Records of the Westem Australian Museum 18: 19-65 (1996),

Early Cretaceous macrofloras of Western Australia

Stephen McLoughlin School of Botany, The University of Melbourne, Parkville, Victoria 3052, Australia

Abstract - Western Australian, Lower Cretaceous, macrofloras from the Broome Sandstone and Callawa Formation (Canning Basin), Nanutarra Formation and Birdrong Sandstone (Carnarvon Basin), Cronin Sandstone (Officer Basin), and Leederville and Bullsbrook Formations (Perth Basin) incorporate a range of lycophytes, ferns, pteridosperms, bennettitaleans, and conifers, The new monotypic genus Roebuckia is established for spatulate fern fronds (R, spatulata) of possible vittariacean alliance. Other newly established species include Phyllopteroides westralensis, Elatocladus ginginensis, and Carpolithes bullsbrookensis, Although the Western Australian fossil suites reveal some specific differences from other Australian late Mesozoic assemblages, several shared index taxa and the high proportion of bennettitaleans support correlation with the Victorian Neocomian Ptilophyllum-Pachypteris austropapillosa Zone (Zone B). Some Western Australian taxa are shared with Indian assemblages but fewer similarities exist with other Gondwanan Early Cretaceous floras, The representation of several hydrophilous fern, lycophyte, and pteridosperm groups together with growth indices from fossil woods implies a seasonal humid mesothermal climate for the Western Australian eratonic margin during the Neocomian-Barremian. Minor differences between the Western Australian assemblages are attributable to local depositional and preservational factors.

INTRODUCTION assemblages of plant macrofossils from subsurface This paper reviews all Western Australian Cretaceous strata in Western Australia. Retallack Cretaceous plant fossils held in the collections of (1977, 1980) and McLoughlin (1993) have the Western Australian Museum and the illustrated differences in palaeosol development Department of Geology and Geophysics of the and the composition of plant fossil assemblages in University of Western Australia together with Australian Permian and Triassic strata according limited material held in the Australian Museum, to variations in local and regional depositional Sydney, As all specimens lack preserved organic settings. Such sedimentological and taphonomic matter, descriptions are based on gross controls also apply to the Cretaceous assemblages morphology, ornamentation, and venation offering an important tool for detailed patterns. This procedure necessarily places interpretation of non-marine sedimentary facies. limitations on the degree of discrimination between Additionally, permineralized, mollusc-bored, some taxa. However, the scarcity of previous driftwood preserved in Lower Cretaceous marine studies on the Western Australian fossil floras sediments has aided the definition of sequence invites this systematic appraisal in order to fill stratigraphic boundaries and systems tracts some major gaps in the knowledge of Australia's (McLoughlin et al. 1994). Palaeobotanical Cretaceous phytogeography. investigations of the diversity of plant groups The relative tectonic stability and deep­ together with anatomical indices such as growth weathering associated with much of the central ring variation are also useful for the categorization and western part of the Australian continent since of palaeoclimates and the identification of major the mid-Mesozoic has removed from outcrops food sources available to the large Cretaceous most of the organic material required for herbivorous dinosaurs. palynological studies. Apart from scarce vertebrate trackways (Colbert and Merrilees 1967; Long 1990), plant macrofossils are virtually the only fossil PREVIOUS STUDIES remains available in outcrop to date and Douglas (1969, 1973), Drinnan and Chambers palaeoenvironmentally categorize Western (1986) and McLoughlin et al. (1995) have provided Australian Cretaceous terrestrial strata. No studies the most detailed studies of Australian Cretaceous dealing with stratigraphic and petroleum fossil plants dealing with macrofloras from the exploration drilling have reported significant Gippsland, Otway and Eromanga Basins. Other 20 s. McLoughlin

113°E

o 600

KILOMETRES

Figure 1 Map of Western Australia showing Phanerozoic sedimentary basins and principal Cretaceous plant macrofossil localities (solid squares). significant contributions include those of Seward McLoughlin et al. (1994) detailed the anatomical (1904), Chapman (1908, 1909, 1914), Walkom (1918, character and palaeoenvironmental significance of 1919a), Medwell (1954a,b), Glaessner and Rao silicified and phosphatized woods from the (1955), and Dettmann et al. (1992). Although Birdrong Sandstone, Carnarvon Basin and several stratigraphic papers have provided lists of Backhouse et al. (1995) described lignitized but fossil plants recovered from Western Australian anatomically similar woods from the Barremian to Cretaceous strata (e.g., Brunnschweiler 1960; Cox earliest Aptian Nakina Formation of the Collie 1961; Playford et aI. 1976) many of these Basin. The Cretaceous palaeobotanical literature identifications are not supported by illustrations or from other Gondwanan continents (especially India descriptions and are commonly erroneous. A small and South America) is extensive. Therefore, assemblage described and illustrated by Walkom synonymy lists provided here deal only with (1944) from strata underlying the Molecap previously figured or described Western Greensand near Gingin is redescribed herein. Australian fossil plants. White (1961a) briefly described and illustrated plant remains recovered from the Canning Basin by Australian Bureau of Mineral Resources CRETACEOUS STRATIGRAPHY regional geological surveys. McLoughlin and Western Australian Cretaceous plant Guppy (1993), and McLoughlin and Hill (in press) macrofossils have been recovered from the Perth, provided preliminary revised lists of Western Carnarvon, Canning, and Officer Basins (Figure 1). Australian Cretaceous plant fossils from several Plant remains are best represented in Early sedimentary basins and briefly discussed some Cretaceous non-marine strata (Figure 2). Due to a palaeoclimatic implications of the floras. dearth of palynological studies and the absence of ------

Early Cretaceous macrofloras 21

marine invertebrate biostratigraphic indices, the However, Feldtman (1963) reported that the plant­ ages of many of these rock units are only broadly bearing horizons occurred within the Strathalbyn constrained. Furthermore, lack of continuous Beds, a term which Playford et al. (1976) regarded outcrops in many areas hinders the mapping and as synonymous with the Leederville Formation. lithological correlation of plant-bearing strata and Wilde and Low (1978) also mapped outcrops in the has led to uncertainties over the application of vicinity of Walkom's plant beds (Cheriton Creek) lithostratigraphic names. For example, Playford et as part of the Leederville Formation. On the basis al. (1976) and Skwarko (1990) assigned Walkom's of lithology and stratigraphic position, the fossils (1944) Gingin fossil assemblage to the Yarragadee are here regarded as belonging to the marine to Formation (Middle Jurassic to Tithonian). continental Leederville Formation (Hauterivian-

LOWER PERTH BASIN CARNARVON CANNING BASIN CRETACEOUS (Dandara~an BASIN (Anketell Shelf & STAGES Trough kGascoyne Broome Platform); ub-basin) OFFICER BASIN

Osborne ALBIAN Formation

Windalia Radiolarite 0 n ete s rezie 0 / ~ .-l «.-l (9 APTIAN iJ u Muderong Dandaragan Sh Sandstone Cl. ::J 0 0 0 .... Birdrong Sst .!.!!c .:::&.c <.!J =0 00 BARREMIAN c: '';= 0·- 0 m eel ..a«l .... "OEm .... -ClJE.... .0 mo "So c ...JLL COLL .... 0 eel m 0 ~ r-- HAUTERIVIAN s: m 0 E Z 0 c m South 0> 0 c m -.~ -0 -c « - c c -Perth 0 0 0 1i5 ~ Shale () .~ "0 1i5 E C "0 VALANGINIAN eel - 0 eel C 0 E LL (j) eel 0 <5 .... (j) 0 0 eel m 0 ~ LL ~ E c W ...... eel eel 0 C Z ~ .... Cii ....0 0.... eel () CO () BERRIASIAN S Parmelia c Formation zeel Fossil leaves _ Permineralized wood 0

Figure 2 Stratigraphic correlation of Lower Cretaceous rock units in the Perth, Carnarvon, and Canning Basins, Western Australia (after Cockbain 1990; Hocking 1990; Middleton 1990) showing the distribution of plant macrofossi! assemblages. 22 s. McLoughlin

Barremian): Figure 2. The Bullsbrook Formation, preserved in ferruginous shale and medium­ exposed 2 km east of Bullsbrook, is a fully grained quartzose sandstone apart from some continental lateral equivalent of the Leederville seeds, megaspores, and axes which are preserved Formation (Figure 2). as natural day or sandstone casts. Organic material The Neocomian-?Aptian shallow marine to was not preserved on any fossils but in some cases paralic Birdrong Sandstone is chiefly exposed latex and silicone casts yielded morphological along the Murchison River near Kalbarri, on the details not clearly evident on negative impressions. flanks of the Kennedy Ranges, and in the Giralia All specimens were illustrated under strong Anticline within the Carnarvon Basin (Hocking et unilateral light directed from the upper left unless al. 1987; McLoughlin et al. 1994). The upper part of otherwise stated. this unit is typically rich in fragmentary Fossil localities are as follows: permineralized gymnosperm wood showing Broome Sandstone - Gantheaume Point, Broome evidence of intensive molluscan borings. Other (material collected by W.H Butler, A. Dawson, Cretaceous marine units in the Carnarvon Basin Land T. Foulkes, J. Long, D. and M. Merrilees, contain sparse fossil woods, and leaf impressions J. Pas, G. Zeck); Quandong Beach, 10 km north occur within the paralic Nanutarra Formation near of Broome lighthouse (material collected by the Ashburton River (Figure 1). Lignified wood is LM. Doust). also locally preserved in the Lower Cretaceous Nanutarra Formation - Cited as "6.5 km north of Nakina Formation, Collie Basin, in the state's Globe Hill, Ashburton River" on specimen southwest (Backhouse et al. 1995). labels (material collected by D.W. McLeod). The Broome Sandstone of the northern Canning This is probably the plant fossil locality 2.5 km Basin was deposited within deltaic to shallow north of the junction of Jubricoo Creek and marine environments whereas the laterally Ashburton River, 8 km west of Nanutarra equivalent Callawa Formation in the basin's indicated by van de Graaf et al. (1980). southwest is fluvial in origin (Towner and Gibson Birdrong Sandstone Near Toolonga Hill 1983). The coeval Cronin Sandstone, outcropping (114°16'6"E 27°34'3"S), Murchison House in the northwestern part of the contiguous Officer Station, 10 km northeast of Kalbarri (material Basin (Figures 1,2) was also laid down in fluvial collected by S. McLoughlin); 1 km northwest of environments (Towner and Gibson 1983). These Black Dam (114°1O'9"E 23°6'3"S), Cardabia plant-bearing units have been regarded as Late Station, 35 km east of Coral Bay (material Jurassic to Early Cretaceous or even fully Jurassic collected by D.W. Haig, R. Howe, G. Ellis, S. on the basis of previous phytostratigraphic McLoughlin). interpretations (Middleton 1990). However, a Leederville Formation - Cheriton Creek, about 500 Neocomian-Barremian age for the plant-bearing m east of MacIntyre Gully, 2.5 km north of portions of these units is proposed herein based on Gingin (material collected by E. de C. Clarke). revised fossil identifications. Bullsbrook Formation - 2.5 km northeast of Bullsbrook Hotel (material collected by A. Page). MATERIAL AND METHODS The majority of material described and figured in this study is held in the collections of the SYSTEMATIC PALAEOBOTANY Western Australian Museum (specimens prefixed WAM) and the Department of Geology and Division Lycophyta Geophysics, University of Western Australia Class Lycopsida (specimens prefixed UWA). Counterparts of some of Walkom's (1944) specimens are lodged with the Order Isoetales Australian Museum in Sydney (specimens prefixed Family Isoetaceae AMF). The Commonwealth Palaeontological Collection housed by the Australian Geological Genus Isoetites Munster, 1842 Survey Organisation, Canberra, contains limited Type species Canning and Officer Basin material from regional Isoetites crociformis Munster, 1842; Jurassic; geological surveys of the 1950s and 60s and has Daiting near Manheim, Germany. been illustrated by White (1961a). The latter material was not re-examined during this study but in most cases White's (1961a) illustrations are Isoetites elegans Walkom, 1944 adequate for identification purposes and her Figures 3A, C-F studied specimens are systematically discussed herein. 1944 Isoetites elegans Walkom; p. 202; plate 1, figs I­ All descriptions are based on impression fossils S. Early Cretaceous macrofloras 23

1986 Megasporangia of lsoetes; White; p. 169; figure Roy, 1964), air channels within the leaves (lsoetites 260. horridus (Dawson) Brown, 1939), the details of megaspores and cuticle (lsoetites choffati Saporta, 1993 lsoetites elegans; McLoughlin and Guppy; p. 1894, lsoetes reticulata Hill, 1987), leaf proportions 14; figure 12. (lsoetites gramineoides Bock, 1962, lsoetites rolandii Holotype Ash and Pigg, 1991), and corm dimensions (lsoetites UWA16687; Leederville Formation (Neocomian­ crociformis Munster, 1842, lsoetites circularis Barremian), Perth Basin, Western Australia (Emmons) Brown, 1958). (Walkom 1944). lsoetes janaianus Banerji, 1989 from the Middle to Upper Jurassic of India has similar leaves and sporangia to l. elegans but is characterized by a 5­ Material lobed rhizomorph whereas corms are not yet UWA16682, UWA16683 (principal sporangia­ available for the latter species. lsoetites bulbiformis bearing specimen), UWA16684, UWA16685, Drinnan and Chambers, 1986 from the Victorian UWA16687 (principal foliage-bearing specimen), Lower Cretaceous, distinguished by its smaller size UWA16690, UWA16703, UWA16711. Counterparts and expanded leaf tips, may represent of some specimens are held in the Australian gymnospermous remains. The Indian Lower Museum (AMF39815: see White 1986, figure 260). Cretaceous lsoetites indicus Bose and Roy, 1964, although represented only by sporophyll bases, is Distribution characterized by its smaller but more numerous Leederville Formation (Neocomian-Barremian), megaspores in each sporangia (Bose and Banerji Perth Basin, Western Australia. Previous 1984). assignation of these specimens to the Jurassic (Walkom 1944; Ash and Pigg 1991) is now deemed erroneous. Genus Nathorstianella Glaessner and Rao, 1955 Type species Description Nathorstianella babbagensis (Woodward) Glaessner Numerous (>30) radiating linear leaves, reaching and Rao, 1955; Lower Cretaceous; Mt Babbage, at least 10 cm long, 5 mm wide, bases not South Australia. preserved, apices pointed, margins entire (Figure 3A). Longitudinal striae evident on some leaves but venation otherwise indistinct. Associated Nathorstianella babbagensis (Woodward) megasporophylls on type locality slabs bear Glaessner and Rao, 1955 clusters of numerous (>30) trilete megaspores. Figure 3B Megasporophylls identical to sterile leaves, cf. 1961 "fragment of stem" White; p. 304; plate 6, showing no expansion around sporangia. figure 3. [1961a]. Megasporangia reach 10 mm long, 2.5 mm wide (Figure 3C). Megaspores 0.5 mm in diameter, circular in polar compression, laesurae prominent; Holotype contact surfaces extending almost to the equator, F15070 (Adelaide University Geology laevigate; distal surface reticulate (moulds coarsely Department); Algebuckina Sandstone granulate: Figure 3D). Associated (Neocomian); Mt Babbage, South Australia microsporophylls bear elliptical microsporangia, 10 (Glaessner and Rao 1955). mm long, 3 mm wide, consisting of densely matted casts of ill-defined microspores (Figures 3E,F). Material WAM P.88.2, WAM P.96.12. Comments Although the earliest records of lsoetites are from Distribution the Upper Triassic (Bock 1962), they underwent a Algebuckina Sandstone (Berriasian-early phase of global diversification during the Early Valanginian), Eromanga Basin, South Australia Cretaceous (Pigg 1992). Ash and Pigg (1991) listed (Dettmann et al. 1992); Broome Sandstone the fossil species previously assigned to lsoetites (Neocomian-Barremian), Canning Basin, Western Munster 1842 and lsodes Linnaeus 1753. All Australia (this study). examples essentially consist of linear leaves or sporophylls either in isolation or attached in Description rosettes to short cylindrical or spherical corms. Fragmentary casts of cylindrical axes (14-17 mm Other species can be distinguished from 1. elegans wide, 24-35 mm long) bearing numerous spirally by possession of serrate leaf margins (lsoetites arranged leaf or root scars (Figure 3B). Scars serratus Brown, 1939, lsoetites serratifolius Bose and diminutive (0.5 mm wide, 0.25 mm high), c. 0.5-1 ------

