Grammatocaulis donponii gen. et sp. nov., a permineralized fern from the Jurassic of Queensland, Australia

Review of Palaeobotany and Palynology, 66 (1990): 147-158

147

Elsevier Science Publishers B.V., Amsterdam

Grammatocaulis donponii gen. et sp. nov., a permineralized fern from the Jurassic of Queensland, Australia William D. Tidwell I and Andrew

C. R o z e f e l d s 2

1Department of Botany, 401 WIDB, Brigham Young University, Provo, UT 84602 (U.S.A.) 2Queensland Museum, P.O. Box 300, South Brisbane, Qd., 4101 (Australia) (Received November 28, 1989; revised and accepted May 23, 1990)

ABSTRACT Tidwell, W.D. and Rozefelds, A.C., 1990. Grammatocaulisdonponiigen. et sp. nov., a permineralized fern from the Jurassic of Queensland, Australia. Rev. Palaeobot. Palynol., 66: 147-158. A permineralized fern stem from Middle to Late Jurassic sediments (Injune Creek Group) in the Wandoan area of southeastern Queensland is assigned to the new genus and species Grammatocaulisdonponii. The stem possesses an ectophloic siphonostele with the xylem divided into an outer zone of long tracheids and an inner zone of short, wide, parenchyma-like tracheids surrounding a pith. Leaf traces are round in cross section in the parencbymatous inner cortex, becoming linear to oblong in the outer cortex and outside of the stem. Petiole bases have sclerotic rings around vascular strands, but lack stipular wings. Some of its outermost petiolar vascular strands assume a C-shape which may be due to distortion. Parenchymatous tissue occurs in the middle of the protoxylem at the ends of the elliptical petiolar vascular strands. At higher levels, the parenchymatous tissue of the strands is found in the metaxylem as well. Nonglandular multicellular scales occur on the petioles. Although the family of this genus is unknown, it is similar anatomically to Grammatopteris of the Anachoropteridaceae from the Late Paleozoic of Europe. If these two genera are related, this specimen suggests more structural diversity, a wider geographic distribution and a longer geologic range for the Anachoropteridaceae than previously known.

Introduction N u m e r o u s fossil ferns are k n o w n f r o m the Jurassic s t r a t a o f Q u e e n s l a n d ( K i d s t o n a n d G w y n n e - V a u g h a n , 1914; W a l k o m , 1917; D u n s t a n , 1920; Hill et al., 1966; G o u l d , 1973, 1974, 1981; Rigby, 1978). All o f the petrified ferns in these r e p o r t s have been assigned to the O s m u n d a c e a e . H o w e v e r , the new genus a n d species d e s c r i b e d here does n o t b e l o n g to this f a m i l y a n d is b a s e d u p o n a p e r m i n e r a l i z e d specimen r e s e m b l i n g Grammatopteris R e n a u l t . T h e specimen w a s ' f o u n d in the W a n d o a n a r e a in Q u a t e r n a r y s t r e a m d e p o s i t s a l o n g H o r s e C r e e k on W a r r a w o o n a S t a t i o n t h a t are r e w o r k e d f r o m the extensive J u r a s s i c - L o w e r C r e t a c e o u s s e d i m e n t s in the region. T h e s e d i m e n t s f r o m which this w a t e r - w o r n fern stem was d e r i v e d are m a p p e d as the M i d d l e to U p p e r Jurassic I n j u n e

0034-6667/90/$03.50

C r e e k G r o u p ( F i g . l ; F o r b e s , 1968; G o u l d , 1968; E x o n , 1971, 1976). A s s o c i a t e d with the fern stem is s o m e p e n t o x y l a l e a n w o o d , u n d e s c r i b e d c y c a d e o i d e a n stems, as well as a b u n d a n t p e r m i n e r a l i z e d c o n i f e r w o o d a n d o s m u n d a c e o u s axes. F r o m the I n j u n e C r e e k G r o u p at the n e a r b y " W a r r e n d o o S t a t i o n " west o f W a n d o a n , G o u l d (1974) r e c o r d e d i m p r e s s i o n s o f Ptilophyllum sp., Taeniopteris spathulata a n d a p o s s i b l e Nilssonia sp. cf. N. elegans as well as Osmundacaulis sahnii ( M i t r e , 1955) o r O. rajmahelense ( G u p t a ) S h a r m a (1973). T h e s e l a t t e r species have been s u b s e q u e n t l y p l a c e d in Millerocaulis (Tidwell, 1986). Osmundacaulis hoskingii G o u l d (1973) was r e p o r t e d f r o m M o u n t O r g a n S t a t i o n a n d Ptilophyllum i m p r e s s i o n s a r e also k n o w n from nearby Bimbadean Station. The specimen of Grarnmatocaulis donponii is d e p o s i t e d in the

