The Middle Triassic Megafossil Flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales, Australia. Part 5. The Genera Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris

2005·17.2 MB·English

by  H M Anderson

The Middle Triassic Megafossil Flora of the Basin Creek
Formation, Nymboida Coal Measures, New South Wales,
Australia. Part 5. The Genera LepidopteriSy Kurtziana,
Rochipteris and Walkomiopteris,
W.B.Keith Holmes' and Heidi M. Anderson^
'46 Kurrajong Street, Dorrigo NSW 2453, Australia (Hon. Research Fellow, University ofNew England,
Armidale NSW 2351); H6 Kurrajong Street, Dorrigo NSW 2453, Australia (Hon. Palaeobotanist, National
Botanical Institute, Pretoria 0001 SouthAfrica).
Holmes, W.B.K. andAnderson, H.M. (2005). The Middle Triassic megafossil flora ofthe Basin
Creek Formation, Nymboida Coal Measures, New South Wales,Australia. Part 5. The genera
Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris. Proceedings oftheLinnean Society
ofNew South Wales 126, 39-79.
Leaves attributed to the gymnosperm genera Lepidopteris, Kurtziana andRochipteris and the
leafsedis incertae Walkomiopteris eskensis (Walkom) Holmes andAnderson gen. et comb. nov.
are described from two quarries in the Basin Creek Formation ofthe Middle Triassic Nymboida
Coal Measures. Based on extensive collections ofleaves, the morpho-genera Lepidopteris and
Kurtziana each reveal a wide range of variation. Lepidopteris is divided into three 'morpho-
species complexes' with intergrading forms based on L. madagascariensis, L. africana and
L. stormbergensis and a new species L. dissitipinnula apparently without links to the three
complexes. The Lepidopteris fertile organs, Peltaspermum and Antevsia are present. Kurtziana
K
is separated into two 'morpho-species complexes' based on K. brandmayri and cacheutensis.
Rochipteris is a diverse genus but ofvery rare occurrence. Six new species are described; R.
obtriangulata andR. tubata display a close-spaced spiral orwhorled arrangement. Seven leaves
ofR. incisa have been examined. R. sinuosa, R. pusilla and R. nymboidensis are represented by
single dispersed leaves. Walkomiopteris eskensis (Walkom) gen. et comb. nov. is redescribed
from additional Nymboida material.
KEYWORDS: Kurtziana, Lepidopteris, Middle Triassic Flora, Nymboida Coal Measures,
Rochipteris, Walkomiopteris.
family fi-om two Nymboida quarries over a period of
INTRODUCTION
almost forty years. Details ofthe Coal Mine Quarry
and the Reserve Quarry together with a summary
In this fifth part of a series describing the early of the geology of the Basin Creek Formation, the
middle Triassic Nymboida flora, leaves assigned to Nymboida Coal Measures and the Nymboida Sub-
three gymnosperm genera: Lepidodopteris with some basin were provided in Holmes (2000).
associated fertile organs, Kurtziana, Rochipteris and
the new genus Walkomiopteris sedis incertae are
illustrated and described. METHODS
Part 1 (Holmes 2000) ofthis series deah with the
Bryophyta and Sphenophyta, Part 2 (Holmes 2001) Inlivingpopulationsofextantplantsthereisoften
with the Filicophyta, Part 3 (Holmes 2003) with a large range ofvariation in leafshape, e.g. juvenile,
fern-like foliage and Part 4 (Holmes and Anderson adult, shade, sun-leaves, etc. Apreserved holotype in
in press) with the genus Dicroidium and its fertile a herbarium seldom exhibits this range ofvariation.
organs Umkomasia andPteruchus. It is only through wide experience in the field that
The material described below is based mainly this variation can be recognised and appreciated. In
on the collections made by the senior author and his the fossil record, leaves constitute the vast majority
1
TRIASSIC FLORA FROM NYMBOIDA SOME GYMNOSPERM GENERA
-
ofpreserved plant organs. Most early palaeobotanical in the 'natural genus' Meyenopteris is untenable. The
taxonomy was based on very limited material in the leal"species Lepidopteris storml^ergensis has priority
field or a few museum speeimens. Original diagnoses over L. natalensis. An additional leaf and ovulate
rarely acknowledged the variation that could occur species have been described from the same Umkl
1
in a 'natural' species. To compound the problem locality (Anderson and Anderson 2003). Until fruit,
of separating fossil leaves into 'natural' species, leaves and stems are found in organic connection, it
the assemblages from one locality may represent is premature to erect a 'natural genus'. In accordance
vegetation from a range of habitats and growing with ICBN (2001) Articles 1.2 the morpho-genus
throughanunknownperiodoftime.Asin Part4ofthis Lepidopteris should be retained for dispersed
series, which dealt with the highly variable morpho- leaves and the morpho-genera Peltaspermum and
genus Dicroidiiim (Holmes and Anderson in press), Antevsia for the dispersed female and male organs
we address the problem ofvariability observed in the respectively.
largecollectionsofLepidopterisandKurtzianaleaves Leaves of the Lepidopteris genus occur in the
by creating 'species complexes'. A 'species complex' Permian of the Northern Hemisphere. The first
includes leavesdisplayingarangeofvariationcentred Gondwanan record is ofL. callipteroides, a branched
on a previously-described species. Intergrading forms leaf form from the basal Narrabeen Group (earliest
often link the 'complexes'. The Selected References Triassic), of the Sydney Basin (Retallack 2002).
are of specimens we consider to represent a mid- This species apparently did not persist through to the
range for each 'species complex'. Leaves illustrated Middle Triassic.For many Gondwanan Lepidopteris
in the Figures should enable comparisons to be made leaves the application ofspecific names has at times
with material from other locations and horizons. beenquestionable. Some speciesareknownonly from
Leaves in the generaRochipteris and Walkomiopteris impressionswhilesome,withbetterpreservationhave
are represented respectively by ten and six specimens been described with cuticle information (Carpentier
only, so are placed in morpho-species based on gross 1935; Townrow 1960, 1966; Baldoni 1972; Baldoni
morphology ofthe material available. and de Cabrera 1977; Anderson andAnderson 1989).
