Elatocladus
ginginen sis McLo ughli n from the Neocomia n-
Barremian of Western Australia (McLoughlin 1996).
The latter has leaves with variably contracted or de-
current bases. Its leaves are distally tapered but lack
a mucronate tip. In contrast, B. richardsii has leaves
with a mucronate apex and decurrent bases that are
not contracted. Elatocladus-like shoots originally
described by Walkom (1918) from the Lower Creta-
ceous of Queensland have subsequently been assigned
to a new species, E. baddowensis (McLoughlin et al.
2000), that is distinguished by its oblong to lanceo-
late leaves with rounded apices and slightly contracted
bases.
Species of Elatocladus described from the York-
shire Jurassic (Harris 1979), although similar in gen-
eral leaf morphology to Bellarinea richardsii, all rep-
resent indeterminate shoots and exhibit either branch-
ing or buds, both of which are absent from the Boola
Boola specimens. Several species of Elatocladus de-
8 NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN AND STEPHEN MCLOUGHLIN
Table 1. Comparison of conifer fossils with similar gross morphology to Bellarinea richardsii from Mesozoic sediments of Australia, India
and New Zealand. Information from Walkom (1921), Sahni (1928), Florin (1952), Townrow (1968), White (1981), McLoughlin (1996) and
McLoughlin et al. (2000).
9
BELLARINEA RICHARDSII FROM SOUTHEASTERN VICTORIA
scribed from the Jurassic and Early Cretaceous of
Antarctica (Halle 1913, Gee 1989, Cantrill 2000a,b,
Cantrill & Falcon-Lang 2001) differ from B. richardsii
by being branched and indeterminate. Of these, E.
australis has lateral shoot units subtended by a rest-
ing bud of scales, which Cantrill (2000a) suggested
may be evidence of deciduousness or at least arrested
growth. The latter is most likely given that Cantrill’s
figure 7.1 shows a shoot of rhythmic growth with large
leaves either side of a region of restricted growth
Associated plant fossils. Bellarinea richardsii is
by far the most abundant of the three conifer foliage
species in the Boola Boola macroflora, but this may
reflect a deciduous habit rather than floristic domi-
nance. The assemblage also contains several other
gymnosperms including bennettitaleans (Otozamites
spp.), pentoxylaleans (Taeniopteris daintreei McCoy),
and pteridosperms of uncertain affinity (Rintoulia
variabilis, Pachydermophyllum austropapillosa and
Komlopteris indica; McLoughlin et al. 2002). Ferns
are represented by around ten foliage-based species
(Douglas 1973) and lycophytes are represented by
abundant detached microphylls and 16 species of
megaspores (McLoughlin et al. 2002). Bellarinea
ri ch ard si i is most commonly associated with
Taenio pteris daint reei, Rintoulia variabilis and
Otozamites spp. in floodbasin siltstones and crevasse-
splay sands (McLoughlin et al. 2002). Based on their
abundance in these sediments, their taphonomy, as-
sociated fossils, and previous palaeoecological inter-
pretations of this plant group (Douglas & Williams
1982), Bellarinea richardsii is interpreted to have
constituted an upper storey tree. It occupied relatively
moist floodbasin environments in alluvial valley sys-
tems during the initial establishment of the Gippsland
Basin rift.
ACKNOWLEDGEMENTS
This study was supported by an Australian Research
Council Large Grant for the investigation of Austral-
ian Mesozoic floras. N.S.N. was supported by a Mel-
bourne Research Scholarship.
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12 NATHALIE S. NAGALINGUM, ANDREW N. DRINNAN AND STEPHEN MCLOUGHLIN
E.