© 1990 - - Elsevier Science Publishers B.V.

148

WD

Quaternary

[IDWELL AND A.C ROZEFELI)S

Alluvium

i

Orallo

Middle to Upper Jurassic

Formatlon

Gubberamunda

Sandstone

injune Creek Group Birkhead

Formation

km

Fig. l. Geologicmap of the collecting site for Grammatocaulis donponii gen. et sp. nov. in southeastern Queensland, Australia. (Map adapted from Exon, 1971 and Forbes, 1968).

Queensland Queensl.

Museum

(QMF)

in

Brisbane,

Systematic description Family Unknown

Grammatocaulis gen. nov. Diagnosis: Fossil fern rhizomes, arborescent or erect axes surrounded by non-stipulate leaf bases

and adventitious roots; ectophloic siphonostele; xylem cylinder composed of an outer, or peripheral zone, of long tracheids, 10 or more tracheids thick in radial diameter and an inner zone of short, wide, parenchyma-like tracheids; leaf gaps lacking; cortex differentiated into inner and outer tissues without sclerotic nests in cross-section; leaf traces initially round becoming linear in shape and tangential to stem; roots diverging from leaf traces in inner stem cortex and from petiolar vascular strands outside of stem.

149

G R A M M A TOCA ULIS DONPONH GEN. ET SP. NOV.

Generitype." G. donponii Grammatocaulis donponii sp. nov. (Plates I-III;

Age: Middle to Late Jurassic Etymology: Specific epithet honors Mr. Norman

Fig.2)

Donpon of Meandarra, Queensland who donated the specimen.

Diagnosis: Tree fern, stem surrounded by leaf bases and roots; stipular wings lacking; siphonostelic; xylem cylinder of inner and outer zones, inner zones consists of short, wide tracheids (parenchyma-like), outer zone of radially aligned, long, pointed tracheids and an occasional radial row of parenchyma cells, protoxylem mesarch; xylem sheath, phloem, pericycle and endodermis external to xylem; cortex, inner parenchymatous, no sclerotic nests, outer sclerenchymatous; leaf traces in cross section, round basally, becoming linear, tangential to stem, straight to C-shaped in the outermost petiolar vascular strands, latter may be oriented adaxially or abaxially; protoxylem clusters divided by parenchymatous tissue at the ends of petiolar vascular strands; multicellular scales on petioles; root traces diarch, arising from leaf traces and occasionally from petiolar vascular strands. Holotype: Q M F 16183 Locality: Warrawoona Station, Kabunga Road, Wandoan, southeastern Queensland, Australia.

Description The specimen, I 15 × 162 mm across and 60 mm thick, is composed of a stem surrounded by petioles and roots (Plate I, 1). The stem is 30 x 40 mm across and flattened. It consists of an ectophloic siphonostele enclosed by inner and outer cortices. The center portion (4 x 6.5 mm in diameter) of the stele comprising the pith is not preserved. It is surrounded by 6-7 layers of crushed cells (Plate III, 1). The xylem cylinder, 11 x 16 mm across, is made up of two distinctly different regions: An inner zone of short, parenchyma-like, tracheal elements and an outer or peripheral zone of long tracheids (Plate I, 3). The inner zone is 1-3 mm wide and contains short, wide tracheids of nearly equal dimensions (Plate I, 4, 5). The tracheids are up to 250 x 400 Ixm in diameter and length and elongated to hexagonal

Fig.2. Gramrnatocaulisdonponiigen. et sp. nov. (A) A stereodiagram of the xylem cylinder illustrating departing traces and the inclusion of parenchyma in the petiolar vascular strand (not to scale). (B) Outermost petiolar vascular strands showing the C-shaped configuration. Structures around them are roots ( × 5). (C) Xylemof an outer petiolevascular strand. Note the gaps at each end which were occupied by parenchyma ( x 40). (B-C transverse sections).