At the Nymboida quarries most specimens are However, as noted by Townrow (1966), there are
preserved as carbonaceous compressions in which problems ofidentification ofspecimens both with or
the gross morphology is usually well-preserved. without cuticle. Rigby (1977) suggested reservingthe
However, spores and cuticles have been destroyed by name L. stormbergensis for all leaves lacking cuticle
a tectonic heating event during the Cretaceous Period and L. natalensis and L. madagascariensis for those
(Russel 1994). withpreservedcuticle, while Retallack in Retallacket
Type and illustrated material is housed in the al. (1977) arguedthat the diverserange ofleaves from
Australian Museum, Sydney. Some additional theCloughers CreekFormationintheNymboidaCoal
specimens are in the collections of the Geology Measures was best placed in L. madagascariensis on
Department, University of New England, Amiidale, thebasis ofthethick leafsubstance and mostly obtuse
NSW, and the type of Walkomiopteris eskensis is pinnules although no cuticle was present. We regard
housed in the collections ofthe Queensland Museum, the Cloughers Creek leaves as being best placed in
Brisbane. our 'L. stormbergensis'' and 'L. africana complexes.
'
In the Nymboida collections, Lepidopteris leaves
are preserved on c. 3% of the catalogued slabs.
SYSTEMATIC PALAEOBOTANY While individual leaves may be identified on gross
morphology with the types ofL. madagascariensis,
Ginkgoopsida S.V.Meyen 1987 L. africana or L. stormbergensis, there are numerous
Peltaspermales F. Nemejc 1968 intergrading forms that link these three 'species'. The
Peitaspermaceae H.H. Thomas ex T.M. Harris 1937 same problem was noted by Holmes 982) for the
(1
Genus Lepidopteris Schimper 1869 Benolong Flora where leaves of L. stormbergensis,
Type species Lepidopteris stutgardiensis Schimper L. africana and L. mortonii formed an intergrading
1869 series. Anderson and Anderson (1989) noted that
at II of their 30 localities where both L. africana
The proposal by Poort and Kerp (1990) to unite and L. stormbergensis were present, in most cases
theleavesofLepidopteris 'natalensis withtheovulate they fonned an unbroken morphological range and
'
organ Peltospermiim thottiasii, which occur together were regarded as one palaeodeme. At the remaining
at the Waterfall locality in the Molteno Fonnation 19 localities only one or other of the species was
(locality Umklll ofAnderson and Anderson 1983), present.
40 Proc. Linn. Soc. N.S.W., 126, 2005
,
W.B.K. HOLMES AND H.M. ANDERSON
As our collections of Lepidopteris have been Description
made from the various sedimentary facies in the Small broad-elliptic bipinnate leaves c. 50-150
mm mm
two Nymboida quarries they were probably sourced long, c. 40-70 wide, leafbase truncate,
mm
from differing vegetation types (Retallack 1977; main rachis 2 wide and tapering to apex,
Holmes 2000) and the Lepidopteris material may sometimes punctate and/or longitudinally striate;
indeed belong to several true species each specific c. 20 pairs ofwell-separated opposite to alternate
to one palaeodeme. However, on our present state straight or arching pinnae, decreasing in length
of knowledge we must accept the collection as basally and apically, are attached at a high angle
representing a single variable population sample. To towards base, at c. 60° in mid-frond and more acute
enable the Basin Creek material to be compared with apically; pinnules not conjoined, oblong, truncate to
that from other localities, we place the Lepidopteris obtusely rounded, attached by whole base to pinna
leaves in three 'species complexes' while noting rachis at c. 60°; first basiscopic pinnule decurrent;
the intergrading forms that link the complexes. A with one or more pirmules attached laterally on
distinct leaf-type with widely-spaced pinnules and rachis between pinnae. These latter are generally
no intergrading links with the three 'complexes' is referred to as 'zwischerfiedem'.
erected as a new morpho-species.A notable feature
of Lepidopteris leaves, with and without preserved Material
cuticle,istheusualpresenceof'blisters'or 'lumps'on AMF126801-3, AMF126805 Coal Mine
AMF
the main and/or pinna rachis (Townrow 1956, 1960, Quarry; 126804 Reserve Quarry
1966; Holmes 1982; Anderson and Anderson 1989,
2003)resultingfromaproliferationofepidermal cells Discussion.
around trichoma bases. Townrow (1960) noted that Typical leaves ofthis complex are not as
in a population of leaves attributed to Lepidopteris numerous as those in the 'L. africana complex'. It
stormbergensis there was a series of leaves with is distinguished from the two complexes below by
rachises ranging from smooth to markedly blistered. the oblong pinnules with obtuse apices separated to
Lumps or blisters are not apparent on the Nymboida the base on the mid-frondpinnae and by the higher
Lepidopteris leaves but some do have punctate or angle ofattachment ofthese pinnules. However,
striate rachises. Townrow (1966) described the main basal and apical pinnae often have coalescing to
rachis of Lepidopteris as having a wing and with coherent pinnules. Figure 2Ashows, on the same
dorsally-attachedpinnae.Arachis wing is notevident slab, portions ofseveral leaves that obviously
in theNymboidamaterial andthe pinnae appear to be represent a single population (palaeodeme). One leaf
attached laterally. is typically 'madagascariensis' while others show
pinnules becoming coherent and grading into the
'Lepidopteris madagascariensis complex', based on form ofthe 'Z. africana complex'.