24 s. McLoughlin Early Cretaceous macrofloras 25

mm apart, arranged in flat (70°-75°) right-handed Material and steep (50°) left-handed spirals (measured with WAM P.65.42, WAM P.88.13, WAM P.89.166, respect to axial orientation). Axes show broad WAM P.89.181, WAM P.96.9. transverse ridges or undulations corresponding to growth increments. Attached roots unavailable. Distribution Leaves unavailable on curated specimens. Broome Sandstone (Neocomian-Barremian), Canning Basin; Lees Sandstone (Neocomian?), Comments Carpentaria Basin, Northern Territory (White Glaessner and Rao (1955) suggested a possible 1961b; Dettmann et al. 1992). relationship between leafless Nathorstianella corms and foliage assigned to Isoetites elegans. Leaves 4 Description

mm wide, 60 mm long and similar to I. elegans L but Frond flabellate, up to 42 mm long and 47 mm lacking sporangia, were observed attached to N. wide; base cuneate, tapering to long petiole (Figure babbagensis axes in outcrops of the Broome 4C); apex rounded, undulate, or broadly dentate. Sandstone at Gantheaume Point near Broome Single vein enters base of frond from petiole, during this study. The specimens are preserved in bifurcates up to six times producing a radiating rock slabs too large to be collected or extracted array of dominant veins; each ultimate dominant without risking damage to the fossils. It seems vein enters base of a marginal tooth (Figure 4A). likely that I. elegans leaves and N. babbagensis Up to five orders of closely anastomosing belonged to the same plant although formal subsidiary veins subdivide lamina between synonymy should wait until fertile material is dominant veins into a meshwork of <0.25 mm2 found attached to the latter. The occurrence of N. aureoles. Petiole 1 mm wide, >6.5 cm long. babbagensis in sandy deltaic sediments of the Sporangia not evident. Broome Sandstone suggests that this plant preferred coastal habitats similar to those occupied Comments by related herbaceous to arborescent Triassic Hausmannia has affinities with reticulate-veined pleuromeian lycophytes (Retallack 1977; Karrfalt pinnatifid dipteridacean fronds such as Goeppertella 1986). Oishi and Yamashita emend. Arrondo and Petriella, 1982, Dictyophyllum Lindley and Hutton, 1834, Thaumatopteris Goeppert, 1841, and Clathropteris Division rteridophyta Brongniart, 1828 (Herbst, 1975). Hausmannia had a Class Filicopsida cosmopolitan distribution during the Jurassic and Cretaceous (Corsin and Waterlot 1979) with several Order Filicales species known from gondwanan continents, Family Dipteridaceae although some are based on scanty material. Hausmannia papilio Feruglio emend. Herbst 1960 Genus Hausmannia Dunker, 1846 fronds from the Early Cretaceous of Argentina are Type species most closely comparable to the Western Australian Hausmannia dichotoma Dunker, 1846; Jurassic; specimens although examples of the former do not near Buckenburg, Bavaria. show long petioles and are more circular with slightly auriculate bases (Herbst 1960). Hausmannia pachyderma Sukh-Dev 1972b and specimens Hausma111lia sp. assigned to Hausmannia crenata (Nathorst) Maller Figures 4A,C from India (Bose and Sah 1968) can be distinguished from the Western Australian form 1961 Hausmannia sp.; White; p. 306; plate 8, figure by their undulate margins and coarsely reticulate 4. [1961a]. venation. The Indian 11. crookshanki Shah and 1993 Hausmannia sp.; McLoughlin and Guppy; p. Singh, 1964 differs by its apically cleft lamina. 14; figure 7. Of the previously described Australian

.... Figure 3 A [soetites e[egans Walkom 1944; Rosette of linear univeined leaves; UWA16687; Leederville Formation; x 0.8; B, Nathorstianella babbagensis (Woodward) Glaessner and Rao, 1955; Corm impression with spirally arranged leaf/root scars; WAM P.88.2; Broome Sandstone; x 3; C, Megasporangia of [soetites e[egans Walkom, 1944; UWA16683; Leederville Formation; x 8; D, Scanning electron micrograph of Isoetites e[egans Walkom, 1944 megaspores; AMF39815; Leederville Formation; x 100; E, Microsporangia (Mi) and megasporangia (Me) of Isoetites elegans Walkom, 1944; UWA16683; Leederville Formation; x 4; F, Scanning electron micrograph of surface of [soetites elegans Walkom, 1944 microsporangium showing ill-defined microspore outlines; AMF39815; Leederville Formation; x 1000. 26 s. McLoughlin Early Cretaceous macrofloras 27

Hausl11amlia species, Herbst (1979) considered that 6 mm wide, 18 mm long; apex rounded to obtusely only H. wilkinsii Walkom, 1928, H. sp. cf. H. pointed, margin entire, basiscopic margin straight defarrariisii Feruglio, 1937, and H. bulbaformis or convex, acroscopic margin straight (Figures Douglas, 1973 were distinguishable. Hausmannia 4B,D). Adjacent pinnule bases contiguous. Pinnules wilkinsii and H. bulbaformis are both differentiated arise at c. 50° from rachilla. Pinnules have from the Western Australian form by their strongly alethopteroid venation; veins depart pinnule dissected lamina (Walkom 1928; Douglas 1973). midrib at 50°-60°, bifurcate once or twice, pass Hausmarlllia sp. cf. H. defarrariisii from the Middle straight to margin, no anastomoses (Figure 4G). Jurassic Walloon Coal Measures (Herbst 1979; Pinnule midribs evanescent c. 3 mm before apex. McLoughlin and Drinnan 1995) differs by its pronounced apical lamina incision and auriculate Comments base with respect to both Northern Territory Cladophlebis foliage is abundant and widespread fronds illustrated by White (1961b, 1966) and the in Permian to Cainozoic strata throughout the forms from the Broome Sandstone. It is likely that world. Many species referred to this genus have both the Northern Territory and Western been assigned to the Osmundaceae based on the Australian Early Cretaceous forms are conspecific. possession of osmundaceous sporangia and spores or by association with permineralized osmundaceous axes. Specimens lacking attached Order Osmundales axial or fertile remains are assigned to a large array Family Osmundaceae of form species based on frond and pinnule shape and venation patterns. The variation of pinnule Genus Cladophlebis Brongniart, 1849 shapes evident on different parts of some extant fern fronds suggests that description of inadequate Type species material may have led to a proliferation of names Cladophlebis albertsii (Dunker) Brongniart, 1849; for species. ?Jurassic; Germany. Cladophlebis Australian cladophleboid fronds from a wide range of localities and ages have been commonly assigned to Cladophlebis australis (Morris) Seward, Cladophlebis sp. cf. C. oblonga Halle, 1913 1904, a species originally erected for Triassic fronds Figures 4B,D,G from Tasmania (Morris 1845). The variation in Australian C. australis pinnule shapes appears to be 1961 Cladophlebis australis (Morris); White; p. 304; roughly equivalent to the range evident between plate 7, figures 1,6. [1961a]. the many South American species (Herbst 1971). 1993 Cladophlebis sp. cf. C. oblonga; McLoughlin and Morris's (1845) original illustrations of C. australis Guppy; figure 2. show slender tapering falcate pinnules quite different from the Western Australian form Material described here. The studied specimens are WAM P.64.5, WAM P.65.40, WAM P.88.9, WAM compared with C. oblonga Halle, 1913 based on P.88.10, WAM P.88.12. their relatively oblong pinnules with rounded or weakly pointed apices although some Antarctic Distribution and South American examples of this species have Broome Sandstone, Canning Basin; Bullsbrook slightly shorter and more falcate pinnules than the Formation, Perth Basin; Algebuckina Sandstone, Western Australian forms (Halle 1913; Herbst 1971; South Australia (all Lower Cretaceous). Arrondo and Petriella 1980). Though poorly preserved and slightly smaller, White's (1961a) Description oblong Cladophlebis australis pinnae from the Fronds pinnate or bipinnate (imparipinnate). Callawa Formation appear to be conspecific with Rachis prominent, striate, <1.5 mm wide, gently the Broome and Bullsbrook fronds as are the South tapering distally. Pinnules alternate, oblong, up to Australian specimens of Glaessner and Rao (1955).

.... Figure 4 A, Hallsmallllia sp.; WAM P.88.13; Broome Sandstone; x 2; B, Cladophlebis cf. oblollga Halle, 1913; UWA10465A; Bullsbrook Formation; x 2; C, Hallsmallllia sp.; Two fragmentary fronds with long slender rachises; WAM P.89.181; Broome Sandstone; x 1; D, Cladophlebis cf. oblollga Halle, 1913; Pinna with preserved apex; WAM P.88.12; Broome Sandstone; x 1.5; E,F,H, Isolated pinnules of Phyllopteroldes westralellsis sp nov.; (E) UWA10473, (F) UWAI0473, (H) UWA10467C; Bullsbrook Formation; all x 3; G, Cladophlebis cf. oblonga fIalle, 1913 showing details of venation; WAM P.88.9; Broome Sandstone; x 2; I, Microphyllopteris gleichenioutes (Oldham and Morris) Walkom 1919a; Enlargement showing details of pinnae; WAM P.88.1; Broome Sandstone; x 3. 28 S. McLoughlin