[ 50

in shape. Their terminal walls are transverse or oblique. They often have round pits between the bars of the reticulate thickenings. The smaller tracheids occur towards the outside of the tissue and most of the larger and elongated ones are toward the inner edge of this tissue. The outer xylem zone is 1.5 mm wide, fairly uniform in width, and has elongated tracheids (Plate I, 6). Transversely, those tracheids near the inner periphery of this zone are smaller and diameters of the tracheids become larger towards the outside. They are up to 100 gm in diam. and 850 gm in length and have scalariform, occasionally reticulate, thickenings (Plate I, 4; Plate IV, 3, 5) with many being irregularly arranged. Clusters of mesarch protoxylem and rows of parenchyma

W D 11DWELL AND A.C. ROZEFELDS

cells occur among the long tracheids (Plate II, 6). These rows originate along the boundary with the short tracheids and are radially oriented. In general, they do not completely traverse this tissue and do not appear to have any relationship to the leaf traces. Although the phloem surrounds the xylem as a continuous ring, it is separated from the latter by a crushed xylem sheath that is 70 tam wide (Plate I1, 6). The phloem is up to 160 p,m wide and is one to three cells thick, forming a single-cell layer around the diverging traces. No protophloem could be discerned. The pericycle is not preserved, but the space it occupied is 100 lam wide and a poorly preserved, single-layered endodermis separates the pericycle from the inner cortex.

PLATE 1

Grammatocaulis donponii gen. et nov. I. 2. 3.

Transverse section of the actual specimen (QMF16183). x 75. Cross-section through two roots (QMFI6183 no.l), x 10. Transverse section of a part of the stem showing the short tracheids (a), long tracheids (b), xylem sheath (c), phloem (d), inner cortex (e), outer cortex not preserved in this view, and a departing trace (f) with two roots (g, h) arising from it (QMFI6183). 4, 5. Short tracheids illustrating the reticulate thickenings. 4. longitudinal section. 5. cross-section. Note how the short tracheids are similar in size to each other and they parallel the long tracheids which occur to the left in 4. Also note how much they resemble parenchyma cells in 5 (QMFI6183 no.3), x 100. PLATE II (see p.1521

Grammalocaulis donponii gen. et sp. nov. 1. Elongated petiolar vascular trace in transverse section. Note that part of the cortex was not preserved and also the outer sclerotic ring (QMFI6183 no.I), x 15. 2. A number of ribbon-shaped multicellular scales from the epidermis of a petiole (QMF16183 no.l). × 15. 3. Longitudinal section of the stele illustrating separating leaf traces. Note separating root trace (arrow) (QMF16183 no.3). × 15. 4. Enlargement of a petiolar vascular strand in transverse section showing the metaxylem tracheids (a), the protoxylem (b), and the parenchyma cells (c) that occur between the protoxylem clusters (QMF16183 no.I). × 80. 5. Transverse section of the inner cortex with round leaf trace and a root trace (arrow) (QMFI6183). × 12. 6. Transverse section of a portion of the stem illustrating the phloem (a), xylem sheath (b) and long tracheids (c), Note the strands of parenchyma (d) and trace (e) (QMF16183 no.l), x 30.

PLATE llI (see p.153)

Grammatocaulis donponii gen. et sp. nov. I. 2. 3.

Transverse section of crushed cells at the interface between the short tracheids and pith (QMF16183 no. l). × 40. Enlargement of leaf trace in transverse section in the inner cortex ( × 35). Longitudinal view of petiolar vascular strand after leaving the stem. Cortex around strand is not preserved (QMFI6183 no. ll. ×15. 4. Overview in transverse section of outermost portion of the specimen illustrating the C-shaped petiolar vascular strand facing adaxially (a) and abaxially (b) ( x 10). 5. Cross section of petiolar vascular strand. Note the parenchyma (arrow) between upper and lower protoxylem clusters (QMFI6183 no.l), x20. 6, 7. Close-up of C-shaped petiolar vascular strands, 6. oriented adaxially and 7. abaxially. Note distortion (QMFI6183 no.6), x 20.