L. madagascariensis Carpentier 1935
Figures 1A,B; 2A-C
Selected references ''Lepidopteris africana complex', based onL.
1935 Lepidopteris madagascariensis, Carpentier, africana (Du Toit 1927) Holmes 1982
P1.3, figs 3,4 Figures 2D; 3A,B; 4A,B; 5A,B
1936 Lepidopteris madagascariensis, Carpentier,
P1.5, fig.4 Selected references.
1966 Lepidopteris madagascariensis, Townrow, 1927 Callipteridium africana, Du Toit, PI.27
Textfig. IE, PI.1, fig.1 1944 Callipteridium argentinum, Frenguelli, PI.1
1975 Lepidopteris madagascariensis, Flint and figs 1,2
Gould, P1.2, figs 1,2 1965 Lepidopteris stormbergensis. Hill et al., P1.T6,
1979 Lepidopteris madagascariensis. Holmes and fig.l
Ash, Fig.5.6, 5.7 1977 Lepidopteris madagascariensis, Retallack et
1983 Lepidopteris madagascariensis, Retallack, al., fig.9D
Fig.5A 1982 Lejp/Joptera (s/r/cawfl. Holmes, Figs 8C, 8D
1995 Lepidopteris madagascariensis, Retallack, 1983 Lepidopteris africana, Anderson and
Figs 2A, 3A Anderson, PI.13, fig.l
1989 Lepidopteris africana,Anderson and
Anderson, p.92, figs 1-3, P1.13, figs 1-10,
P1.43,figs 1-16
Proc. Linn. Soc. N.S.W., 126, 2005 41
2
TRIASSIC FLORA FROM NYMBOIDA SOME GYMNOSPERM GENERA
-
1998 Lepiclopleris mcidagascariensis, Gnaedinger 1975 Lepidopteris stormbergensis, Flint and Gould,
and Herbst, figs I4a-c P1.2, figs 1,2
2001 Lepiclopleris mcidagascariensis, Gnaedinger 1977 Lepidopteris madagascariensis, Retallack et
and Herbst, fig. 1 la al. fig. 9A
2003 Lepidopteris africana, Anderson and 1982 Lepidopteris stormbergensis. Holmes, fig.8A
Anderson, p.157, fig.1 1983 Lepidopteris stormbergensis, Anderson and
Anderson, PI.13, figs 2,3
Description 1989 Lepidopteris stormbergensis, Anderson and
Small to medium-sized bipinnatifid leaves 120- Anderson, p.93, figs 1-4, P1.26, figs 2-5, PI. 27,
mm mm
>170 long, 25-70 wide, with a truncate figs 1-4
mm
leaf-base 4 wide, tapering gradually to apex; 1998 Lepidopteris madagascariensis, Gnaedinger
pinnae closely spaced, longest at 2/3 ofleaflength and Herbst, PI.3, fig.h, figs 14a,c
where attached at c. 45° to main rachis, apically the 2003 Lepidopteris stormbergensis, Anderson and
pinnae decrease in length and become more acute, Anderson, p.157, fig.4
basally the pinnae have a higher angle ofattachment,
become shorter, with rounded apices and entire to Description.
undulate margins; pinnae with coherent pinnules Large bipinnate to tripinnatifid leaves, broad-
mm mm
forming a serrate margin; the basiscopic base ofthe oblanceolate, to 400 long and 180 wide.
mm
pinnae strongly decurrent along the main rachis to Rachis to 5 in mid-frond; pinnae opposite to
the acroscopic base ofthe pinna below, leaving no alternate, longerpinnae at mid-frond attached at c.
space for zwischerfiedem. 80°-45°, closely spaced to overlapping on larger
and tripinnatifid fronds; pinnules on mid-portion of
mm mm
Material mid-pinnae 6-25 long, 3-6 wide, tapering
AMP AMF
126806 Reserve Quarry; 126807-1 to acute ornarrow obtuse apex, margin entire to
Coal Mine Quarry serrate. On the largest leaves (Figures 8B, 9B)
the pinnules are deeply lobed to pinnatisect. First
Discussion basiscopic pinnule attached to base ofpinna rachis
The leaves illustrated in Figures 3A,B and or strongly decurrent on main rachis; nil to three
4A were exposed on one bedding plane and surely zwischerfiedem between pinnae on main rachis in
represent a single population (palaeodeme). Figure mid-portion ofleaf
4B shows anotherbeddingplane assemblage showing
many 'Z. africana' leaves of varying size together Material
with a fragment of a leaf with larger separated AMF126816-21,AMF126851, all Coal Mine
pinnules that approaches 'L. stormbergemis' but Quarry.
with pinnules coalescing distally and apically. Large
leaves with pinnules becoming less coherent form Discussion
AMF
intergrading links between X. africana complex' and The leafassemblage preserved on 126819,
AMF AMF
X. stormbergensis complex' (Figs 5C; 6A,B; 7A). 126821 and 126851 are parts and
counterparts from the same bedding plane and show
fronds ranging from bipinnatifid to tripinnatifid and
''Lepidopteris stormbergensis complex', based onX. include the largest in the collection (Figure 8B).
stormbergensis (Seward 1903) Townrow 1956 Zwischerfiedem preserved on the tripinnatifid leaf
Figures 6C; 8A,B; 9A,B (Figure 9B) are broad-elongate with a lobed margin.