talbragarensis
Characters
Shoot
length
Leaf
arrangement
Maximum
leaf length
Maximum
leaf width
Leaf shape
Leaf
orientation
Leaf
cross-section
Leaf base
Leaf apex
Stomatal
distribution
Arrangement
of stomates
Subsidiary
cells
Guard cells
Papillae
Epidermal
cells
Affiliated
reproductive
structures
Type
formation
or locality
Age of type
formation
127 mm
spiral; basally
twisted to
pseudodistichous
c. 60 mm
2 mm
linear; in some
cases slightly
reflexed or
inflexed
20-90° from axis
uncertain;
probably keeled
in most cases
markedly
decurrent
(specimens with
transverse striae
on leaves here
excluded)
blunt obtuse or
rounded
not available
not available
not available
not available
not available
not available
B. barklyi
c. 100 mm
spiral; basally
twisted to
pseudodistichous
32 mm
3 mm
linear to
lanceolate
40-90° from axis
uncertain;
probably keeled
slightly
contracted,
non-petiolate,
twisted, strongly
decurrent
subacute to
obtuse
hypostomatic;
0.5 mm broad
stomatiferous
band in centre
of lamina either
side of midvein
haplocheilic
stomates with
mostly oblique
apertures in
weakly defined
longitudinal rows
4-6; never shared;
slightly papillate;
slightly raised to
form weak Florin
ring
slightly sunken
present only on
subsidiary cells
generally
rectangular or with
slightly sinuous
anticlinal walls
no obvious
associated fruits
lower Eumeralla
Formation,
Otway Basin,
Aptian
associated with
120 mm long
cones comprised
of c. 30 spirals of
cone scales each
bearing a
?solitary seed
Purlawaugh
Formation,
Surat Basin,
?Bajocian-
?Callovian
E. ginginensis
51 mm
spiral; basally
twisted to
pseudodistichous
20 mm
1 mm
linear; in some
cases slightly
reflexed or
inflexed
35-60° from axis
uncertain;
probably keeled
slightly
contracted,
non-petiolate,
twisted, slightly
decurrent
rounded or blunt
obtuse
not available
not available
not available
not available
not available
not available
no obvious
associated fruits
Leederville
Formation,
Neocomian-
Barremian
E. mccoyi
>25 mm
spiral; basally
twisted to
pseudodistichous
11 mm
1 mm
linear; generally
reflexed
50-75° from axis
uncertain;
probably keeled
slightly
contracted,
twisted, broadly
decurrent
acute to almost
obtuse
not available
not available
not available
not available
not available
not available
no obvious
associated fruits
lower Eumeralla
Formation,
Otway Basin,
Aptian
40-80° from axis
M.
podocarpoides
30 mm; shoots
commonly
retained in
connection with
parent shoots
spiral; basally
twisted to be more
or less distichous
except near base
of ultimate shoots
where they
remain spiral
15 mm
4 mm
oblanceolate;
commonly
reflexed
triangular;
strongly keeled
below midvein
tapered; markedly
decurrent
variable: rounded,
acute, or slightly
mucronate
hypostomatic or
very unequally
amphistomatic;
stomata restricted
to zones in
uncertain position
monocyclic;
arranged in
longitudinal,
though often
irregular rows
apertures mostly
longitudinally
orientated
4-6; never shared
but sometimes
adjacent; bearing
hemispherical
papillae; Florin
ring absent
slightly sunken
generally present
only on
subsidiary cells
mostly rectangular
with slightly
sinuous anticlinal
walls
seed cone spike-
like; 30 mm long;
bearing 8-12
spirally arranged
units comprising a
bract and axillary
ovuliferous scale
bearing two
stalked seeds
Clent Hills,
New Zealand
?Middle Jurassic
E. planus
linear; commonly
reflexed
not available
not available
not available
not available
not available
not available
no obvious
associated fruits
26 mm
spiral; basally
twisted to
pseudodistichous
>90 mm
1.2 mm
60-90° from axis
uncertain;
probably keeled
not significantly
contracted,
strongly
decurrent
acutely pointed
to rounded
?Barremian-
?Aptian
Vemavaram
Formation
Cauvery-Palar
Trough, India
E.
baddowensis
>41 mm
spiral; basally
twisted to
pseudodistichous;
opposite-
subopposite
17 mm
2.5 mm
oblong to
lanceolate;
straight or slightly
inflexed
45-80° from axis
uncertain;
probably flattened
slightly contracted;
weakly decurrent
rounded or blunt
obtuse
not available
not available
not available
not available
not available
not available
no obvious
associated fruits
Maryborough
Formation,
Maryborough
Basin, Australia
Aptian
B. richardsii
10 mm
84 mm
spiral; basally
twisted to
pseudodistichous
16 mm
2 mm
linear;
straight, inflexed
or reflexed
40-90° from axis
uncertain
twisted, broadly
decurrent
obtuse,
commonly
mucronate
no obvious
associated fruits
Rintoul Creek
Formation,
Gippsland Basin,
Neocomian
hypostomatic;
arranged in at
least two
longitudinal
files; apertures
longitudinally
orientated
papillate
sunken
present, only on
rectangular
subsidiary cells
probably flat or
slightly keeled
0.3 mm broad
stomatiferous
band in centre
of lamina either
side of midvein
Australia Australia Australia Australia Australia
Western