GRAMMA TOCA ULIS DONPONII G EN. ET SP. NOV,

151

PLATE I

The inner cortex is 1.5 mm wide and is not well preserved. It is composed of thin-walled ovoid or rounded parenchyma cells of near equal dimensions without any sclerotic sacs or nests being

present. The tissue had split before or during preservation and much of it has disintegrated. The outer cortex (10-15 mm wide) is irregularly shaped and consists of relatively thick-walled fibers

152

P L A T E II

(for explanation see p.150)

\~, 1) I I D W E I I

AND A.(

R O Z E F t - [ 1)5;

GRAMMA TOCA ULIS DONPONII G E N . E T SP, N O V .

153

PLATE III

i!i!~~

~ii

~

,,

÷~~i~~~ ~!

~ilil

(for explanation see p. 150)

(15-30 pm wide x 150-350 ~tm long) that surround the traces as they depart the stem. Leaf traces arise directly from the xylem without

definite leaf gaps. A slight bulge on the edge of the xylem cylinder signifies the base of the trace. This tissue extends outward and upward to form the

154

trace. Transversely, the outer tracheids associated with the trace become somewhat radially elongated towards the trace. The leaf traces make an adaxial angle of 30 ° with the long axis of the stem and after leaving the vascular tissue, they give rise to one or two lateral roots (Plate I, 3; Plate II, 3; Plate III, 3; Fig.2A). The leaf trace has a circular transverse shape at its base (Plate II, 5; Plate IlI, 2) with a cluster of mesarch protoxylem elements at its center. Outwardly, it becomes tangentially flattened into a ribbon-like vascular strand with exarch protoxylem at each end of the trace. A group of parenchyma cells occurs in the center of the protoxylem clusters, which in turn sometimes surround this parenchymatous tissue. Generally, however, the clusters are separated by the parenchyma and thus, remain open (Plate IV, 1, 2; Fig.2C). The inner cortex of the petiole consists of small to large, thin-walled parenchyma cells and is continuous with the inner cortex of the stem. As the traces pass through the outer cortex, they become surrounded by a wide, homogeneous, sclerotic ring. Although the petioles are roundish (80 m m x 80 ram) in transverse section, they are often distorted (30 mm x 60 mm) and they lack stipular wings (Plate II, 1). The petiolar vascular strands outside the stem are similar to the leaf traces. The protoxylem groups at the ends of the elliptical strands occasionally surround, but most often are separated by parenchyma (Plate I1, 4, Plate III, 5; Plate IV, 1, 2). However, at high levels of the strands, parenchyma cells also occur in the metaxylem, as well as protoxylem (Fig.2C). The xylem of the strands is surrounded by a parenchymatous sheath of thin-walled cells. Although a few

w I1 III)WIL[ AND A ( ROZEI=[LI)S

elements are present suggesting that the phloem may have surrounded the xylem of the strand, this tissue is rarely preserved. A row of cells with dark contents representing the endodermis occurs outside of these tissues. The inner and outer ground tissue around these strands is similar to that in the stem. The uniseriate epidermis is generally preserved as a discontinuous dark line. Pinna traces were not observed. The vascular strands of the outermost petioles become C-shaped to some degree with their orientation varying from the C facing the stem to its facing outwardly due to twisting of the petioles (Plate III, 4, 6, 7: Fig.2B). Protoxylem occurs at the tips of the arms of these strands. Non-glandular epidermal scales (80 185 gm wide) attached to the epidermis of the petioles (Plate If, 2; Plate IV, 4) are unbranched, multicellular and ribbon-shaped. They fill much of the space between petioles, particularly among the older ones. The diarch roots usually arise from the leaf traces and occasionally from petiolar vascular strands. They are variously shaped from elongated to four-sided due to distortion from growing in the limited space between the petioles (Plate I, 2). The xylem of the roots is two to three tracheids wide and their phloem, pericycle and endodermis are not preserved. The inner cortex of these organs is thin-walled parenchymatous tissue and their outer cortex is sclerenchymatous.