This assemblage demonstrates the large range of
Selected references. variation even within a single population.
1903 Callipteridium stormbergense, Seward, PI.7,
fig.5 Lepidopteris dissitipinnula Holmes andAnderson
1927 Lepidopteris stiittgardensis, Du Toit, PI.28 sp. nov.
1956 Lepidopteris stormbergensis, Townrow, figs Figures lOA-C
lA, IB
1960 Lepidopteris stormbergensis, Townrow, text Diagnosis
A
figs 5C,F,G medium-sized Lepidopteris leafwith sub-
1965 Lepidopteris stormbergensis. Hill et al., P1.T6, opposite slightly arching to recurved pinnae;
fig.2 pinnules well-spaced, elongated-linear with obtuse
42 Proc. Linn. Soc. N.S.W., 126, 2005
W.B.K. HOLMES AND H.M. ANDERSON
apices, margins parallel, entire to lobate. The ovulate organ Peltaspermum had a wide
LaurasianandGondwanan distribution. Itis generally
Description accepted as the female fructification ofthe plant that
Based on two specimens, both with base and hoxQLepidopteris leaves (Thomas 1933; Harris 1937,
apices missing, length preserved to 110 mm; rachis Townrow 1960, Anderson and Anderson 2003).
mm
at base of preserved section 2-3 wide; pinnae Poort and Kerp (1990) revised the Peltaspermum -
mm
elongate-lanceolate, alternate, c. 12 apart, longest Lepidopteris associationbasedonwesternandcentral
pinnae to 65 mm. Specimen AMFl13528 (Figures European material. They proposed the creation of
10A,B) has slightly recurved pinnae attached at c. the 'natural genus' Peltaspermum by emending the
60"; AMFl26823 (Figure IOC) has lower pinnae diagnosis of Peltaspermum to include Lepidopteris
attached at right angles and arching slightly and the leaves. As Peltaspermum and Lepidopteris are both
upperpinnaeattachedatc. 60°,butthismayalsobean morpho-taxa under ICBN (2001) Article 1.2 a new
artifact ofpreservation. Pinnules opposite, spaced c. name would be required for a 'natural genus'.
onepinnulewidth apart, decurrent, straightorslightly Despite the large number ofLepidopteris leaves
arched, apex rounded-obtuse, margins parallel, entire in the Nymboida collection, Peltaspermum is known
to lobate. Pinnules are longest at mid-pinna, to 12 only from two incomplete strobili and two detached
mm mm
long, 1-2 wide, decreasing in length basally peltate discs.
and apically, basal basiscopic pinnule not decurrent
on main rachis; one or two pairs ofnarrow, elongate Peltaspermum cfmonodiscum Anderson and
zwischerfiedem on main rachis between pinnae. Anderson 2003
Figures llA-E
Holotype
AMFl13528 and counterpart AMFl13529 Description
Australian Museum, Sydney. Based on two incomplete sfrobili. Axes as
mm mm mm
preserved c. 40 and 25 long, 2 wide;
mm mm
Type Locality six discs c. 4 in diameter attached at 8-10
Coal Mine Quarry. Basin Creek Formation, intervals singly or opposite each other by a peduncle
mm mm
Nymboida Coal Measures, Middle Triassic. c. 5 long. Each disc is c. 6 wide, pendant,
mm
showing four decurved, linear lobes 4 long,
mm
Other material 1 wide. As the fossils represent sideways-
AMFl26823 Coal Mine Quarry. compressed discs, the number oflobes in life would
be eight. A single detached disc and its counterpart
Name Derivation (Figures IID, E) show a radially symmetrical disc c.
mm
dissitus, (Lat.) well-spaced; referring to the 9 in diameter with ten linear lobes around the
mm mm
well-separated pinnules. circumference, each c. 1 wide and 2-3 long.
A possible peduncle protrudes from one side of the
Discussion disc but its point ofattachment is uncertain.
Lepidopteris dissitipinnula differs from all
described Lepidopteris morpho-species by the Material
elongated-linear well-spaced pinnules and perhaps AMF126824-6 Coal Mine Quarry.
is closest to L. madagascariensis. However, at
Nymboida there are no intergrading forms to link Discussion
L. dissitipinnula with the 'Z. madagascariensis Anderson andAnderson (2003 pp. 152, 158, 159)
complex'. Both specimens of L. dissitipinnula are describedandillustrated fromthe Molteno Formation
preserved in a white sandstone matrix in contrast to of South Africa some reasonably intact strobili with
all otherLepidopteris, material which is preserved in lobed receptacles attached singly to an axis. They
black to grey shales and mudstones, thus suggesting refer to the receptacles as 'discs'. Their specimens
they were sourced from ecologically separated have 11 or 12 lobes. The Nymboida material with
populations. 8-10 lobes is otherwise closely comparable.