Walkom's (1919a) Early Cretaceous C. australis (Neocomian-Barremian); Bullsbrook, Perth Basin, specimens from Queensland are also comparable Western Australia. in terms of pinnule shape but appear to have more prominent, but less divided, secondary veins. The Etymology Western Australian fronds have similarly Signifying the Western Australian distribution of proportioned pinnules and venation patterns to the species. Cladophlebis sp. cf. C. oblonga Halle, 1913 of Drinnan and Chambers (1986) although the Victorian Material examples have more sharply acute apices. WAM P.96.11. UWAI0461D, UWAI0462A, The oblong shape and generally rounded pinnule UWA10464A, UWAI0467C, UWAI0470A-B, apices of the Western Australian specimens UWA10472A, UWA10473B-D, E-F, UWAI0474C. distinguish them from many of the Indian Mesozoic Cladophlebis species which typically have either shorter elliptical or more narrow and Distribution elongate (lanceolate to falcate) pinnules often with Broome Sandstone, Canning Basin; Bullsbrook acutely pointed apices (Bose and Sah 1968; Sukh­ Formation, Perth Basin; Cronin Sandstone, Officer Dev 1972a; Zeba-Bano 1980; Roy 1968; Bose and Basin (all Neocomian-Barremian). Banerji 1984; Kasat 1970). Herbst (1971) reviewed South American Diagnosis Mesozoic fronds assigned to 17 Cladophlebis Fronds imparipinnate; pinnules narrowly species. Of these, C. denticulata Brongniart, 1828 elliptical to lanceolate, base tapering acute with and C. patagonica Frenguelli, 1947 have similarly short petiolule, apex rounded, margin entire or sized oblong pinnules to the Western Australian gently undulate; secondary veins once or twice specimens but are distinguished by their dichotomous, 15-20 veins per cm, intersecting prominently pointed acute apices (Baldoni 1980a; margin at 20°-40°. Longobucco et al. 1985). The Western Australian specimens compare favourably to Archangelsky's Description (1964b) Cladophlebis sp., C. oblonga Halle, 1913, C. Fronds pinnate (imparipinnate). Rachis slender pintadensis Herbst, 1966, and C. haiburnensis var «2 mm) wide. Pinnules narrowly elliptical to rectimarginata Herbst, 1966 and it appears that few lanceolate, reaching 10 mm wide, 40 mm long, individual features enable consistent differentiation inserted alternately on rachis at c. 40° (Figure 5A). of these species in the absence of fertile or cuticular Base tapering acute, joined to rachis by short «2 material. mm petiole), apex rounded, margin entire or gently South African Early Cretaceous Cladophlebis undulate. Pinnules show weakly defined bezuidenhoutensis Gianniny and Wiens and C. evanescent midrib. Secondary veins once or twice dunbrodiensis Gianniny and Wiens (in Anderson dichotomous, arch gently from indistinct midvein and Anderson, 1985) fronds clearly differ from the to intersect margin at 20°-40°, 15-20 veins per cm; Western Australian form by their undulate pinnule no anastomoses (Figures 4E,F,H). margins. Other South African species differ by their smaller, more elongate, contracted or falcate Comments pinnules (Anderson and Anderson 1985). Cantrill and Webb (1987) re-evaluated the genus Phyllopteroides and indicated its likely affinity to the Osmundaceae based on the association of Genus Phyllopteroides Medwell, 1954b sterile P. dentata fronds with osmundaceous Type species sporangiate remains (Caecumen expansa Cantrill and Phyllopteroides dentata Medwelt 1954b; Early Webb, 1987). Five species (viz., P. dentata Medwell, Cretaceous; Killara, Otway Basin, Victoria. 1954b, P. lanceolata (Walkom) Medwell, 1954b, P. serrata Cantrill and Webb, 1987, P. laevis Cantrill and Webb, 1987, and P. macdymontae McLoughlin, Phyllopteroides westralensis sp. novo Drinnan and Rozefelds, 1995) have been described Figures 4E,F,H; 5A from Australia. The entire-margined lanceolate Western Australian leaves compare most closely 1%1 Linguifolium denmeadi Jones and de Jersey; with P. lanceolata identified from Albian strata of White; p. 302; plate 5, figure 3. [1961a]. central Queensland (Walkom 1919a). However, P. 1993 Phyllopteroides lanceolata; McLoughlin and westralensis sp. novo differs by its consistently entire Guppy; figures 4, 11. pinnule margins, substantially lesser venation angle, and generally lesser vein densities. Holotype Phyllopteroides dentata, P. serrata and P. UWAI0473C (Figure 5A); Bullsbrook Formation macdymontae can be distinguished by their strongly Early Cretaceous macrofloras 29 dentate or serrate margins. Phyllopteroides laevis Description leaflets may also have entire margins, however, Frond at least bipinnate (Figure 5B). Rachis they differ from P. lanceolata by their more broadly slender «2 mm wide). Pinnae linear, opposite to elliptical shape. White's (1961a) Lingui.folium subopposite, arched distally, up to 2.5 mm wide, denmeadi from the Cronin Sandstone, Officer Basin, 30 mm long, inserted on rachis at 70°-90°. Pinnules is here regarded as a detached P. westralensis minute (1 x 1 mm), showing full basal attachment, pinnule. semicircular, alternate, apices rounded (Figure 41). Cantrill and Webb (1987) suggested that the No sporangia evident. successive stratigraphic ranges of P. laevis, P. serrata, and P. dentata in Victoria represented Comments elements of an evolutionary lineage. They also Arber (1917) established Microphyllopteris for suggested that the Queensland P. lanceolata arose fronds resembling extant Gleichenia ferns but from the same Neocomian-Aptian stock but lacking definitive evidence for a relationship to that evolved independently of the Albian P. dentata of genus. However, many workers have continued to Victoria. The Neocomian-Barremian P. westralensis assign such fronds to Gleichenites Goeppert, 1836 or may represent a separate lineage derived from Gleichenia Smith, 1793 even in the absence of fertile entire-margined P. laevis stock which became material. Microphyllopteris gleichenioides (Oldham restricted to the lowlands of the continent's and Morris) Walkom 1919a and comparable forms western margin. Few fossil fronds outside have been reported from Upper Jurassic to mid­ Australia are comparable with P. westralensis Cretaceous strata in Queensland (Walkom 1919a, although Thinnfeldia chunakhalensis Sah and Sukh 1928; McLoughlin et al. 1995). Gleichenites Dev, 1958 may be referable to Phyllopteroides based nanopinnatus (Douglas) Drinnan and Chambers on its P. laevis-like pinnules. 1986 from the Lower Cretaceous of Victoria has larger and more dissected pinnules than M. gleichenioides. Order Gleicheniales Gleichenites sp. of Arrondo and Petriella (1980) from the Argentinian Jurassic is a much larger Family Gleicheniaceae frond than M. gleichenioides but has similar Genus Microphyllopteris Arber, 1917 pinnules and may be closely related to the Australian species. Gleichenites sanmartinii Halle Type species emend. Herbst 1962a and G. vegagrandis Herbst, Microphyllopteris pectinata (Hector) Arber, 1917; 1962a from the Cretaceous of Argentina and Lower Jurassic; Mataura Falls, New Zealand. Antarctica also have similar diminutive semicircular pinnules although these are slightly inclined distally (Herbst 1962a, b). Microphyllopteris gleichenioides Some Indian examples of 'Gleichenites (Oldham and Morris) Walkom, 1919a gleichenoides' and Microphyllopteris sp. illustrated by Figures 41; 5B Sah (1965) and Bose and Sah (1968) differ slightly from the Australian M. gleichenioides in having 1993 Microphyllopteris gleichenoides; McLoughlin pinnules which are strongly deflected towards the and Guppy; figures 5, 6. pinna apex (insertion angle about 60°). Lectotype No. 4444 (Geological Survey of India, Calcutta); Rajmahal Series (?Lower Cretaceous), Rajmahal Order ?Pteridales Hills, India. Selected by Bose and Sah (1968). Family ?Vittariaceae Material Roebuckia gen. novo WAM P.88.I. Type species Distribution Roebuckia spatulata sp. nov.; Broome Sandstone Broome Sandstone (Neocomian-Barremian), (Neocomian-Barremian); Gantheaume Point, Canning Basin; Dalrymple Sandstone (Upper Broome, Canning Basin, Western Australia. Jurassic), Laura Basin, Queensland (Walkom 1928); Burrum Coal Measures (Albian), Maryborough Diagnosis and discussion Basin, Queensland (Walkom 1919a); Rajmahal See under Roebuckia spatulata below. Series and equivalents (Lower Cretaceous) of Peninsula India (Seward and Sahni 1920; Bose and Distribution Sah 1968). Known only from the type formation. 30 s. McLoughlin Early Cretaceous macrofloras 31

Roebuckia spatulata sp. novo and species. Superficially, the specimen resembles Figures 5E,F a ginkgophyte leaf in terms of its divergent dichotomous venation. However, Mesozoic 1993 Indeterminate spatulate frond; McLoughIin ginkgophytes typically have broadly flabellate, and Guppy; p, 14; figure 9 (lower part), leaves with much finer and more closely spaced Holotype venation (Sah 1952, 1953; Baksi 1968; Mehta and WAM P.88,15 (Figures 5E,F); Broome Sandstone Sud 1953; Sah and Jain 1965; Bose 1958; Bose and (Neocomian-Barremian); Gantheaume Point, Sukh Dev 1960; Anderson and Anderson 1989), Broome, Canning Basin, Western Australia. The Broome Sandstone specimen is clearly a pteridophyte owing to the presence of rows of Etymology elliptical sori between the ultimate veins. Species Genus: after Roebuck Bay near Broome, Species: of the extant tropical epiphytic or lithophytic denoting frond shape, vittariacean fern genus Antrophyum Kaulfuss, 1824 have pendant spatulate fronds similar to Roebuckia Material but their veins are commonly reticulate with WAM P,88,15, flanking dichotomous or reticulate sori (Clifford and Constantine 1980; Piggott 1988). Roebuckia Distribution spatulata resembles individual oblanceolate Broome Sandstone (Neocomian-Barremian); pinnules of Adiantopteris oshimaensis Kimura, Canning Basin. Ohana and Aiba 1990 but is a larger leaf with more robust and less dense venation. Combined diagnosis for genus and species Frond oblanceolate, margin entire, apex rounded, base acute, petiolate. Single robust vein in petiole, repeatedly dichotomous and gently divergent Incertae sedis across lamina. Veins terminate at apex or distal Genus Aculea Douglas, 1973 part of lateral margin, Longitudinally elliptical sori inserted in depressions and arranged in 1-2 rows Type species between each pair of veins on distal portion of Aculea bifida Douglas, 1973; Lower Cretaceous; frond. Boola Boola Forest; Victoria.

Description Single available spatulate frond 13 mm wide, 56 Aculea bifida Douglas, 1973 mm long, Frond margin entire, apex rounded, base Figure 50 tapering acutely to a 15 mm long petiole (Figure 5E), Single vein emerges from base, bifurcates up cf. 1961 Stenopteris tripinnata (Walkom); White; p, to five times, Veins gently diverge distally, 302; plate 4, figure 6. [1961a], terminating at apical margin or distal part of lateral margin; no anastomoses, no marginal vein. Sori Holotype GSV60879 (Geological Survey of Victoria), impressions evident on distal portions of leaf (best Strzelecki Group, Zone B (Neocomian); Boola seen on latex cast). Sori elliptical, inserted in depressions, reaching 05 mm wide, 2 mm long, Boola Forest, Gippsland, Victoria. arranged in 1-2 rows between adjacent veins (Figure 5F), Sori have granular appearance (on Material casts) but details of sporangia or spores UWA120489. unavailable, Distribution Comments Strzelecki Group, Zones B-C (Neocomian­ Although only a single leaf of the type species is Aptian), Gippsland Basin, Victoria (Douglas 1973); known, the specimen is deemed sufficiently Broome Sandstone (Neocomian-Barremian), distinctive to warrant the erection of a new genus Canning Basin, Western Australia.

~ Figure 5 A, Phyllopteroides westralensis sp, nov,; Portion of frond; UWA10473C; Bullsbrook Formation; x 15; B, Microphyllopteris gleichenioides (Oldham and Morris) Walkom 1919a; WAM P,88.1; Broome Sandstone; x 1; C, Sphenopteris warraglllensis McCoy (in Stirling), 1892; WAM P,89,176; Broome Sandstone; x 1; D, AClllea In(ida Douglas 1973; UWA120489; Broome Sandstone; x 2; E, Latex cast of Roebllckia spatlllata gen, et sp, nov" WAM P,88.15 (left) overlying a fragmentary BlIcklandia sp, axis with spiral rhomboid leaf scars, WAM P,96,8 (upper right); Broome Sandstone; x 2; F, Enlargement of distal portion of latex cast of Roebllckia spatlllata frond showing elliptical sori between veins; WAM P,88,15; x 6; G, Sphenopteris sp, A; WAM P,89,186; Isolated pinna; Broome Sandstone; x 2; H, Sphenopteris sp, A; WAM P,89,167; Broome Sandstone; xl. 32 s. McLoughlin

Description Neotype Bipinnate, probably elliptical to rhomboid­ NMVP21341 (National Museum of Victoria), shaped frond. Rachis slender «1 mm wide) with Strzelecki Group, Zone C (Barremian-Aptian); prominent central longitudinal ridge (on Jeetho Valley, Victoria. Designated by Douglas impressions). Pinnae lanceolate, reaching 6 mm (1973). wide, 35 mm long, departing rachis at 20°-25°, alternate. Pinnules, needle-like or slightly flattened Material (Figure 50), sub-opposite to alternate, <1 mm WAM P.89.176. wide, reaching 12 mm long, with single median vein persisting to apex; base decurrent, apex pointed acute. Distribution Broome Sandstone (Neocomian-Barremian), Canning Basin. Comments Although only a single sterile frond portion is available from the Broome Sandstone, it shows the Description deeply dissected lamina and univeined linear Fragmentary frond at least tripinnate, gross pinnules typical of Early Cretaceous Victorian shape uncertain. Rachis reaching 1.5 mm broad examples of Aculea bifida (Douglas 1973; Drinnan with a prominent central longitudinal groove (on and Chambers 1986). Dissected foliage of this type mould). First-order pinnae ovate to lanceolate had previously been assigned to Sphenopteris or bearing alternately arranged second-order pinnae Stenopteris species (Seward 1904; Chapman 1909). proximally but giving way to ultimate pinnules White's (1961a) Stenopteris tripinnata (Walkom) distally (Figure 5C). Lanceolate second-order from the Cronin Sandstone is comparable to A. pinnae depart midrib at about 20°. Lanceolate bifida although her figured specimen appears to ultimate pinnules arise from second-order pinnae have more numerous pinnules arranged in a more alternately at about 20°, bases decurrent, apices regular bipinnate fashion. Walkom's (1919a) pointed acute, margins entire. Venation Stenopteris laxum (Tenison-Woods) from the sphenopteroid but poorly defined. Maryborough Basin may also belong to this species. The Indian Sphenopteris metzgeroides Harris Comments of Bose and Banerji (1984) and Antarctic S. The single Broome Sandstone specimen nordenskjoldii Halle, 1913 may be synonymous with compares favourably with Sphenopteris A. bifida based on frond morphology but both lack warragulensis McCoy, (in Stirling) 1892 from fertile material comparable to the Australian Victoria (McCoy in Stirling 1892; Douglas 1973; species. Drinnan and Chambers 1986) although the details of the fine serrations on pinnules are poorly defined and sori are not preserved. Medwell's Genus Sphenopteris (Brongniart) Sternberg, 1825 (1954a) Sphenopteris hislopi Oldham and Morris Type species probably also belongs to S. warragulensis. The Sphenopteris elegans (Brongniart) Sternberg, 1825; narrow, elongate, acutely inserted pinnules Carboniferous; Silesia. differentiate this form from most other Sphenopteris species. Walkom's (1919a) S. flabellifolia Tenison­ Woods, 1883 from the Queensland Cretaceous is Discussion closely comparable to S. warragulensis and would Numerous species have been assigned to this have nomenclatural priority if examination of the cosmopolitan form genus based on subtle Queensland and Victorian specimens established differences in pinnule shape and venation pattern. their conspecificity. Sphenopterid fronds may represent foliage of true South American sphenopterid fronds (Herbst ferns or seed ferns in the absence of fertile material. 1964; Baldoni 1980a; Baldoni and de Vera 1980; They are here assigned to the Pteridophyta based Petriella and Arrondo 1984) generally differ from on the occurrence of fertile sphenopterid foliage S. warragulensis by their squat rhomboid or found in coeval strata elsewhere in Australia elliptical pinnules. Many Indian Jurassic­ (Drinnan and Chambers 1986). Cretaceous sphenopterid species can also be separated from S. warragulensis by their more compact elliptical to rhomboid pinnules (Jain 1968; Sphenopteris warragulensis McCoy, (in Stirling) Bose 1958; Bose and Sah 1968). Roy's (1968) S. 1892 specifica (Feistmantel) appears to have a lesser Figure 5C degree of frond dissection and more elliptical or 1993 Sphenopteris sp. A; McLoughlin and Guppy; oblanceolate ultimate pinnules than S. figure 3. warragulensis. Early Cretaceous macrofloras 33

Sphenopteris sp. A Incertae sedis Figures 5G,H Genus Thinnfeldia Ettingshausen, 1852 1961 'Neorhachopteris minuta' White; p. 303; plate 5, figure 4. [1961a]. Type species TI1innfeldia rhomboidalis Ettingshausen, 1852; cf. 1961 Ruffordia mortoni Walkom; White; p. 304; Early Jurassic; Steierdorf, Germany. plate 5, figures 6,7. [1961a].