Comparison and discussion

Grammatocaulis donponii is similar to the Paleozoic fern Grammatopteris in having an inner xylary zone of shorter and an outer zone of longer

PLATE IV

Grammatocaulis &mponii gen. et. sp. nov. 1. Transverse section of one end of a petiolar vascular strand illustrating the metaxylem (a), protoxylem clusters (b), parenchyma between the protoxylem (c) and the sclerotic ring (aft (QMF16183 no.2), x 30. 2. Enlargement of tigure 1. x 80. 3. Longitudinal section through the long tracheids, Note the scalariform thickenings of the tracheid walls (QMFI6183 no,3), x 50. 4. Transverse section of a number of multicellular, epidermal scales attached lo the epidermis of the petiole (darker area near the top) IQMFI6183 no.I), ×30. 5. Longitudinal section of the long tracheids illustrating scalariform and reticulate thickening on thc trachcid walls (QMFI6183 no.3). × 80.

GRA M M A T O C A U L I S D O N P O N I I GEN. ET SP. NOV.

[ 55

PLATE IV

tracheids and in having linear, oblong petiolar vascular strands tangential to the stem. Three species, Grammatopteris rigollotii Renault

(1896) from the Lower Permian of France, G. balo daufii (Beck) Hirmer (1927) from the Upper Carboniferous (Stephanian) of Hilbersdorf of the

15('~

G.D.R. and G. bertrandii Corsin (1937) from the Lower Carboniferous of France, are known. The stem of G. bertrandii is presently unknown, whereas, G. rigollotii and G. baldaufii are composed of small, upright, radially symmetrical axes considered to be small-sized tree ferns (Sahni, 1932), but not necessarily arborescent (Miller, 1971). The protostele of the stems of these species consists of a central zone of tracheids shorter and wider than those of an outer zone of long tracheids. The leaf traces are elliptical to barshaped, tangential to the stele, and without curvature, either ab- or adaxial. The petiole bases are helically arranged around the stem and cylindrical in shape. Sahni (1932) emphasized the sclerotic nests in the cortex, the slightly invaginated protostele and the straight band-shaped foliar bundles of this genus. The basic differences between Grammatocaulis donponii and the species of Grammatopteris are that G. donponii is siphonostelic, has round rather than elliptic leaf traces in its inner cortex, has no sclerotic nests, has a two part cortex rather than three as in G. baldat~ffii and G. rigollotii (Corsin, 1937), lacks rudimentary gaps that are present in G. baldaufii, has protoxylem clusters that are separated by parenchyma cells at each end of the petiolar vascular strand rather than peripherally adaxial as in G. baldaufii and G. rigollotii and lacks the sclerotic fundamental tissue that is found in the petioles of G. bertrandii. The pith of Grammatoeaulis donponii is not preserved and although some cell layers around the edge of this space have collapsed, there is no evidence that this space was filled by short tracheids. The fact that the tracheids around this area are very well preserved suggests that the pith was constructed of thinner walled parenchyma cells that disintegrated prior to fossilization. This interpretation parallels that of Eggert (1964) who postulated that the inner zone of small tracheids in Grammatopteris represented a stage in the evolution of a pith. This may have occurred in protostelic ancestors of G. donponii, where the cells in their centers lost the ability to form secondary walls, and therefore, remained parenchyma cells. According to Kidston and Gwynne-Vaughan (1909, 1910), the tracheids were either transformed