Genus Peltaspermum Harris 1937 Peltaspermum spA
Type species. Peltaspermum rotula Harris Figures 11F,G
1937
Proc. Linn. Soc. N.S.W., 126, 2005 43
TRIASSIC" FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA
Description whether the sporangial sacs are sessile or shortly
One specimen and its counterpart shows a pedunculate. The second specimen, AMF126829
mm
spherical disc 18 in diameter with c. 13-14 (Figure 12C), is of two detached clusters and some
mm
broad obtuse lobes around the margin, each separated scattered sporangial sacs to 5 long. Associated
by an incision or ridge reaching from half to two with the sporangia is a detached oval-shaped
thirds distance to the centre, which is marked with indeterminate ovule.
mm
an irregular-shaped abcission scar c.1.8 in
diameter. Material
AMF AMF
126828 and 126829 Coal Mine
Material Quarry.
AMF AMP
26852 and counterpart 26853 Coal
1 1
Mine Quarry. Discussion
Antevsia sp. A has some similarities to Antevsia
Discussion mazenodensis Anderson and Anderson (2003) from
Peltaspermum sp. A differs from P. cf. the Molteno Formation but the preservation is not
monodisciim by the larger size and less incised lobes. sufficient for specific determination.
Similardetachedpeltoiddiscshaveawidedistribution
and are associated in Gondwana deposits with the Order Matatiellales Anderson and Anderson 2003
peltaspermaceous genera Lepidopteris (Harris 1937; Family Matatiellaceae
Holmes and Ash 1979; Holmes 1982, Anderson Genus ATi/r/z/owo Frenguelli 1942a
and Anderson 2003) and Scytophyllum (Zamuner Type species Kiirtziana cacheutensis (Kurtz)
et al. 1999) and in the Northern Hemisphere with Frenguelli 1942a
Lepidopteris, Tatarina, Comia, Pachydermophyllum
and Scytophyllum (Meyen 1987). This isolated In frond morphology and venation pattern
Nymboida disc has insufficient diagnostic features to Kurtziana differs from all other Gondwanan
place it in any known species. ginkgoopsid leafgenera. Based on mutual occurrence
Anderson and Anderson (2003) have given a Grade
Genn^Antevsia Harris 1937 2 affiliation of Kurtziana leaves with the female
Type species Antevsiazeilleri (Nathorst) Harris strobilis Matatiella and placed the leaf genus in the
1937 Order Matatiellales.
The genus Kurtziana was erected by Frenguelli
Antevsia strobili have been recorded from (1942a) for unforked pinnate leaves with pinnae
Rhaetic localities in Sweden (Antevs 1914), having contracted pinna bases and attached laterally
Greenland (Harris 1932) and from the UpperTriassic to the rachis. These leaves from Argentina were first
Molteno Formation of South Africa (Anderson and illustrated by Kurtz (1921, PI. 16, figs 198-199) as
Anderson 2003). Antevsia has been linked at those Daneacacheutensis.Asecondspecies,K. brandmayri
occurrences with Lepidopteris on the basis ofsimilar Frenguelli (1944), was erected forleaves inwhichthe
cuticles (Antevs 1914; Harris 1932) and the same pinnae were closely-spaced to imbricate and attached
A
distinctive blistering on the strobilis axes as on the to the dorsal surface of the rachis. very large
foliar rachises (Anderson andAnderson 2003). leaf ft-om Chile has recently been described as K.
paipotensis Herbst andGnaedinger(2002). Kurtziana
Antevsia sp A leaves with preserved cuticle have been described by
Figures 12A-C Petriella and Arondo (1982) andArtabe et al. (1991).
Herbst and Gnaedinger (2002) have erected the new
Description morpho-genus Alicurana for Kurtziana leaves with
Two fragmentary specimens from Nymboida preserved cuticle. To date Kurtziana is best known
AMF
show clusters of sessile sporangia. 126828 from South American localities.
(Figures 12A,B) is a portion ofa strobilis overlain by Kurtziana leaves are also known fi-om South
a fragment ofa Sphenobaiera leaf. The curved axis, Africa (Du Toit 1927; Anderson andAnderson 1983).
mm
which maybe an almost completebranch, is c. 60 In a recent publication, Anderson and Anderson
mm mm
long, 1.4 wide at the base andtapering to 0.8 (2003) illustrated five species and noted the presence
distally. Blisters are not apparent. Clusters of up to of 16 species, generally ofrare occurrence, from the
mm
five irregularly elliptic microsporangia to 2 long Molteno Formation ofSouth Africa. From Australia,
are scattered along the branch axis. It is not certain the leaf"Thinnjeldia 'eskensis Walkom (1928) is here
44 Proc. Linn. Soc. N.S.W., 126, 2005
W.B.K. HOLMES AND H.M. ANDERSON
transferred to Kurtziana cacheutensis. overlapping oblong pirmae with obtuse apices and
Kurtziana leaves are represented on c. 2% of constrictedauriculatebases attachedatahighangleto
catalogued slabs in the Nymboida collection. They the rachis. Other specimens with a dorsal attachment
arepreservedas impressions andduetotheirprobable ofthepinnaeandcontractedbasesdifferbythepinnae
coriaceous nature, only rare examples show clear being not so closely-spaced, with a more acute angle
details of the venation. One bedding plane in Coal ofattachment and by the tapering ofthe pinnae to a
Mine Quarry was covered with complete Kurtziana narrowerrounded-acute apex.