1993 Sphenopteris sp. B; McLoughlin and Guppy; Thinnfeldia cf. talbragarensis Walkom, 1921b figure 8. Figures 6A,B 1944 Thinnfeldia talbragarensis Walkom; Walkom; p. Material 203; plate Il, figure 8. WAM P.64.14, WAM P.89.167, WAM P.89.170, WAM P.89.173, WAM P.89.175 WAM P.89.186. 1993 Thinnfeldia sp.; McLoughlin and Guppy; figure 10.

Distribution Material Broome Sandstone (Neocomian-Barremian), UWA16690, UWA10472C. Canning Basin. Distribution Leederville Formation and Bullsbrook Formation Description (Neocomian-Barremian), Perth Basin. Frond elliptical, at least bipinnate, 50 mm wide, 85 mm long. Rachis stout (2 mm wide) at base, Description tapering distally, straight or slightly sinuous. Frond at least bipinnate, >90 mm long, 60 mm Pinnae arranged alternately at 35°-50° to rachis, wide, elliptical. Rachis robust «3 mm wide), linear or arched distally, reaching 4 mm wide, 50 longitudinally striate. Pinnae opposite, ovate to mm long (Figure 5H). Rachilla with prominent elliptical, L:W ratio 2:1 to 4:1, arising from rachis at central longitudinal ridge (on moulds). Pinnules 60°-80°. Pinnae range from undulate-margined to inserted alternately by contracted or slightly deeply dissected into 4-7 pinnules (Figure 6A). decurrent base. Pinnules rhomboid to elliptical, Pinnule arrangement catadromous. Pinnules reaching 2 mm wide, 4 mm long; apices rounded elliptical, wide-obovate, or rhomboid, reaching 5 (Figure 5G). Venation dichotomous and gently mm wide, 8 mm long; subopposite to alternate; divergent from base. apex rounded; base decurrent; venation odontopteroid (Figure 6B). Comments This species is distinguished from S. Comments warragulensis by its small elliptical to rhomboid Walkom (1944) assigned material from the rather than lanceolate pinnules. Sphenopteris travisi Leederville Formation to Thinnfeldia talbragarensis, Stirling and Sphenopteris sp. of Drinnan and a species erected for Jurassic fronds from New Chambers (1986) and Ruffordia mortoni Walkom, South Wales, but these specimens, together with 1928 have more deeply dissected pinnules than additional specimens from the Bullsbrook Sphenopteris sp. A. The Jurassic Coniopteris sp. of Formation, differ from the eastern Australian form Hill et al. (1966) has similar small rhomboid in having ovate-elliptical rather than lanceolate pinnules but Sphenopteris sp. A has much longer pinnules. Walkom's (1944: plate Il, figure 8) and proportionately narrower pinnae. illustration appears to be upside-down. Thinnfeldia Herbst's (1964) Argentinian Sphenopteris sp. has media Tenison-Woods, 1883 figured by Jack and more rounded pinnules than the Western Etheridge (1892) from the Burrum Coal Measures, Australian form but the incomplete specimens of Queensland, and the Victorian Thinnfeldia sp. cf. T. the former inhibit close comparisons. Kasat's (1970) indica Feistmantel, 1876 of Drinnan and Chambers Cladophlebis sp. cf. C. longipennis Seward from the (1986) which is probably synonymous with T. Indian Early Cretaceous has a very similar gross maccoyi Seward, 1904 differ from the Western frond structure to Sphenopteris sp. A but its Australian fronds by their less-dissected lanceolate pinnules possess distinct midribs. Other Indian and pinnules. South African Sphenopteris species figured by Jain The Western Australian specimens bear some (1968), Sukh-Dev (1972a), Bose and Sah (1968), and similarities to Ruflorinia pilifera Archangelsky, Anderson and Anderson (1985) differ from the 1964a from the Cretaceous of Argentina, however, Western Australian specimen by their undulate, the former specimens lack cuticular details toothed, or deeply dissected pinnules. necessary for close comparison to the South ------~-~- -

34 s. McLoughlin Early Cretaceous macrofloras 35

American species. Thinnfeldia indica of Zeba-Bano Description et al. (1979) has proportionately more elongate and Leaves simple, up to 2 cm wide and 8.3 cm long, less dissected leaves than 1'. sp. cf. 1'. talbragarensis. lorate, linear, or spatulate. Margin entire or gently However, Jain's (1968) ?DicroidiulIl specimen from undulate, apex rounded obtuse, base tapering the Early Cretaceous of eastern India has very acute, petiolate. Midrib longitudinally striate, similar pinnule shapes and venation to the Western robust (up to 2 mm wide), gently tapering distally, Australian fronds and may be conspecific. persistent. Secondary veins closely spaced, depart midrib at >60°, bifurcate once or twice then pass straight to margin at 75°-90°. Lamina rarely shows Incertae sedis undulate folds. Attached fertile organs and cuticle Order Pentoxylales unavailable. Family Pentoxylaceae Comments Genus Taeniopteris Brongniart 1832 Drinnan and Chambers (1985) provided a detailed account of this Early Cretaceous species Type species based on material from Victoria. They suggested a Taeniopteris vittata Brongniart, 1832; Jurassic; relationship between 1'. daintreei leaves and both Whitby, England. the pollenate fructification Sa/mia laxip/lOra Drimlan and Chambers, 1985 and the ovulate pentoxylalean Taeniopteris daintreei McCoy, 1874 fructification Carnoconites cramvellii Harris, 1962, Figures 6C,F,G; 7A based on their co-occurrence in the Koonwarra fossil bed of the Gippsland Basin and similar co­ 1944 Taeniopteris spatulata McClelland; Walkom; p. fossilized forms in New Zealand, New South 203; plate Il, figure 9. Wales and India (Blashke and Grant-Mackie 1976; 1961 Taeniopteris cf. 1'. elongata Walkom; White; p. White 1981; Visshnu-Mittre 1953; Rao 1981). 303; plate figures lB, 2B. [1961a]. Drinnan and Chambers (1985) provisionally excluded White's (1981) New South Wales Jurassic 1993 Taeniopteris daintreei; McLoughlin and Guppy; Taeniopteris spatulata McClelland, 1850 from 1'. figure 14; figure 18. daintreei due to the discrepancy in the ages of the Victorian and New South Wales specimens. Lectotype Differences in the preservation of associated NMVP12270 (National Museum of Victoria); fruiting bodies also inhibited close comparison. Strzelecki Group, Zone C (Valanginian-Albian); However, Drinnan and Chambers (1985) noted that Cape Paterson, Victoria. Selected by Drinnan and previous distinctions between Victorian Cretaceous Chambers (1985). 1'. daintreei and Indian Jurassic-Cretaceous 1'. spatulata cuticle morphologies are probably less Material significant than initially suggested (Rao 1943, WAM P.64.12, WAM P.64.15, WAM P.74.26, Douglas 1969; Visshnu-Mittre 1957). The Western WAM P.74.27, WAM P.74.28, WAM P.74.29, WAM Australian specimens are morphologically 1'.74.32, WAM P.74.33, WAM P.74.34, WAM 1'.88.8. indistinguishable from Victorian 1'. daintreei leaves. UWA16685, UWA16682, UWA16683, Taeniopteris daintreei-like leaves are common UWA10374A,B,E, UWA10375, UWA10376, UWA­ components of several other Middle Jurassic to 10461A,B,C, UWA10463, UWA10464B,F,G,J,K, Early Cretaceous floras within Australia and New UWA10466A, UWA10470C,D, UWA10472B, Zealand (Dun 1898; Walkom 1919b, 1944; UWA10474A, UWA10477A,C. Glaessner and Rao 1955; White 1961a, b; Gould 1980; Blaschke and Grant-Mackie 1976). Several Distribution authors (e.g., Chapman 1909; Glaessner and Rao Widespread; probably upper Lower Jurassic to 1955, Douglas 1969; Baldoni and de Vera 1980; Lower Cretaceous of Australasia. Western Drinnan and Chambers 1985) have figured or Australia: Broome Sandstone, Canning Basin; described leaves which show varying degrees of Cronin Sandstone, Officer Basin; Bullsbrook and lamina dissection/ in some cases making them [,eederville Formations, Perth Basin. gradational with forms attributable to Nilssollia

.. Figure 6 A,B, Thil1l1{i'ldia cf. talbragarel1sis Walkom, 1921; (A) UWA16690, Portion of frond, Leederville Formation, x 1; (B) UWA10472C, Isolated pinna, Bullsbrook Formation; x 4; C,F,G, Portions of variably dissected Tael1iopteris damtrm McCoy, 1874 leaves; (C) UWA24640B, Bullsbrook Formation, x 1.5; (F) UWA120484, Broome Sandstone, x 3; (G) WAM P.74.26a, Bullsbrook Formation, x 1.5; D,E,H,t Plilophylllllll clIlchens" Morris (in Grant) emend. Bose and Kasat, 1972; (E) WAM P.88.7, Broome Sandstone, x 2; (H) WAM P.88.4, Broome Sandstone, x 0.9; (I) UWAl18836, Bullsbrook Formation, x 3. 36 s. McLoughlin

Brongniart 1825. Several specimens from the Description Bullsbrook Formation also display variable Fragmentary pinnate fronds with broad striate dissection but differ from Nilssonia species herein rachis «5 mm wide). Pinnules subopposite, which show distinct gaps between lamina truncate elliptical, semicircular, oblong, or slightly segments (pinnules). falcate, up to 15 mm wide and 25 mm long; apex rounded, asymmetrical; full basal attachment; inserted laterally? or adaxially? on rachis (Figure Incertae sedis 7F). Pinnule bases expanded slightly, basiscopic Pinnate frond margin rounded convex, acroscopic margin Figure 6D straight or slightly concave. Generally >10 veins arising from pinnule base, each bifurcates no more Material than twice (rarely more than once), passing sub­ UWA10475. parallel to apical and lateral margins without anastomoses. Distribution Bullsbrook Formation (Neocomian-Barremian), Comments Perth Basin, Western Australia. The full basal insertion of pinnules on the rachis distinguishes this leaf-form from Otozamites species Description which have pinnules with contracted bases (Taylor A single poorly preserved pinnate frond of and Taylor 1993). White's (1961b) Pterophyllum uncertain affinities occurs in the Bullsbrook fissum Feistmantel leaves are pinnatifid in contrast Formation assemblage. Frond 4 cm long, 2.2 cm to the pinnate Nilssonia sp. A. Several Nilssonia and wide, base and apex absent. Rachis broad (3 mm), Ctenozamites species from the Yorkshire Jurassic longitudinally striate, gently tapering distally; (Harris 1964) have similar sized fronds to the pinnules linear with full basal attachment, margins Western Australian examples but differ in the entire or gently undulate, venation indistinct details of their pinnule shapes and venation. (Figure 6D).

Remarks Nilssonia sp. B This specimen may represent a fern, seed-fern, or Figure 7B cycadophyte frond. The apparently undulate 1993 Nilssonia sp. B; McLoughlin and Guppy; p. 14; margins on some pinnules invite comparison to figure 17. Douglas's (1969) Pachypteris austropapillosa recorded from coeval "Zone B" floras of the Material Victorian Early Cretaceous but the specimen is too WAM P.88.5, WAM P.88.6. poorly preserved for identification. Distribution Broome Sandstone (Neocomian-Barremian), Division Cycadophyta Canning Basin. Order Bennettitales Description Genus Nilssonia Brongniart, 1825 Fragmentary elliptical to oblanceolate pinnate Type species fronds, petiolate; rachis up to 2 mm wide. Pinnules Nilssonia brevis Brongniart, 1825; Rhaetian; Hoer, with full basal attachment, semicircular to oblong, Sweden. apex rounded. Basiscopic margin of pinnules gently convex, acroscopic margin straight or slightly concave (Figure 7B). Pinnules alternate, up Nilssonia sp. A to 12 mm wide, 22 mm long. Numerous closely Figure 7F spaced, fine, indistinct, veins emerge from rachis, 1993 Nilssonia sp. A; McLoughlin and Guppy; p. diverge gently towards apex and lateral margins; 14; figure 15. bifurcations rare, anastomoses absent.

Material Comments WAM P.96.3. This leaf form differs from Nilssonia sp. A by its more divergent and closely spaced venation. Distribution White's (1961a) and Walkom's (1919a) Nilssonia Nanutarra Formation (Neocomian-?Aptian), schaumbergensis Dunker from the Broome Carnarvon Basin. Sandstone and Burrum Coal Measures respectively Early Cretaceous macrofloras 37 both have broader and shorter lamina segments 30°--65°, up to 3 mm wide, 40 mm long (Figures than either of the Western Australian Nilssol1ia 71,J). Pinnule bases rounded, apices pointed acute species described here. White's (1981) Nilssol1ia or truncate. Pinnules aligned in a common plane or compta (Phillips) Brongniart also has substantially slightly imbricate. Several veins enter base of shorter wedge-shaped pinnules. By contrast, pinnules, diverge slightly, pass straight to apical Nilssol1ia plutovillensis Walkom, 1928 has narrower margin, no anastomoses. pinnules with more acute apices. Comments Ptilophyllum acutifolium is distinguished from Genus Ptilophyllum Morris emend. Bose and other bennettitalean fronds in the collection by its Kasat,1972 very narrow and elongate pinnules which may be straight or gently falcate. This species is common Type species within the Broome Sandstone and Nanutarra Ptilophyllum acutifolium Morris, (in Grant) 1840; Formation and was identified by White (1961a) as Early Cretaceous; south of Charivar Range, India. Zamites sp. Ptilophyllum acutifolium is widely recorded from the Indian Early Cretaceous (Morris Discussion in Grant 1840; Feistmantel 1877a, b, c, 1879; Sahni Otozamites Braun, in Munster 1843 is traditionally and Rao 1933; Bose and Kasat 1972; Sukh-Dev and distinguished from Ptilophyllum by the Zeba-Bano 1980; Baksi and Naskar 1981; Bose and asymmetrical, expanded (auriculate) bases of its Banerji 1984). South American Jurassic and Early pinnules (Seward 1917). However, the genera may Cretaceous fronds have also been assigned to this be synonymous as several Ptilophyllum species also species (Arrondo and Petriella 1980; Longobucco et possess asymmetrical auriculate pinnules (Bose al. 1985) although both the Australian and South and Kasat 1972). In the absence of cuticular data, American forms tend to have longer pinnules than Western Australian bennettitalean fronds with the Indian examples. narrow pinnules can be compared with a wide range of Otozamites and Ptilophyllum species (Bose and Kasat 1972; Bose 1974).