\~, I) IIDWE[.I. A N D A . C ROZEFI-~LI)S

simultaneously into parenchyma cells forming a homogeneous pith, or only some of the tracheids lost their characteristics, resulting in a mixed pith. Because there is no evidence for a mixed pith in G. donponii, perhaps its pith evolved simutaneously, although it is difficult to imagine a uniform change as a tissue rather than a gradual loss of secondary walls on a cell by cell basis. The parenchyma tissue separating the protoxylem groups at the ends of the petiolar vascuolar strands in G. donponii is continuous from its presence in the leaf traces of the outer cortex to the higher levels of the strands. It is similar to that in Diplolabis roemeri (Solms-Laubach) Gordon (1911) but, unlike the latter species, there is no definite closure of the tissue or pinna being given off. This tissue is not present in Grammatopteris and appears to be a vestigal feature in Grammatocaulis. A "root felt" (Sahni, 1932) occurs around the periphery of the specimen. Most of the roots are vertically oriented and thus, are cut transversely which indicates an erect or arborescent growth habit (Miller, 1971). Roots arising from leaf traces are not unusual in ferns, however, roots originating from the abaxial surface of each trace, and only the trace, as in Grammatocaulis donponii, can be an important characteristic (Phillips and Andrews, 1966). This feature occurs in Tubicaulis q/'ricanus Holden and Croft (1962) and A urealcaulis crossii Tidwell and Parker (1987). Roots arise from petioles in Catenopteris simplex Phillips and Andrews (1966), occasionally in G. baldaufii and in Botryopteris, their source is the foliar members associated with incipient shoot formation (Phillips and Andrews, t966). Roots are also derived from the stem in C. simplex, but not in G. donponii. The C-shaped petiolar strands of the outermost petiole bases in G. donponii may be due to distortion, The older petioles appear not to be rigid; they meander between one another with some twisting until these C-shaped strands face outwardly (abaxial), as well as inwardly (adaxial) (Plate Ill, 4, 6, 7; Fig.2B). Beck (1920) mentioned strands with slight adaxial curves were of common occurrence in G. baldau/ii and attributed their shape to being deformed. However, Phillips and

157

G R A M M A TOCA ULIS DONPONII GEN. ET SP. NOV.

Andrews (1966) noted that, although the protoxylem elements generally formed a slight ridge at the ends o f the xylem strand in G. baldaufii, the protoxylem groups occasionally formed prominent adaxial projections which were considered atypical by Sahni (1932) but give a small adaxial curvature to the xylem strand. Grammatocaulis donponii is also anatomically similar to Asterochlaenopsis kirgisica (Stenzel) Sahni (1930) from the Permian o f Kirgis Steppes o f western Siberia (Andrews and Boureau, 1970). They differ in that Asterochlaenopsis has a stellate mixed pith, permanently closed peripheral loops in its petiolar vascular strands, aphlebiae and lacks multicellular scales or epidermal hairs on its petioles. A l t h o u g h Grammatopteris is assigned to the A n a c h o r o p t e r i d a c e a e (Eggert, 1964), it has been c o m p a r e d to early members o f the O s m u n d a c e a e (Sahni, 1932). The double protoxylem o f G. rigollotii, the absence o f adaxially curved petiole strands and stipular wings, and the sclerotic cortex o f G. rigollotii, make any relationship between them unlikely (Miller, 1971). Grammatocaulis differs from members o f the O s m u n d a c e a e by the lack o f stipular wings and curvature in most o f the strands in Grammatocaulis and by having protoxylem groups terminating the ends o f the strands in this latter genus that does not occur a m o n g o s m u n d a c e o u s forms. Like Grammatopteris (Eggert, 1964), without the frond and fertile c o m p o nents, the relationship o f Grammatocaulis cannot be definitely answered at present. The occurrence o f Grammatocaulis donponii in the Southern Hemisphere is significant. It appears much later in the fossil record than the Grammatopteris species from the Carboniferous and Permian o f Europe which suggests that, if they are related, the modification o f G. donponii from a possible Grammatopteris progenitor p r o b a b l y occurred during the interval o f the long migration o f its ancestors from the N o r t h e r n to the Southern Hemisphere. H o w this occurred and the route these ancestors took remains uncertain.

Acknowledgements We wish to thank Dr. T o m Phillips o f the University o f Illinois and Dr. Charles N. Miller, Jr.

o f the University o f M o n t a n a for reviewing the manuscript and Mr. D o u g M o o r e o f Glendora, California and Mr. Paul Cary o f Microtec Engineering Lab, Clifton, C o l o r a d o for their assistance with the thin-sectioning.