leaves, perhaps indicating an autumnal deposit of a
deciduous plant. The Nymboida specimens exhibit a 'Kurtziana cacheutensis complex', based on
range of variation in frond and pinna size, in pinna Kurtziana cacheutensis (Kurtz) Frenguelli 1942a
shape and manner ofattachment to the rachis. Some
agree closely with the types of^. cacheutensis and Figures 14A-D;15A; 16A,B; 17A
K. brandmayri, others with K. cacheutensis sensu
Herbst and Gnaedinger (2002) from collections that Selected references
also exhibited a variation in leaf form. We have 1928 'Thinnfeldia' eskensis, Walkom, P1.27, fig. 2,
separated the leaves into two 'species complexes' P1.28, fig. 1
based essentially on the perceived dorsal or lateral 1942aKurtziana cacheutensis (Kurtz) Frenguelli,
attachment of the pinnae to the rachis, a feature Pl.l
sometimes obscuredby the manner ofpreservation. 1975 Dicroidium eskense, Flint and Gould, PI. 3,
fig. 3
'Kurtziana brandmayricomplex', based on 1983 Kurtziana cacheutensis, Anderson and
Kurtziana brandmayri Frenguelli 1944 Anderson, PI. 9, fig. 5
Figures 13A,B 2002 Kurtziana cacheutensis, Herbst and
Gnaedinger, Fig.lA-H
Selected references.
1944 Kurtziana brandmayri, Frenguelli, text fig. 2, Description
mm
PI. 4, figs 1,2 Elliptic-ovate pinnate leaves, 100-200
mm
1965 ''Thinnfeldia'" eskensis. Hill et al., PI. T5, figs long, 40-100 wide, pinnae attached laterally to
mm
3,4 a striated rachis 2-3 wide, opposite to alternate,
1991 Kurtziana brandmayri, Artabe et al. Fig. 1 separated by c. one pinna width, linear-oblong,
mm
2002 Kurtziana brandmayri, Herbst and tapering slightly to acute rounded apex, to 45
mm
Gnaedinger, figs 2A-C; PI. 4, figs D-F long, 9 wide in midleaf, pinna acroscopic base
contracting to midvein, basiscopic base contracted
Description. or variously decurrent. Pirmae inserted laterally on
Kurtziana leaves with elliptic lamina, to c. rachis at c. 60° becoming more acute apically.
mm mm mm
240 long, 100 wide, rachis to 4 wide,
decreasing in width distally, striate and sometimes Material
punctate, with expanded leafbase. Pirmae opposite to AMF126832-9 Coal Mine Quarry.
sub-opposite, closely spaced to overlapping, sessile,
with contracted auriculate or caudate bases, attached Dicussion
to the dorsal surface ofthe rachis; oblong to linear- This is the most common form ofKurtziana at
ovate or tapering to rounded-acute apex, 40-50 Nymboida, sometimes forming monotypic autumnal
mm mm
long, 8-18 wide; basal pinnae broad-oval, depositsonabeddingplane(Figure 15A)orassociated
increasing in length to mid-portion ofleaf Angle of with leaves ofthe coniferRissikia (Figure 16B). This
attachment ofpinnae to rachis, from 80° near base to species complex is separated from theK. brandmayri
75° in mid-leafandbecoming more acute apically. complex by the lateral attachment of the pinnae to
the rachis. Some specimens (Figures 15A and 16A)
Material are closely comparable with the illustrated type of
K
AMF126830-1 Coal Mine Quarry. cacheutensis (Frenguelli 1942a). However, there
is a wide range of variation in leaf size, pinna size
Discussion and shape, spacing and inclination ofpinnae to rachis
The Nymboida specimen on Figure 13A agrees and the degree of contraction or decurrence of the
K
closely with the type of brandmayri (Frenguelli basiscopic base ofthe pirma. Figure 17A shows two
1944, PI. 4, figs 1,2) by the closely-spaced to leaves with extreme decurrent pinnae in the apical
Proc. Linn. Soc. N.S.W., 126, 2005 45
TRIASSIC FLORA FROM NYMBOIDA SOMF GYMNOSPERM GENERA
-
halfofthe leafand contacted pinnae in the basal half thus Rochipteris Herbst et al. (2001) has priority over
Some leaves (e.g. Figure 14C) appear to have pinnae Kannaskoppifolia. Kannaskoppifolia may be used as
with asymmetrical laminae but this is probably an a genus for leaves attached to a stem.
artifact ofpreservation caused by the inrolling ofone Anderson and Anderson (2003) regarded
edge ofthe pinna. Kannaskoppiafolia (= Rochipteri.s) as "ubiquitous,
diverse, long-lived, relatively frequent but generally
lacking in abundance". Barone-Nugent et al. (2003)
Order Petriellales Taylor et al. 1994 noted that their Rochipteris species appeared to be
Family Kannaskoppiaceae Anderson and Anderson distinct between separate basins and showed a strong
2003 degree of intra-Gondwanic provincialism in marked
Genus Rochipteris Herbst, Troncoso and contrast to Dicroidium species, which are widely
Gnaedinger 2001 distributed throughout Gondwana (Retallack 1977;
Type species Rochipteris lacerata (Arber) Herbst et AndersonandAnderson, 1983; HolmesandAnderson,
al. 2001 in press). Forty years ofcollecting at Nymboida has
yielded the six new species described below, but, with
Flabellate leaves with anastomosing venation theexceptionoftwospecies,theothersarerepresented
have long been known from the Triassic floras of by a single specimen only. The species have been
Gondwana. Early records were from Queensland by distinguished on leaf morphology, venation pattern
Carruthers (1872, as Cyclopteriscuneata) and Shirley and vein density. Two of the Nymboida species are
(1898, as Sagenopteris cuneata), from Tasmania important in demonstrating, for some species at least,
by Johnston (1888 as Cyclopteris aiistralis), from that Rochipteris foliage is inserted on the stem either
Victoria by Chapman (1927 as Psygmophyllum as a close spiral or a terminal whorl.
fergusoni), from New Zealand by Arber (1917 as
Chiropteris lacerata), from Chile by Solms-Laubach
(1899 as Chiropteris copiapensis) and South Rochipteris obtriangulata Holmes andAnderson sp.