Ptilophyllum cutchense Morris, (in Grant) Ptilophyllum acutifolium Morris, (in Grant) 1840 emend. Bose and Kasat 1972 Figures 71,J Figures 6E,H,I; 7C,D 1%1 Zamites sp.; White; p. 305; plate 8, figure 3. 1944 Ptilophyllum pecten (Phillips); Walkom; p. 204; [1961a]. plate 11, figure 6. 1993 Ptilophyllum acutifolium; McLoughlin and 1%1 Ptilophyllum pecten (Phillips); White; p. 302­ Guppy; p. 14; figure 16. 305; plate 6, figure 3D; plate 8, figure 1. [1961a]. 1%1 Otozamites bengalensis Oldham and Morris; Lectotype White; p. 305; plate 7, figure 5. [1961a]. V21330, British Museum (Natural History); Jabalpur Series equivalents (Lower Cretaceous); 1993 Otozamites bengalensis; McLoughlin and Kutch, India. Selected by Bose and Kasat (1972). Guppy; p. 14; figures 13, 14.

Material Lectotype WAM P.64.16, WAM P.89.162 WAM P.96.2, V20191 (9943), British Museum (Natural WAM P.96.4. History); Rajmahal Series equivalents (?Lower Cretaceous); Kutch, India. Selected by Bose and Distribution Kasat (1972). Lower Cretaceous of India; possibly Jurassic and Lower Cretaceous of South America; Nanutarra Material Formation, Carnarvon Basin; Broome Sandstone, WAM P.64.6, WAM P.64.7, WAM P.64.8, WAM Canning Basin (both Lower Cretaceous), Western P.64.13, WAM P.70.46, WAM P.86.57, WAM Australia. P.86.172, WAM P.88.4, WAM P.88.7, WAM P.88.1I, WAM P.89.169, UWA16684, UWA1I8836. Description Fragmentary, broadly elliptical pinnate fronds. Distribution Rachis stout (1-2.5 mm wide). Pinnules Broome Sandstone, Canning Basin; Leederville subopposite to alternate, linear, straight or arched and Bullsbrook Formations, Perth Basin (all distally, inserted on adaxial surface of rachis at Neocomian-Barremian). 38 S. McLoughlin Early Cretaceous macrofloras 39

Description Ptilopl1yllum boolellsis (Douglas> Douglas, 1969 honds linear, pinnate, petiolate, up to 16 mm Figure 7C wide, 155 mm long. Rachis stout, up to 1 mm thick. 1993 Ptiloplzyllum boolensis; McLoughlin and Pinnules inserted adaxially on rachis. 1'innule base Guppy; p. 14; figure 21. slightly auriculate or contracted (Figure 7C). Adjacent pinnules slightly imbricate. 1'innule bases Holotype abutting alternately on adaxial surface of rachis GSV58781 (Ceological Survey of Victoria); forming a zig-zag pattern on impressions (Figures Strzelecki Group, Zone B (Neocomian); Boola 6E, 7D). 1'innules oblong or falcate, apices pointed Boola Forest, Gippsland, Victoria (Douglas 1969). acute, slightly inclined distally (Figures 6H,I). I'innules arise at 50 0 --7()o from rachis, reaching 8 x Material 2 mIn. Veins emerge from centre of pinnule base, WAM 1'.96.13. dichotomize, slightly diverge, intersect apical and lateral margins without anastomoses. Distribution Comments Broome Sandstone (Neocomian-Barremian), Canning Basin; Strzelecki Group, Zone B In gross morphology these fronds are similar to a number of bennettitalean foliage species including (Neocomian), Victoria. Ptilophyllum pecten (1'hillips) Harris, 1969, P. horridum Roy, 1963, P. canthifera Douglas, 1969, and Description Otozamites exhislopi Bose, 1974. However, the Single diminutive lanceolate pinnate frond contracted pinnule bases, slightly expanded fragment preserved as an impression in siliceous basiscopic pinnule lamina, pinnule imbrication, siltstone. Rachis slender «1 mm wide). 1'innules and relatively short, slightly falcate pinnules with attached to upper surface of rachis. 1'innules very obtuse apices suggests that the Western Australian small «3 mm wide, <5 mm long), circular to specimens are better assigned to P. cutchense Bose elliptical, bases truncate to auriculate, apices and Kasat, 1972, a common Jurassic? to Early broadly rounded (Figure 7G). Pinnules inserted Cretaceous species of India and Australia (Walkom alternately at about 50° to rachis; adjacent pinnules 1944; Glaessner and Rao 1955; White 1961a,b; Bose slightly overlap producing a zig-zag pattern along and Kasat 1972). Most previous Australian studies rachis. Single vein enters centre of pinnule base have assigned such remains to Otozamites and dichotomizes two or three times, ultimate bengalensis (Oldham and Morris) Schimper, 1870 veins diverging straight to lateral and apical but Bose and Kasat (1972) transfered the type margins. specimens of that species to P. cutchense. Walkom's (191721, 1919(1) Ptilophyllum pecten specimens from Comments the Lower Cretaceous Stanwell and Burrum Coal Although Douglas' (1969) Ptilophyllllm boolensis Measures may also belong to P. clltclzense based on specimens were defined partly on cuticular pinnule morphology. features not available for the Western Australian Several Otozamites species identified by Walkom specimens, the Broome Sandstone frond (Figure (1921b) from the Yarragadee Formation, northern 7G) appears to be attributable to this species based Perth Basin, have much broader and either on its size and circular to elliptical pinnules. substantially longer or shorter pinnules than P. Otozamites pecten Sitholey, 1984 from the ?Jurassic cutchellse and are probably of Middle to Late of India is a similar diminutive frond with Jurassic age based on palynological studies of that rounded, partly overlapping, pinnules. unit (Backhouse 1988). Furthermore, its squat, sinuous-walled, papillate

~ Figure 7 A, damlrccl McCoy, 1874; WAM P.64.15; Broome Sandstone; x 2; B, Nilssonia sp. B; Base of frond; WAM 1'.885; Broome Sandstone; x 2; C,D, Ptilop!zyllllm clllc!zCIISC Morris (in Grant) emend. Bose and Kasat, 1972; (C) UWA16684, Leederville Formation, x 15.; (D) WAM P.88.172; Broome Sandstone; x 1; E, Zamitcs sp; isolated pinnule; UWA118826; Bullsbrook Formation; x 2; F, Nilssonia sp. A; fragmentary pmnules; WAM ['.96.3; Nanutarra Formation; x 1.5; G, Plilop!zyllllm boolcnsis (Douglas) Douglas, 1969; WAM P9613; Broome Sandstone; x 6; H, Arallcaria sp. B; WAM P.90.1; latex cast of axis with imbricate leaves; Broome Sandstone; x 5; I,J, Plzlopllllllllm aClIiifolilllll Morris (in Grant), 1840; (I) WAM P.96.2, apical portion of frond, Nanutarra Formation, x 2; (J) WAM P.89.162, Broome Sandstone, x 1, K silicon cast of AmllCilrta sp A, slender branches with elongate rhomboid leaves; WAM P.96.7; Broome Sandstone; x 3. 40 S. McLoughlin epidermal cells are closely comparable to the the Western Australian Cretaceous on the basis of epidermal!cuticular morphology of P. boolensis leaf size and shape and axis thickness. These suggesting a close relationship. Ptilophyllum species may represent different portions or stages boolensis is restricted to Douglas' (1969) "Zone B" of maturity of the same plant. Until intermediate flora in Victoria and its occurrence within the or more complete remains are found the foliage Broome Sandstone suggests that this unit is, at least types are described separately. partly, also of Neocomian age.

Genus Zamites Brongniart, 1828 Araucaria sp. A Figure 7K Type species Zamites gigas (Lindley and Hutton) Morris, 1843; 1993 Araucaria sp. A; McLoughlin and Guppy; Jurassic, Scarborough, England. Selected by figure 22. Andrews (1970). Material WAM P.63.31, WAM P.96.7. Zamites sp. Figure 7E Distribution Material Broome Sandstone (Neocomian-Barremian), UWA118826. Canning Basin.

Distribution Description Bullsbrook Formation, Perth Basin (Neocomian­ Slender terminal axes <4 mm wide, reaching 36 Barremian). mm long, bearing spirally inserted simple scale­ like leaves closely adpressed to axis (Figure 7K). Description Branches rare, irregular. Leaves small (1 mm wide, Linear pinnule 46 mm long, 5 mm wide, base 2 mm long), elliptical, ovate, or rhomboid, with contracted, apex rounded obtuse, margin entire prominent ridge along midline of abaxial surface (Figure 7E). Several veins emerge from base, rarely (on casts). Leaves tightly imbricate; overlapping by dichotomize immediately above base producing 10 about one-third those in distally adjacent spiral. ultimate veins which pass parallel along the lamina Leaf apices acutely pointed, commonly flexed and slightly converge near apex. Complete frond adaxially. and cuticular data not available. Comments Comments Araucaria sp. A differs from other Western The single available pinnule is similar to the Australian araucarians by its very small, linear parallel-veined leaflets of Zamites or longitudinally elongate, rhomboid leaves that Pseudoctenis fronds from the Early Cretaceous of typically show a ridge along the centre of the South Africa (Anderson and Anderson 1985) and abaxial leaf surface casts. In this respect the leaves India (Sahni and Rao 1933) but specific are very similar to Brachyphyllum sp. of Banerji and identification is not possible without complete Pal (1986) from the Upper Jurassic Sarnu Hill fronds or cuticular data. Formation, India. Araucarioid axes from the Yaragadee Formation (Middle to Late Jurassic), Division Pinophyta Perth Basin, described as Pagiophyllum or Brachyphyllum species or coniferous fragments by Class Pinopsida Arber (1910), Walkom (1921b), and White (1986) Order Pinales have narrower leaves that are not adpressed to the axis and belong to a separate araucarian species. Family Araucariaceae A number of other Australian and South Genus Araucaria Jussieu 1789 American forms, including Brachyphyllum gippslandicum McCoy (in Stirling), 1892, Araucaria Type species sp. of Drinnan and Chambers (1986), Brachyphyllum Araucaria araucana (Molina) K. Koch, 1873; extant; sp. of Baldoni (1980a), and Nothopehuen brevis Del Chile. Fueyo, 1991, are morphologically similar to Araucaria sp. A but their precise relationships will Discussion only be resolved when cuticular data and Three species of Araucaria are recognized from fructifications become available. Early Cretaceous macrofloras 41

Araucaria sp. B spirally arranged rhomboid leaves similar to those Figures 7H; 8A-C,F of Araucaria sp. B. However, closer comparison is not possible owing to the lack of cuticular details 1993 Araucaria sp. B; McLoughlin and Guppy; or pollenate cones associated with the Western figure 26. Australian material.

Material WAM P.89.188, WAM P.89.189, WAM P.89.190, WAM P.90.1, WAM P.90.3, WAM P.96.14. Araucaria sp. C Figure 8E

Distribution Material Broome Sandstone (Neocomian-Barremian), WAM P.88.16. Canning Basin. Distribution Description Broome Sandstone (Neocomian-Barremian), Axes slender to robust (2-9 mm wide, reaching Canning Basin. at least 70 mm long) bearing spirally arranged, broadly rhomboid, scale-like leaves (Figures 7H, Description 8F). Axes linear or variably curved, giving off Robust bifurcate axis clothed in spirally lateral branches at irregular intervals (Figures arranged?, flattened, lanceolate or linear leaves 8B,C). Leaves squat, remaining attached and (Figure 8E). Length from base to bifurcation 14 cm, adnate to axis, basally contiguous forming a terminal branches both 8 cm long. Maximum width rhomboid, pentagonal or hexagonal pattern on axis of axis 20 mm. Leaves reaching 4 mm wide, 30 mm impressions (Figures 8A,C). Leaves attached to axis long, smooth or with indistinct longitudinal striae, by broad base, apex contracted to a short «0.5 possibly contracted slightly at base, apices pointed mm) spine. Variations in the average basal width acute. Detached leaf imprints on same slab reach 5 of leaves occurs along some axes reflecting mm wide, 60 mm long. seasonal growth. Leaves most often apically compressed, apparently fleshy or leathery in original state, venation indistinct. When laterally Comments compressed, leaves slightly overlap bases of those The available specimen is too poorly preserved in succeeding spiral. for close comparison to other illustrated material. It probably represents an araucarian conifer branch bearing dense, flattened, spirally arranged, Comments lanceolate to linear leaves similar to the modem Numerous Australian Mesozoic axes bearing Australian Bunya Pine (Araucaria bidwilli Hooker, rhomboid leaves, and often assigned to 1843). The leaves and branched axis of Araucaria BraclzyplzyIlum or AIlocladus species (e.g., Seward sp. C are much more robust than the other species 1904; Chapman 1904; Walkom 1917b, 1919a, 1921a, of Araucaria described from the Broome Sandstone. 1928; Medwell 1954a; Hill et al. 1966; Townrow 1967; White 1961a, 1981), may belong to the Araucariaceae, Podocarpaceae, Taxodiaceae, Cupressaceae, or Cheirolepidaceae. In this Araucarian cones instance, association with araucarian cone scales Figure 80 favours an araucarian affinity, however, further morphological details and attached fertile material Material will be needed to clearly establish the affinities of WAM P.89.185, WAM P.89.187. Araucaria sp. B. BrachyphyIlum rholllbicum Feistmantel of Sahni Distribution (1928) is closely comparable to Araucaria sp. B Broome Sandstone (Neocomian-Barremian), whereas his B. mamillare Brongniart has more Canning Basin. elongate rhomboid leaves. The Early Cretaceous Indian BracJlyphyllrwl bansaensis and B. eikaiostomum Description of Sukh-Oev and Bose (1974) are differentiated on Two cone impressions available, both c. 5 x 4 cm, epidermal/cuticular details but are otherwise longitudinally elliptical in outline (Figure 80). difficult to distinguish from Araucaria sp. B. Baksi's Cones feature >15 transverse ridges (on casts) (1968) Bracllyphyllum rllOmbiculIl (Lindley and marking positions of closely spaced cone scale Hutton) Sahni is also very similar but its leaves are spirals (ridges 1.5 mm apart). Individual cone more axially elongate. The South African Araucaria scales poorly differentiated, apices extended as 6 rogersii (Seward) Anderson and Anderson, 1985 has mm long flanges or spines that arch distally at 42 s. McLoughlin Early Cretaceous macrofloras 43