References Andrews, H.N. and Boureau, E., 1970. Classe de Coenopteridopsida. In: E. Boureau (Editor), Filicophyta. Trait~ de Paleobotaniqu6 IV(l). Masson, Paris, 519 pp. Beck, R., 1920. l]ber Protothamnopterisbaldaufiinov. sp., einen neuen verkieselten Farnaus dem Chemitzer Rotliegenden. Abh. Math.-Phys. K1. S~ichs. Akad. Wiss., 36(5): 513-522. Corsin, P., 1937. Contribution a l'l~tudes des Fougbres Anciennes du Groupe des Inversicatenales Pt. 1. l~tude du genre Grammatopteris., G. Sautai, Lille, pp.l-80. Dunstam G., 1920. Northwestern Queensland; geological notes on the Cloncurry-Camooweal-Burketown-Boulia area. Publ. Geol. Surv. Queensl., 265, 48 pp. Eggert, D.A., 1964. The question of the phylogenetic position of the Coenopteridales. Bull. Torrey Bot. Club, 21(5): 38-57. Exon, N.F., 1971. Roma, Queensland 1:250,000 geological series explanatory notes. Rep. Bur. Miner. Resour. Geol. Geophys. Aust., 40 pp. Exon, N.F., 1976. Geology of the Surat Basin in Queensland. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., No. 166, 160 pp. Forbes, V.R., 1968. Taroom Queensland 1:250,000 geological series explanatory notes. Rep. Bur. Min. Resour. Geol. Geophys. Aust., 20 pp. Gordon, W.T., 1911. On the structure and affinities of Diplolabis roemeri (Solms). Trans. R. Soc. Edinburgh, 47: 711-736. Gould, R.E., 1968. The Walloon Coal Measures: a compilation. Queensl. Gov. Min. J., 69:509 515. Gould, R.E., 1973. A new species of Osmundacaulis from the Jurassic of Queensland. Proc. Linn. Soc. N.S.W., 98(2): 86-94. Gould, R.E., 1974. The fossil flora of the Walloon Coal Measures: A survey. Proc. R. Soc. Queensl., 85(3): 33-41. Gould, R.E., 1981. The coal forming flora of the Walloon Coal Measures. Coal Geol. (Austr.), I(3): 83 105. Hill, D., Playford, G. and Woods, J.T., 1966. Jurassic fossils of Queensland, Brisbane. Queensl. Palaeontogr. Soc., Brisbane, 32 pp. Hirmer, M., 1927. Handbuch der Pal/iobotanik. Bd. I.R. Oldenbourg, Munich, 708 pp. Holden, H.S. and Croft, W.N., 1962. The morphology of Tubicaulis africanus sp. nov. a fossil fern from Tanganyika. Bull. Br. Mus. Nat. Hist. Geol., 7:199-211. Kidston, R. and Gwynne-Vaughan, D.T., 1909. On the fossil Osmundaceae. Pt III. Trans. R. Soc. Edinburgh, 46: 651-667. Kidston, R. and Gwynne-Vaughan, D.T., 1910. On the fossil Osmundaceae. Pt. IV. Trans. R. Soc. Edinburgh, 47: 455-477. Kidston, R. and Gwynne-Vaughan, D.T, 1914. On the fossil Osmundaceae. Part V. Trans. R. Soc. Edinburgh, 50: 469-480. Miller, C.N., Jr., 1971. Evolution of the fern family Osmunda-

[58

ceae based on anatomical studies. Contrib. Mus. Paleontol. Univ, Michigan, 23(8): t05 169. Mittre, V., 1955. Osmundites sahnii sp. nov. a new species of petrified osmundaceous rhizomes from India. Paleobotanist, 4:113 118, Phillips, T.L. and Andrews, H . N , 1966. Catenopteris simplex gen. et. sp. nov.~ a primitive pteridophyte from the Upper Pennsylvan[an of Ilinois. Bull. Torrey Bot. Club, 93(2): 117~128. Renault, B., 1896. Bassin houiller et permian d ' A u t u n el d'Epinae, Bd. II, pp.45 47, Rigby, J.F., 1978. Jurassic plant fossils from the Walloon Coal Measures at Rosewood Consolidated Colliery. Queensl, Gov. Min. J., 79: 526-529, Sahni, B., 1930. On Asterochlaenopsis, a new genus of zygopterid tree-ferns from western Siberia, Philos. Trans. R. Soc. London, 218B: 447-471.

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