Africa by Seward (1903 as Chiropteris cuneata). nov.
Other records may be found in Etheridge (1895), Figures 18A-C; 19A-D
Chapman and Cookson (1926), Frenguelli (1942b,
1944) and Menendez (1951). The taxonomy of the Diagnosis
group was confused. Retallack (1980) discussed the Obtriangular lamina, lateral and distal margins
problems and subsequently recognised six species straight, entire; angle of divergence 20°-30°;
under an emended diagnosis for Ginkgophytopsis. veins sub-parallel, dichotomising c. five times and
Herbst et al. (2001) made a detailed analysis of anastomosing twice in distal half of lamina; leaves
the significant differences between the essentially attached in close spirals or whorls of eight to ten;
Northern Hemisphere genus Ginkgophytopsis and venation density 20-25 per 10 mm.
the Gondwanan leaves. For the Gondwanan leaves
they erected the new genus Rochipteris in which Description
was included five species based mainly on material Basedonaslabbearingimpressionsofonealmost
from Argentina and Chile. Barone-Nugent et al. complete whorl andtwo incomplete whorls plus other
(2003) redescribed leaves from the Leigh Creek Coal isolated single leaves. The leaves are obtriangular, c.
mm mm
Measures of South Australia and the Ipswich Coal 40 long and 12 wide at the truncate apex.
Measures of Queensland as Rochipteris etheridgei Lamina diverging from the acute sessile base at c.
and R. ginkgoides respectively. 20°-30°, Lateral margins straight, leafapex truncate,
In a recent significant publication on Gondwana straight and entire. Veins sub-parallel, dichotomising
Triassic gymnosperms, Anderson and Anderson close to base and then four or five more times to leaf
(2003) described some remarkable material apex; from c. mid-lamina the veins converge and
from the Molteno Formation of South Africa. conjoin with adjacent veins, usually twice, to form
Included were specimens of flabellate leaves with in'egular linear elliptical areoles; venation density at
anastomosingvenationandwith eitherfemaleormale 2/3 leaflength 20-25 per 1 mm. Foliage arranged in
fructifications attached to a stem. The female strobili a close spiral or a terminal whorl of 8-10 leaves but
were described as Kannaskoppia, the male strobili leafbases and stem not visible.
as Kannaskoppianthus and the attached leaves as
Kannaskoppifolia. Under ICBN (2001), Article 1.2, Holotype
AMF AMF
detached leaves are regarded as morpho-taxa and 126840 and counterpart 126842;
46 Free. Linn. Soc. N.S.W., 126, 2005
W.B.K. HOLMES AND H.M. ANDERSON
paratypeAMF126841, Australian Museum, Sydney. Name Derivation
tubata (Lat.), trumpet; referring to the
Type Locality expanding outline ofthe leaflamina.
Coal Mine Quarry, Nymboida. Basin Creek
Formation, Nymboida Coal Measures, Middle Discussion
Triassic. Rochipteris tubata is arranged in a whorl of
leaves similar to R. obtriangularis but unfortunately
Name Derivation in neither species is their attachment to the stem
obtriangiilata (Lat.), obtriangular; referring to visible. The venation in both species is similar but
the reversed triangular leafform. R. tubata is separated on the basis ofthe expanding
lamina and rounded leafapex. The whorls of leaves
Discussion in R. obtriangulata and R. tubata suggest a strong
The straight lateral margins of the expanding relationship to the stems bearingKannaskoppia fruits
laminae and the truncate entire apex differentiates R. (Anderson and Anderson 2003), which have small
obtriangiilata from all other described Rochipteris groups orwhorls ofleaves attached at intervals along
species. They are close to the leaves from the the slender stem.
Molteno Formation locality Umklll illustrated as
Kannaskoppifolia sp. C by Anderson and Anderson
(2003). Rochipteris incisa Holmes andAnderson sp. nov.
Figures21A-C;22A
Rochipteris tubata Holmes andAnderson sp. nov.
Figures 20A-C Diagnosis
Medium-sizedcuneate leafwith archedapex; one
Diagnosis to six deep incisions to below mid-lamina forming
Vase-shaped lamina, lateral margins concave, sub-parallel lobes; venation parallel, occasionally
distal margin convex-rounded, entire; angle of bifurcatingorconjoiningto formextremelyelongated
divergence at base 15° increasing to 60°-90° linear areoles. Venation density c. 18 per 10 mm.
apically; veins sub-parallel, dichotomising from near
base, anastomosing in distal 2/3 of lamina. Foliage Description
in a close spiral or whorl ofc. 7 leaves; vein density Based on three leaves from the Reserve Quarry
20-25 per 10 mm. and eight from Coal Mine Quarry. Leaf cuneate, to
mm mm
115 long and c. 70 wide; lateral margins
Description straight or slightly concave, diverging from base at
Based on one almost complete whorl of seven c. 45°-80°; apex semicircular, deeply incised to form
mm mm
leaves. Lamina vase-shaped; leaves to 30 long, a number of linear segments from 8-10 wide,
mm
19 wide, rising from an attenuated base at c. incisions reaching to 1/3 distance from the lamina
15° and expanding distally in a curve to 60°-90°. base, distal ends of segments entire or with a minor
Lateral margins concave, apical margin convex- incision. Veins fine and parallel, dichotomising and
rounded, slightly undulate. Venation sub-parallel, occasionally conjoining to form extremely elongated
dichotomising c. five times from near base and areoles; density ofveins c. 18 per 10 mm.
anastomosing in distal 1/3 of lamina. Seven leaves
apparently attachedin a close spiral orterminal whorl Holotype
but leafbases and stem not visible. Density ofveins AMF126844 Australian Museum, Sydney.
c. 20-25 per 10 mm.