periphery of cone. Cone attachment point not most South American and eastern Australian cone evident. Seeds and foliage not available. scales (Walkom 1919a; Baldoni 1981; Drinnan and Chambers 1986). They also appear to have poorly Comments developed lateral membraneous wings although Both of the Broome Sandstone cones are isolated this may be a preservational flaw. The Broome and can not be linked to any of the araucarian cone scales are most similar to Araucarites sp. B of foliage species described above. Latex casts failed Seward (1904) and the "winged seed type 1" of to reveal significant morphological details of cone Drinnan and Chambers (1986). Indian Mesozoic scale shapes although their large size suggests that cone scales assigned to Araucarites parsorensis Lele, the cones are ovulate rather than pollenate. 1955 and Baksi's (1968) A. cutchensis Feistmantel Anatomical details are not preserved in the are not as apically expanded as the Western Western Australian specimens preventing Australian specimens. Araucarites janaianus Bose comparison to permineralized araucarian cones and Banerji, 1984 is similar in size and ovule from the Jurassic? of India (Bohra and Sharma position to the Western Australian scales but it 1980). An araucarian cone from the Kennedy possesses distinct apical spines. Ranges (Merlinleigh Sandstone: Eocene), Western Australia, figured by McLoughlin and Hill (in press) is of similar dimensions but has more Family Podocarpaceae distinctive cone scale impressions which in some Genus Elatocladus Halle emend Harris, 1979 cases have left prominent spirally arranged rhomboid scars on the cone axis. Type species Elatocladus heterophylla Halle 1913; ?Lower Jurassic or ?Lower Cretaceous; Hope Bay, Araucarian cone scales Antarctic Peninsula. Figures 9A,B 1993 Araucarian cone scale; McLoughlin and Elatocladus ginginensis sp. novo Guppy; figure 20. Figures 9C-E,G Material 1961 Elatocladus plana (Feistmantel); White; p. 302­ WAM P.96.5, WAM P.96.6. 303; plate 6, figures lA, 2A. [1961a). 1993 Rissikia sp.; McLoughlin and Guppy; p. 14; Distribution figure 24. Broome Sandstone (Neocomian-Barremian), Canning Basin. Holotype UWA16703; Leederville Formation (Neocomian­ Description Barremian); Cheriton Creek, Gingin, Western Detached cone scale complex (fused ovuliferous Australia. scale, seed, and bract). Scale cuneate, base pointed acute to truncate, apex truncate, no terminal spine Material evident (though possibly removed); width 13 mm, WAM P.88.14, WAM P.89.174, WAM P.89.192, length 12-14 mm. Scales with elliptical to oblong WAM P.96.1O, UWA10464C-E, UWA10466B-E, indentation or cast (2.5 mm wide, 8 mm long) near UWA10467A,B, UWA10468A, UWA10474B, base marking position of seed (Figures 9A,B). Cone UWA10475, UWA10477B, UWA16703. scale narrowly winged. Distribution Comments Broome Sandstone, Canning Basin; Leederville Two cone scales are preserved in the Broome and Bullsbrook Formations, Perth Basin (all Sandstone assemblages. Similar isolated cone scales Neocomian-Barremian). are found throughout Gondwanan Jurassic and Cretaceous strata but their affinities with foliage­ Diagnosis based species are generally indeterminate. The Slender distal shoots bearing univeined, spirally Broome specimens lack apical spines in contrast to inserted, basally twisted, linear leaves in distichous

arrangement. Leaves flattened; apices bluntly not allow close comparison. Other Australian rounded; bases slightly contracted to slightly specimens assigned to E. plana (Hill et al. 1966; decurrent. Walkom 1919a); and Rissikia talbragarensis White, 1981 have either substantially longer or broader Description leaves which show a dense but regular spacing not Slender terminal axes (reaching 3 mm wide and evident in E. ginginensis. 51 mm long) bearing numerous univeined linear Elatocladus conferta (Oldham and Morris) Sahni, leaves (Figures 9C,E,G). Leaves spirally inserted 1928 and E. kingianus Bose, Kutty, and although basally twisted to lie in a common plane. Maheshwari, 1982 specimens of Halle (1913), Sah Leaves inserted on branches at 40°-90°. Leaves up (1965), Maheshwari and Singh (1976), Bose and to 1 mm wide, 20 mm long, flattened, straight or Banerji (1984), and Walkom (1928) have slightly flexed distally, apices rounded. Leaf bases substantially smaller leaves than E. ginginensis. slightly contracted but not significantly petiolate, Permineralized remains assigned to E. salmii slightly decurrent (Figure 90). Vishnu-Mittre, 1957 from the Indian Jurassic have similar decurrent bases but the absence of anatomical details in the Western Australian Comments specimens prevents closer comparison. Extra­ Harris 1979 emended Elatocladus Halle, 1913 to Gondwanan Elatocladus species are more readily exclude the morphology of the leaf bases as a distinguishable on gross leaf morphology or diagnostic character thus incorporating specimens cuticular details (e.g., Miller and Lapaska 1985; previously assigned to both Elatocladus and Rissikia Kimura and Ohana 1988, 1989; Kimura et al. 1991). Townrow, 1967. It should be noted, however, that some workers (e.g., Anderson and Anderson, 1989) have maintained a distinction between these genera based on the presence of contracted leaf bases in Incertae sedis Elatocladus and decurrent leaf bases in Rissikia. In Genus Carpolithes Brongniart 1822 the absence of attached reproductive organs the Western Australian material is insufficiently Type species abundant or well preserved to resolve this Carpolithes thalictroides Brongniart, 1822; Tertiary; taxonomic issue and the specimens are assigned to France and England. a new species within Harris' (1979) broader definition of Elatocladus. Numerous species from the Gondwanan Mesozoic attributed to Elatocladus, Carpolithes bullsbrookensis sp. novo Rissikia, or other podocarpacean genera have often Figures 9F,H-K been distinguished by slight differences in leaflet size, shape, leaf arrangement, and petiole 1993 Carpolithes sp. McLoughlin and Guppy; figure characters (Petriella and Arrondo 1984). 19. Elatocladus ginginensis sp. novo lacks cuticle but is Holotype distinguished from other species by the size of its UWAI0469A; Bullsbrook Formation slender leaves, rounded leaf apices, and decurrent (Neocomian-Barremian); 2.5 km northeast of lamina bases which are not as significantly Bullsbrook Hotel, Perth Basin. contracted as those of Elatocladus longifolium Baldoni, 1980b, E. plana (Feistmantel) Seward 1919, Material or Morenoa fertilis Del Fueyo, Archangelsky, and Taylor 1990. White's (1961a) Elatocladus plana UWAI0374C,D, UWA10462B, UWAI0464H,I, UWA10468B, UWAI0469A,B-F, UWA10473A. specimens from the Cronin Sandstone have identical le"lflet characters and are reassigned to E. gillgillelh,IS. TILe E. plana specimens of Glaessner Distribution and Rao (1955) from the Early Cretaceous of South Bullsbrook Formation (Neocomian-Barremian); Australia are described as having leaves with full Perth Basin. basal attachment, a feature not consistent with r:latocladus. Their specimens may be synonymous Diagnosis with E. ginginensis although their illustrations do Ovate to elliptical wingless seeds, reaching 5 x 7

... Figure 9 A,B, Araucarian cone scales; (A) WAM P.96.5; (B) WAM P.96.6; Broome Sandstone; both x 4; C-E, G, Elatocladlls gillgilli'llsis sp. nov.; detached planated foliage-bearing twigs; (C) UWA16703, x 1.5; (D) UWA118835, x 5; (E) UWA10466C, x 2; (G) UWA10464E, x 2; all from Bullsbrook Formation; F, H-K, Carpoltthi's bllllsbrookCllsis sp. nov.; seed impressions and internal casts; (E) UWA10469F; (H) UWA104688; (I) UWA10462B; (J) UWA10469C; (K) UWA10469A; all from Bullsbrook Formation; all x 8. 46 s. McLoughlin mm, with a rounded or obtusely pointed apex and Description smooth surface apart from a prominent vertical Seed positioned basally on wing, wrinkled, ridge (on internal casts). ovate, c. 3.5 mm wide, 5 mm long. A flange «1 mm wide) around lateral margins of seed expands Description distally to form the 8.5 x 17 mm longitudinally Isolated seeds of indeterminate affinity. Seeds striate oblong wing (Figure lOA). ovate to elliptical, reaching 5 mm wide, 7 mm long, wings absent; internal casts ovoid (Figures 9H-K). Comments Prominent vertical ridge commonly evident on The single available seed impression differs from internal casts but surface otherwise lacks those figured by Drinnan and Chambers (1986) ornamentation (Figures 9F,J,K). Base truncate or from the Victorian Cretaceous by its distally rounded, apex rounded or closing to an obtuse expanded wing. The basal part of the seed has point. broken away leaving only a small portion of the fertile body. The seed does not show organic Comments connections to any other plant remains and its Seeds of this type are common in the Bullsbrook affinities remain uncertain. Formation and are frequently preserved in association with Taeniopteris daintreei leaves although no physical connection is evident. White Genus Bucklandia Presl (in Sternberg), 1825 (1981, 1986) figured similar though smaller seeds Type species with a longitudinal ridge preserved in Bucklandia anomala (Stokes and Webb) Presl (in pentoxylalean (Carnoconites) cones from the Sternberg), 1825; ]urassic; Sussex, England. ]urassic Talbragar fish beds in New South Wales. Seeds belonging to Carnoconites cranwellii Harris, 1962 fruits from the Victorian Early Cretaceous (Drinnan and Chambers 1985) are also Bucklandia sp. substantially smaller (1 mm diameter) than the Figure 5E Western Australian seeds and differ by their 1961 Bucklandia sp.; White; p. 305; plate 8, figure 2. polygonal outline. This may simply be due to [1961a]. crowding of seeds in the Victorian cones. Larger, more loosely packed Carnoconites cones may have 1993 Bucklandia-type axis; McLoughlin and Guppy; borne more rounded seeds like White's (1981) p. 14; figure 9 (upper part). examples from Talbragar. Seeds of Carnoconites Material compactum Srivastava, 1946 are similar in size WAM P.96.8. although the species was defined upon permineralized material which does not clearly Distribution show the seed shape in lateral view (Sahni 1948; Broome Sandstone (Neocomian-Barremian), Vishnu-Mittre 1953). Compressed Carnoconites Canning Basin. laxum Harris, 1962 fruits from New Zealand show only indistinct seeds not favourable for comparison to Carpolithes bullsbrookensis. Carpolithes sp. B of Description Bose and Banerji (1984) has a longitudinal ridge Axis impression bearing spirally arranged, like C. bullsbrookensis but is much larger with an broadly elliptical, leaf scars. Leaf scars 4 mm wide, acute apex. Seeds belonging to Rissikia 2 mm tall, lacking obvious vascular strands. Scars talbragarensis White, 1981 are of similar size to C. raised above surface of axis (on cast), separated by bullsbrookensis but lack a longitudinal ridge. 0.5 mm grooves; arranged in 35° right-handed spirals, 50° left-handed spirals measured with respect to axial direction. Leaves and anatomical Winged conifer seed details not preserved. Figure lOA 1993 Conifer seed with wing; McLoughlin and Comments Guppy; figure 25. The single stem fragment (upper right of Figure 5E) is similar to a number of ]urassic and Material Cretaceous axis casts assigned to Bucklandia Presl WAM P.89.191. (in Sternberg), 1825. White (1961a) figured a larger axis from the same locality (Gantheaume Point, Distribution Broome) showing well developed spirally arranged Broome Sandstone (Neocomian-Barremian), elliptical leaf scars. The Western Australian Canning Basin. examples lack the anatomical details necessary for Early Cretaceous macrofloras 47 comparison to Indian Bucklandia species described Distribution by Seward (1917), Sahni (1932), Bose (1953a, b) and Broome Sandstone (Neocomian-Barremian), Sharrna (1969a). Anatomically preserved Bucklandia Canning Basin. axes have been associated with Williamsonia fructifications and Ptiloplzyllum-type foliage in Description India (Sahni 1932) suggesting a bennettitalean Irregular ?bifurcate axis bearing circular to affinity. Although a Roebuckia spatulata frond elliptical protuberances set in surface depressions occurs on the same slab as the Bucklandia specimen (on mould) separated by broad ridges and folds (Figure 5E), no connection is evident or inferred. (Figure 1OC). Protuberances reaching 3 mm wide, 7 mm long, roughly arranged along axis in en eclzelon bands. Sporangia, leaves, and cuticle not evident. Ribbed axes Figure lOB Comments 1993 Striate axis; McLoughlin and Guppy; figure A single available specimen is similar to axes 23. figured by Seward (1904) and Chapman (1909) from the Lower Cretaceous of Victoria. Chapman Material (1909) inferred a relationship between his WAM P.63.30, WAM P.65.38, WAM P.65.43, Rlzizomopteris axes and Taeniopteris leaves. No such WAM P.65.46, WAM P.65.47, WAM P.65.48, WAM affinity can be deduced from the present material. P.66.3, WAM P.66.4, WAM P.66.6, WAM P.66.7, The fossil probably represents a fern rhizome WAM P.66.8, WAM P.66.9, WAM P.66.1O, WAM bearing circular to elliptical scars of abscissed P.66.12. frond rachises although no vascular traces are evident on the abscission zones. Rlzizomopteris Distribution rajmalzalensis Gupta, 1955 is most closely Broome Sandstone, Canning Basin; Bullsbrook comparable to the Broome Sandstone specimen but Formation, Perth Basin (both Neocomian­ has C-shaped rather than circular to elliptical Barremian). rachis scars. Bose (1958) figured additional material comparable to Gupta's (1955) specimens Description and considered that the axes belonged to conifers. Several indeterminate longitudinally ribbed or Coniferous axes figured by Bancroft (1913) and striate axes are preserved as casts (figure lOB) and Sahni (1931) have transversely elongate abscission impressions. Axes reaching 4.5 cm wide, 13.5 cm scars in contrast to the oblique or longitudinally long, cylindrical or flattened through compaction, elongate scars of Gupta's (1955), Bose's (1958), or commonly with external? longitudinal ribs the Broome Sandstone Rlzizomopteris species. averaging 2.5 mm apart. No axis nodes, attached leaves, fruits, or roots evident. Genus Mesembrioxylon Seward, 1919 Comments These axes lack diagnostic features necessary to Type species assign them to any plant group, however, their size Mesembrioxylon woburnense (Stopes) Seward, 1919; suggests that they may be the remains of a robust Cretaceous; Wobum, England. gymnosperm. The ribbed surface of some specimens invites comparison to equisetalean axes (e.g., Roy 1968; Drinnan and Chambers 1986), Mesembrioxylon sp. however, the Western Australian axes lack the Figures lOD-H transverse nodes typical of that group. 1993 Silicified wood with Teredo burrow casts; McLoughlin and Guppy; figure 1. Genus Rhizomopteris Schimper, 1869 1994 Fossil wood; McLoughlin et al.; p. 453; figs 8a-m, lOa,b. Type species Rlzizomopteris lycopodioides Schimper, 1869; Material Carboniferous; near Dresden, Germany. UWA120186-UWA120237.

Rhizomopteris sp. Distribution Figure 10C The described specimens are derived from the Birdrong Sandstone (Neocomian-?Aptian), Material Camarvon Basin. Other fossil woods have been WAM P.89.184. reported from the Nanutarra Formation 48 S. McLoughlin Early Cretaceous macrofloras 49

(Neocomian-?Aptian) and Yarraloola Conglomer­ groups is abundant in coeval Western Australian ate (Neocomian-?Barremian) of the Carnarvon strata. Identification of fossil conifer woods is Basin, Dandaragan Sandstone (?Barremian­ largely based on pitting arrangements on the radial ?Aptian) and Molecap Greensand (Cenomanian­ walls of longitudinal xylem tracheids and ray cells Santonian) of the Perth Basin, and Nakina but due to poor preservation such details are not Formation (Barremian-Iowermost Aptian) of the available for most Western Australian specimens. Collie Basin (Simpson 1912; Cox 1961; Playford et The absence of typically araucariacean closely al. 1976; Hocking 1990, Buckhouse et al. 1995). spaced multiseriate hexagonal bordered pits on radial tracheid walls and sporadic presence of Description uniseriate bordered pits with oblique apertures Woods typically fragmentary, seldom exceeding suggests that these woods have podocarpacean 50 cm long and 15 cm diameter, randomly affinities although cupressacean woods also have orientated in sediments. Woods show extensive similar pitting arrangements (Greguss 1955). The biogenic borings (Figures lOE,H). Woods preserved prominent but variable growth banding and by siliceous and phosphatic permineralization, occasional development of false growth rings strongly weathered and infiltrated by iron-oxide implies a distinctly seasonal climate but with and day minerals. Only secondary xylem tissues considerable intra- and inter-seasonal variability are preserved. Prominent growth banding evident McLoughlin et al. (1994). (Figures lOG,H) but details of pitting arrangement Cox (1961) illustrated infilled molluscan borings on individual cell walls often poorly defined or from the Nanutarra Formation (Early Cretaceous, absent (Figure 100). Relatively sparse uniseriate Carnarvon Basin) both with and without traces of xylem rays consist of 1-12 (av. 4) cells (Figure lOF). associated fossil wood. The casts are probably Uniseriate or rarely biseriate bordered pits are remnants of teredinid or pholadid boring casts left sparsely evident on radial walls of xylem tracheids. behind after the degradation of coniferous Tangential walls not pitted. Pits have oblique slit­ driftwood. One wood specimen in the Birdrong like apertures where discernible. Growth bands Sandstone assemblage shows a cross-section of an variable in width (0.06-12 mm) both between and unidentified bivalve preserved within an within specimens. False growth rings sporadically excavation chamber (Figure lOE). developed in some specimens. In most cases middle lamella (initially consisting of pectinoid Saprophytic fungi compounds between the primary walls of adjacent Figures 11A-I cells) has been degraded producing a wood texture that appears to consist of detached or partially 1994 Fungi within wood; McLoughlin et al.; p. 455; detached tracheids. Cambium, phloem, and bark figures 9a-1. tissues absent. Primary xylem and pith poorly preserved or masked by iron oxides. Material UWA120187, UWA120217, UWA120228, UWA120229, UWA120231, UWA120232. Comments The small pith content of the plants (pycnoxylic axes) suggests that the woods derive from the Distribution dominant arborescent elements of the Australian Birdrong Sandstone (Neocomian-?Aptian), Early Cretaceous vegetation (viz., the podocarp or Carnarvon Basin. araucarian conifers) rather than fern, pteridosperm, or bennettitalean groups that typically contain a Description large proportion of pith within the stem (manoxylic Several fossil wood specimens from the Birdrong axes). Permineralized wood belonging to podocarp Sandstone, Carnarvon Basin, show zones of partly and araucarian conifers is common in Gondwanan degraded cells or cavities which are circular, Cretaceous sediments (Sahni 1931; Jefferson 1983; elliptical, or irregular in transverse section and Francis 1986; Frakes and Francis 1990; Francis and elliptical in longitudinal section. The cavities are Coffin 1992) and the foliage attributable to these up to 4 mm in axial length and 0.2-0.9 mm in

~ Figure 10 A, Winged seed with portions of base and apex removed; WAM P.89.191; Broome Sandstone; x 6; B, Cast of ribbed axis; WAM P.63.30; Broome Sandstone; x 1; C, Rhizomopteris sp.; WAM P.89.184; Broome Sandstone; x 2; D,F,G, Mesembrioxylon sp.; (D) radial section with three groups of ray cells, UWA120l87, x 60; (F) tangential section showing rays generally comprising 2-4 cells, UWA120199, x 30; (G) transverse section showing transition from latewood (bottom) to earlywood (top), UWA120186, x 30; all from Birdrong Sandstone; E,H, Transverse sections of Mesenzbrioxylol1 wood showing variation in growth rings and intensive bivalve borings filled by calcite and/or glauconitic sand. Note preserved bivalve within bored cavity (E: arrowed); (E) UWA120220, x 2; (H) UWA120208, x 1.5; both from Birdrong Sandstone. 50 S. McLoughlin Early Cretaceous macrofloras 51

diameter. The degraded ZOnes occur commonly, from the Callawa formation, is a pinna bearing though not exclusively, within the latewood oblanceolate pinnules some of which appear to (Figures lIA,B,C). The cavities are either filled have elliptical fertile bodies along their acroscopic with secondary iron oxides or silica or occasionally margins. Similar fronds from the Victorian and contain ramifying weakly septate filaments Indian Early Cretaceous have been assigned to reaching 450 pm in length and 4-15 pm in width Onychiopsis paradoxus Bose and Sukh Dev, 1961 (Figure lID). The filaments branch irregularly, (Drinnan and Chambers 1986). White's (1961a) have an external bullate texture, and appear to Cladophlebis albertsi (Dunker) from the Leveque contain irregularly spaced and poorly defined Sandstone [now regarded as part of the Broome septa (Figures lIE-G)). Filaments often occur in Sandstone (Yeates et al. 1984)] differs from intimate contact with the degraded wood cells of Cladophlebis sp. cf. C. oblonga by its pointed the cavity margin (Figure lIF). pinnules but her specimen is too poorly preserved for close comparison to other described species. White's (196lc~.) Dichopteris delicatula Seward from Comments the Broome Sandstone does not have preserved Such filaments and spindle-shaped wood cavities venation and may be a Sphenopteris or Cladophlebis are morphologically similar to saprophytic fungal species. Her Nilssonia schaumbergensis Dunker, also hyphae and white pocket rot cavities described from the Broome Sandstone has pinnules of from other modern and fossil woods (Blanchette variable width. The specimen may be a Taeniopteris 1980; Stubblefield et al. 1985; Stubblefield and daintreei leaf showing irregular lamina dissection Taylor 1986, 1988; Weaver et al., in press). Jefferson like some forms illustrated herein. White's (1987) figured fungal hyphae and possible spores Cycadolepis sp. from the Cronin Sandstone is too in Early Cretaceous conifer wood from Alexander poorly preserved for definite identification. White Island, Antarctica, and Halle's (1913) sketch of (1961a) also mentions but does not figure several Mesozoic fossil wood from the Antarctic Peninsula other plant taxa from the Canning and Officer also shows elliptical markings reminiscent of Basin Lower Cretaceous including: Carpolithus pocket rot. The Birdrong Sandstone woods circularis, Cladophlebis sp., Taeniopteris howardensis incorporate the first examples of fossilized Walkom, Dictyophyllum davidi Walkom, saprophytic fungi recorded from Western Australia Pterophyllum sp., Sphenopteris superba Shirley, (McLoughlin et al. 1994). Brachyphyllum mamillare Lindley and Hutton, and Ginkgoites digitata Brongniart. These identifications Other plant remains (Table 1) have not been assessed in the present White (1961a) figured a number of Western study. Australian plant macrofossils many of which have Walkom (in McWhae et al. 1958) identified been reassigned here to different taxa on the basis specimens referrable to Nilssonia, Otozamites, and of information from new material in the Western Elatocladus from the Nanutarra Formation. The Australian Museum and University of Western whereabouts of this material is unknown and nO Australia collections (Table 1). However, a few of revision of the identifications can be attempted. her figured specimens are not matched by However, these genera are common elements of conspecific material in the Western Australian the Early Cretaceous Broome Sandstone flora and collections. Her Pagiophyllum peregrinum Schimper it is likely that several species are shared between from the Callawa Formation, although similar to these formations. Walkom (in McWhae et al. 1958) Araucaria sp. B, has leaves with distinctly also reported fossil leaves from the Yarraloola mucronate apices. White's (1961a, plate 6, figure 4) Conglomerate (Early Cretaceous, Carnarvon Basin) "fine frond" is too incomplete for confident but provided nO identifications and the location of identification but it may represent the fragmentary his specimens is unknown prohibiting systematic remains of an Aculea or Sphenopteris frond or an re-evaluation. Elatocladus terminal twig. Her Pachypteris sp., also Brunnschweiler (in Traves et al. 1956) tentatively

.... Figure 11 Aspects of fungal attack on, and hyphae preservation within, Mesembrioxylon (conifer) wood from the Birdrong Sandstone. (A) UWA120231, transverse section of wood showing fungal-degraded pockets, x 2. (B) UWA120187, transverse section of wood showing fungal-degraded pockets positioned within latewood, x 6; (C) UWA120221, transverse section of wood showing initial stages of cell breakdown and pocket formation, x 12; (D) UWA120186, details of a pocket with numerous phosphate coated fungal hyphae and a spherical body (upper left), x 60; (E) UWA120217, branched hyphae within a pocket, x 60; (F) UWA120232, details of fungal hyphae represented by a thin central filament surrounded by a bullate­ textured phosphate coating, x 120; (G) UWA120186, details of an expanded area near the tip of a fungal hypha, x 600; (H) UWA120132, fungal hyphae in contact with wood tracheids along the margin of a pocket, x 120; (I) UWA120232, branched fungal hyphae in pocket, x 120. ------_ .. _------

52 S. McLoughlin

Table 1 Tentative revision of Lower Cretaceous plants illustrated or identified by White (1961a) from Canning and Officer Basins collections compiled during regional geological surveys of the 1950s and 1960s.

White's (1961a) identification! illustration Formation Tentative revised identification

Stenopteris tripinnata Walk. (PI. 4, fig. 6) Cronin Sst cf. Aculea bifida Linguifolium denmeadi Jones and de Jersey Cronin Sst Phyllopteroides westralensis (PI. 5, fig. 3) 'Neorhachopteris minuta' (PI. 5, fig. 4) Callawa Fm Sphenopteris sp. A Pagiophyllum peregnnum Sch. (PI. 5, fig. 5) Callawa Fm ?Araucaria sp. Ruffordia mortoni Walk. (PI. 5, figs 6,7) Callawa Fm cf. Sphenopteris sp. A Elatocladus planus (Feist.) (PI. 6, figs lA, 2A) Cronin Sst Elatocladus ginginensis Taeniopteris cf. elongata Walk. (PI. 6, Cronin Sst Taeniopteris daintreei figs lB,2B) Ptilophyllum pecten (Phillips) (PI. 6, fig. ID; Cronin Sst, Ptilophyllum cutchense PI. 8, fig. 1) Broome Sst Cycadolepis sp. (PI. 6, fig. 2e) Cronin Sst not identified stem (PI. 6, fig. 3) Callawa Fm NathorstianeIla babbagensis fine frond (PI. 6, fig. 4) Callawa Fm not identified Pachypteris sp. (PI. 6, fig. 5) Callawa Fm Onychiopsis sp. Cladophlebis australis (Morr.) (PI. 7, figs 1,6) Callawa Fm, Cladophlebis cf. oblonga Broome Set Cladophlebis albertsi (Dunk.) (PI. 7, fig. 2) Broome Sst ?Cladophlebis sp. Dichopteris delicatula Sew. (PI. 7, fig. 3) Broome Sst ?Sphenopteris or Cladophlebis sp. Nilssonia schaumbergensis Dunk. (PI. 7, fig. 4) Broome Sst Nilssonia or Taeniopteris sp. Otozamites bengalensis O. and M. (PI. 7, fig. 5) Broome Sst PtilophyIlum cutchense Bucklandia sp. (PI. 8, fig. 2) Broome Sst Bucklandia sp. Zamites sp. (PI. 8, fig. 3) Broome Sst PtilophyIlum acutifolium Hausmannia cf. buchU Andrae (PI. 8, fig. 4) Broome Sst Hausmannia sp. Taeniopteris sp. Cronin Sst not assessed Carpolithus circularis Cronin Sst not assessed . Hausmannia sp. Cronin Sst not assessed Cladophlebis sp. Broome Sst not assessed Taeniopteris howardensis Walk. Broome Sst not assessed Dictyophyllum davidi Walk. Broome Sst not assessed Pterophyllum sp. Broome Sst not assessed Sphenopteris superba Shirley Broome Sst not assessed Brachyphyllum mamillare 1. and H. Broome Sst not assessed Ginkgoites digitata Brongn. Callawa Fm not assessed Brachyphyllum foliage Callawa Fm not assessed

identified ?Johnstonia, Ginkgo, Cladophlebis, late Mesozoic (Bose and Jana 1979; Bose and Pagiophyllum, ?Brachyphyllum, Taeniopteris, Banerji 1984). Specimens referred to Otozamites, Sphenopteris, ?Samaropsis, Dictyophyllum and Pterophyllum and Pseudocycas may belong to one of indeterminate wood from the Callawa Formation, the Ptilophyllum species described above. Canning Basin. The presence of Johnstonia (=Dicroidium) would denote a Triassic age for this Comparisons between the Western Australian unit but the identification is probably incorrect. assemblages Brunnschweiler (1960) listed but did not describe A significant number of plant taxa are shared or illustrate the following taxa from the Broome between the various Western Australian Early Sandstone at Gantheaume Point: Ptilophyllum. Cretaceous formations (Table 2). Although there is pecten, Cladophlebis .australis, Otozamites cf. evidence of intra-Australian floral provincialism in bengalensis, Pseudocycas sp., Hausmannia sp., the Early Cretaceous (Dettmann et al. 1992), the ?Cordaites sp., Pterophyllum sp., ?Microphyllopteris Western Australian marginal basins all lay within sp., Sphenopteris cf. superba Walkom, ?Dictyophyllum latitudes of about 45°-55° during that period sp., Cladophlebis cf. albertsi, Taeniopteris cf. (Veevers et al. 1991) and probably experienced howardensis, and Nilssonia sp. Most of these taxa similar climatic conditions. Minor differences are probably attributable to species described between the Western Australian assemblages are herein. The identification of Cordaites, a Palaeozoic probably attributable to limited sampling, local genus, is almost certainly incorrect. The depositional factors and environmental setting. For Dictyophyllum specimens may represent example, the fully fluvial Cronin Sandstone fragmentary portions of Hausmannia fronds appears to contain a higher representation of although the former is not uncommon in the Indian gymnosperms than units deposited in deltaic or Early Cretaceous macrofloras 53

Table 2 Distribution of fossil plant taxa in the studied stratigraphic units. See text for author citations.