Type Locality
Holotype Reserve Quarry, Nymboida. Basin Creek
AMF126843 Australian Museum, Sydney. Formation, Nymboida Coal Measures.
Type Locality Other Material
Coal Mine Quarry, Nymboida. Basin Creek AMF126827, 126862, Reserve Quarry;
Formation, Nymboida Coal Measures, Middle AMF126863-66, Coal Mine Quarry.
Triassic.
Name Derivation
incisa, (Lat.), incised, referring to the regularly
Proc. Linn. Soc. N.S.W., 126, 2005 47
TRIASSIC FLORA FROM NYMBOIDA SOME GYMNOSPF^RM GFNF:RA
-
incised distal margin. AMP126845 Australian Museum, Sydney.
Discussion Type Locality
Rochipferis incisa shows some similarities in Coal Mine Quarry, Nymboida. Basin Creek
shape and outline to the attached Moltcno leaves Formation, Nymboida Coal Measures, Middle
Kaiinuskoppifoliavinciilaris(AndersonandAnderson Triassic.
2003). It differs from all other Nymboida Rochipleris
spp in lamina shape and venation details. Rochipferis Name Derivation
etheridgei (Arber) Barone-Nugent et al. is similar in sinuosa (Lat.) sinuous; referring to the venation.
outline to R. incisa but differs by its larger size, the
less-deeply incisedsegments,thesinuatevenation and Discussion
by the presence ofa broadly-flared leafbase (Barone- The gross morphology, sub-sinuous venation and
Nugent et al. 2003, fig. 33, PI. 1, figs 2-5). paucity of anastomoses distinguishes this leaf from
Onthesameslabandadjacenttotheholotypeof^. other Nymboida Rochipleris species. Rochipleris
incisaisasmallerspathulateleafwithvenationsimilar sinuosa closely resembles the specimens from the
to that ofR. incisa (Figure 2IB). This spathulate form Llantenes Formation of Argentina attributed to
is similar to leaves from Argentina that have been Chiropleris copiapensis Steinmann and Solms by
placed in Rochipleris cimeata (Carruthers) Herbst Menendez (1951, PI.3, figs 1^). However, those
et al. 2001. That species was based on Cyclopteris specimens have been synonymised with Rochipleris
cimeata Carruthers (1872), a poorly preserved leaf lacerala (Arber) as a new combination by Herbst
fragment with both apex and base missing. We et al. (2001). Rochipleris lacerala was described
believe that our spathulate leaf may belong to the originally from New Zealand by Arber (1917) as
same population asR. incisa andperhaps represents a Chiropleris lacerala and after detailed discussion by
juvenile or immature stage ofdevelopment. Retallack (1980, 1983) was transferred to the genus
Ginkgophylopsis. Rochipleris lacerala sensu Herbst
etal. (2001) is largerthanR. sinuosa, is deeply incised
Rochipleris sinuosa Holmes andAnderson sp. nov. into several parallel-sided segments and has straight
Figures 23A-C parallel venation that dichotomises and anastomoses
to form long areoles. Rochipleris copiapensis
Diagnosis (Solms-Laubach) sensu Herbst et al. (2001) is a large
A small flabellate leaf, diverging at c. 45° from leaf divided into two equal segments with straight,
short expanded leaf base; divided into two major bifurcating and anastomosing venation. In outline
segments by a deep incision; one segment again and venation pattern, R. sinuosa differs from the
divided by a shallower incision; veins radiating ten illustrated but undescribed Kannaskoppifolia
from base, sinuous, approaching and diverging from = Rochipleris leaves from the Molteno Formation
each other, occasionally dichotomising but rarely (Anderson andAnderson 2003).
anastomosing. Vein density c. 18 per 10 mm.
Description Rochipleris nymboidensis Holmes andAnderson sp.
Basedonasingleleafwithapicalmarginmissing. nov.
Leafaspreserved,42mmlong,25mmwide;flabellate, Figures 24A-D
expanding at c. 45° from a short flared leaf base 4
mm
wide. Lamina divided into two major segments Diagnosis
by a medial incision commencing at c. 12 mm from A small cuneate leaf, lateral margins concave,
the leafbase; the left segment is again divided by a apical margin convex, entire; venation dense, straight
mm
narrow incisioncommencingat 18 from thebase. andparallel,dichotomising,veryrarelyanastomosing;
The lamina lateral margins are slightly concave, the vein density c. 30-35 per 10 mm.
apical margin is missing. Veins run in a sub-sinuous
manner parallel to the lateral margins, dichotomising Description
A
occasionally, approaching and diverging from each Based on a single specimen. cuneate leaf 63
mm mm
other but rarely forming a true anastomosis. The long, with leafbase missing, 50 wide at the
apparent areoles are elongate-elliptic. Vein density in entire to slightly undulate apex. Angle ofdivergence
the upper portion ofthe lamina c. 18 per 10 mm. from base c. 25°, increasing to 90° at lamina apex.
Holotype Venation very fine, parallel, straight, dichotomising
48 Proc. Linn. Soc. N.S.W., 126, 2005