~z ~Z ~Z ~ ~ ,..JO 00 p::;0 l:lZ Z ~~ ~~ <:>-< ZO ~O ~<: 1:0<: 0<:...... """ 0"""p::;tfl :::E~ 0:::E ~:::E Z~ 00 00 ~p::; ,..Jp::; p::;Z oZ ~O 00 >-<<: P::;<: SPECIES ,..J>z.. 1:O>z.. ~~ I:Otfl I:Otfl

Isoetites elegans + Nathorstianella babbagensis + Hausmannia sp. + Cladophlebis cf. oblonga + + Phyllopteroides westralensis + + Microphyllopteris gleichenioides + Roebuckia spatulata + Aculea bifida + Sphenopteris warragulensis + Sphenopteris sp. A + Thinnfeldia cf. talbragarensis + + Taenipteris daintreei + + + Pinnate frond + Nilssonia sp. A + Nilssonia sp. B + Ptilophyllum acutifolium + + Ptilophyllum cutchense + + + Ptilophyllum boolensis + Zamites sp. + Araucaria sp. A + Araucaria sp. B + Araucaria sp. C + Araucarian cones + Araucarian cone scales + Elatocladus ginginensis + + + Carpolithes bullbrookensis + Winged conifer seeds + Bucklandia sp. + Ribbed axes + + Rhizomopteris sp. + Mesembrioxylon sp. + Saprophytic fungi + coastal plain environments. The latter units are Formation) flora of Western Australia (Arber 1910; richer in lycophytes and pteridophytes probably Walkom 1921b). Of these, the Otway, Gippsland, owing to milder local maritime climates and wetter and Eromanga Basin floras are the best known. riparian environments, or higher water tables in Douglas (1969, 1971) proposed four assemblage lowland settings. zones (Zones A-D) for Victorian Neocomian to Early Albian strata based on plant macrofossils and Relationships with other Australian floras tied these to the independent palynostratigraphic Other significant Australian mid- to late zonation of Dettmann (1963); later refined by Mesozoic floras include the Gippsland-Otway Dettmann and Playford (1969), Dettmann and Basin assemblages of Victoria (Douglas 1969, 1973; Douglas (1976), and Helby et al. (1987). Douglas' Drinnan and Chambers 1986), the Talbragar flora (1969) megafloral zones are largely based on the of New South Wales (Walkom 1921a; White 1981), presence or absence of key foliage species hence the Eromanga, Clarence-Moreton, Laura, Styx, and their recognition may be strongly influenced by Maryborough Basin floras of Queensland (Walkom local palaeoenvironmental, depositionai, and 1917a,b, 1918, 1919a, 1928; Hill et al. 1966; Gould seasonal factors. The collective Western Australian 1978, 1980; McLoughlin and Drinnan 1995; assemblages bear greatest similarity to Douglas' McLoughlin et al. 1995), the Bauhinia Downs flora (1969) Zone B (Neocomian) flora due to the of the Northern Territory (White 1961b), the Mt presence of the key species Taeniopteris daintreei Babbage flora of South Australia (Glaessner and and Ptilophyllum boolensis. Phyllopteroides Rao 1955), and the Mingenew (Yarragadee westralensis is an entire-margined form similar to P. 54 s. McLoughlin laevis from the Victorian Zone B (and equivalent Sandstone, Laura Basin, initially assigned to the strata in Queensland). This favours an Early Cretaceous by Walkom (1928) are now regarded as Cretaceous age but P. westralensis requires further Jurassic in age (Gould 1975) and only share one study to determine its phylogenetic relationships species (Microphyllopteris gleichenioides) with the with the range of eastern Australian species. Western Australian assemblages. The Middle to Another diagnostic Victorian Zone B species, Late Jurassic floras of the Yarragadee Formation, Pachypteris austropapillosa is absent from the Perth Basin, Western Australia (Arber 1910; Western Australian assemblages although a poorly Walkom 1921a; McLoughlin and Hill, in press) preserved pinnate frond from the Bullsbrook contain Otozamites fronds and Pagiophyllum Formation (Figure 6D) may be a related form. branchlets that differ from the Early Cretaceous Douglas's (1969) latest Jurassic or earliest fossils. Several other fragmentary plant fossils have Cretaceous Zone A floras are characterized by been recorded from the Yarragadee assemblages bennettitalean leaf forms not recognized in but the flora requires systematic revision. Western Australia. Victorian Zone B floras are also A small plant assemblage of probable Late rich in bennettitaleans but only one Western Jurassic age from the Hooray Sandstone in the Australian form (Ptilophyllum boolensis) is shared northwestern Eromanga Basin, Northern Territory suggesting significant intra-Australian floristic (Gould 1978), is dominated by Bennettitalean provincialism. Nevertheless, the abundance of fronds (Ptilophyllum sp.) and conifer twigs but no bennettitaleans in Zone B suggests better species are shared with the Western Australian correlation with the Western Australian floras than Cretaceous floras. does the succeeding Zone C where bennettitaleans The Maryborough Formation of the are entirely absent. The Victorian Zone C (?late Maryborough Basin, Queensland, contains a small Neocomian to early Albian) floras also differ from macroflora showing some similar elements the Western Australian assemblages by the [Taeniopteris spp., Ptilophyllum pecten (= presence of ginkgophytes, sphenophytes, ?Ptilophyllum cutchense)] to the Western Australian angiosperms, and a range of distinctive assemblages (Walkom 1918). However, the pteridophyte and conifer species (Drinnan and Maryborough Formation floras are more similar to Chambers 1986). A few species (Taeniopteris the Gippsland Basin Zone C (?late Neocomian to daintreei, Sphenopteris warragulensis, and Aculea earliest Aptian) macrofloras in terms of the conifer, bifida) are shared between these assemblages but ginkgophyte and equisetalean groups represented. some of these forms may be long-ranging. Douglas' The Albian Burrum Coal Measure flora from the (1969) Zone D (middle-?late Albian) floras are Maryborough Basin (Walkom 1919a) has a number dominated by conifer and fern species which are of generic similarities to Western Australian not represented in, Western Australia. Australian assemblages but there are few conspecific elements. Cenomanian floras mark the decline' of'many The presence of angiosperms and ginkgophytes in gymnosperm groups and diversification of the the Burrum flora suggests closer correlation with angiosperms (McLoughlin et al. 1995). the Victorian Zone C-D floras. Similarly, the Styx The Talbragar Fish Bed flora (Merrygoen Coal Measures of central Queensland are also Ironstone Member of the Purlawaugh Formation) characterized by the presence of angiosperms and illustrated by Walkom (1921a) and White (1981) is are probably of Albian age (Walkom 1919a; generally regarded as late Early Jurassic in age Dettmann and Playford 1969; Dettmann and (Hind and Helby 1969; Gould 1975) but contains a Douglas 1976). number of superficially similar plant fossils to the The little-studied Stanwell Coal Measures west Western Australian Early Cretaceous assemblages of Rockhampton, Queensland, are associated with including species of Taeniopteris, Thinnfeldia, marine strata yielding Neocomian bivalves Brachyphyllum (Araucaria herein), Nilssonia, aild (Skwarko 1968 and early to middle Albian Rissikia (= Elatocladus). However most forms can be palynofloras (Dettmann and Playford 1969). They distinguished at species level and the generic also contain a macroflora consisting of Equisetites similarities may be due to palaeoecological sp., Cladophlebis sp., Phyllopteroides laevis, similarities between these assemblages. The Early Taeniopteris spp. (cf. Taeniopteris daintreei), and Middle Jurassic floras of the Marburg Ptilophyllum pecten (cf. Ptilophyllum cutchense), Formation, Walloon Coal Measures, and Araucarites polycarpa Tenison-woods, and equivalents in the Clarence-Moreton Basin of New Elatocladus plana (cf. Elatocladus ginginensis), South Wales and Queensland (Walkom 1915, showing strong similarities to the Western 1917a,b, 1919b; Hill et al. 1966; Gould 1971, 1980; Australian assemblages (Walkom 1915, 1917a,b; McLoughlin and Drinnan 1995) also share several Hill et al. 1966; Cantrill and Webb 1987). Cantrill genera with the Western Australian Cretaceous and Webb (1987) suggested a Neocomian age for assemblages but clear species-level distinctions are the Stanwell Coal Measures owing to the apparent. Plant assemblages from the Dalrymple occurrence of the index fossil Phyllopteroides laevis Early Cretaceous macrofloras 55 and its distribution in Neocomian strata of the [Gleiehenites (=Mierophyllopteris), Hausmannia, and Surat and Gippsland Basins. Cladophlebis] and Conifers [Arauearia, Arauearites The floras most closely comparable to the (cone scales and cones), and Elatocladus]. This Western Australian assemblages are those from the Indian zone differs from the Western Australian Algebuckina Sandstone (Eromanga Basin) at Mt assemblages by its possession of ginkgophyte and Babbage, South Australia, and from the Lees Paehypteris species and its lack of Phyllopteroides, Sandstone and overlying beds (Carpentaria Basin), Roebuekia, and Nathorstianella species. The at Bauhinia Downs, Northern Territory. The Dietyozamites-Pterophy11 um-Anomozamites Algebuckina Sandstone incorpo!ates Middle Assemblage Zone is best represented in the Jurassic to Neocomian sediments (Moore 1986; Jabalp~r. Formation (Chaugan Forest district) of Wiltshire 1989). The macrofloras at Mt Babbage Madhya Pradesh (Crookshank 1936), the Lower occur in the upper part of this unit and are Rajmahal Formation (intertrappean beds 1-3, probably of Neocomian age. Although requiring Rajmahal Hills) of Bihar (Bose and Sah 1968; systematic revision, the Mt Babbage flora shares Sharma 1969b, 1971, 1975; Zeba-Bano et al. 1979), several taxa with the Western Australian and the lower Bhuj Formation of Kutch (Bose and assemblages (Glaessner and Rao 1955). Most Banerji 1984) together with a number of lesser­ notable are the co-occurrences of Nathorstianella studied'units in peninsula India, Sri Lanka, and babbagensis, Ptilophyllum eutehense, and Taeniopteris Nepal (see Sukh-Dev 1987). Certain genera (and daintreei. (White 1961b, 1966). The Bauhinia Downs perhaps species) from Sukh-Dev's (1987) flora likewise shares several key taxa with Western succeeding Alloeladus-Braehyphyllum-Pagiophyllum Australia including Ptilophyllum eutehense, Assemblage Zone and Weiehselia-Onyehiopsis­ Mierophyllopteris gleiehenioides, Hausmannia sp., and Gleiehenia Assemblage Zone are also shared with Taeniopteris daintreei, and owing to the abundance the Western Australian floras but these of bennettitaleans and lack of ginkgophytes, is of assemblages appear to be more akin to the probable Neocomian rather than Aptian age as Victorian Zone C (?late Neocomian-early Albian) suggested by Dettmann et al. (1992). floras. A Neocomian-Barremian age is proposed for the The Tico (Baquer6 Formation) Flora of Patagonia collective Western Australian terrestrial plant (Argentina) has a number of conifer, fern, and assemblages described here. Accumulation of plant bennettitalean genera in common with the Western remains in several regions along the continent's Australian Cretaceous floras (Herbst 1962a; western margin during the Neocomicin-Barremian Archangelsky 1963a,b,c, 1964a,b; Archangelsky and probably occurred as part of a single phase of Baldoni 1972) but the number of shared taxa are widespread fluvio-deltaic sedimentation fewer than between the Western Australian and ~ssociated with continental breakup, marginal Indian assemblages. The Baquer6 Formation has basin sag, and regional and global sea-level been assigned a late Barremian to early Aptian age fluctuations at that time (McLoughlin, et al. 1994). based on its palynoflora (Archangelsky et al. 1984) and most notably differs from the Western Comparisons with other Gondwanan floras Australian Early Cretaceous assemblages by its The Western Australian Cretaceous floras share possession of angiosperm remains (Archangelsky several genera and possibly some species with the 1963c; Romerro and Archangelsky 1986). It Rajmahal, Kutch, and Jabalpur floras of India. The probably has greater floristic affinities with the Rajmahal Series has been variously assigned to: the Victorian Zone C assemblages. lower, middle, or upper Jurassic according to A number of other lesser-studied Early traditional palaeobotanical studies (Feistmantel Cretaceous floras from Argentina show distinct 1877a; Halle 1913; Sahni and Rao 1933); the Lower similarities to the Western Australian assemblages. Cretaceous based on palynostratigraphy (Playford Though dated as Aptian on the basis of associated and Cornelius 1967); and the Albian according to ammonoid faunas, the Kachaike Formation, Santa radiometric (K-Ar) dating (McDougall and Cruz, has yielded Mierophyllopteris, Hausmannia, McElhinny 1970). Sukh-Dev (1987) re-evaluated the Cladophlebis, and Ptilophyllum species strikingly stratigraphic relationships of the Indian Mesozoic similar to those from the Broome Sandstone macrofloras and established 12 assemblage zones (Longobucco et al. 1985): The Apeleg Formation, from the Early Triassic to latest Cretaceous. He Chubut, also contains species of Taeniopteris, reassigned the lower Rajmahal assemblages to the Cladophlebis, and Ptilophyllum similar to those from Lower Cretaceous. Sukh-Dev's (1987) Western Australia and has been assigned a D ietyozamites-P terophy11 um-Anomozamites Hauterivian-Barremian age (Baldoni and de Vera Assemblage Zone corresponds most closely to the 1980). Although some elements (e.g., Tieoa, Zamites, Western Australian Cretaceous floras by its high and Cyeadolepis spp.) differ, the Spring Hill proportion of bennettitaleans, Taeniopteris, and the Formation of southern Argentina and Chile also occurrence of comparable species of ferns has a number of bennettitaleans, ferns, and 56 s. McLoughlin conifers, which are comparable to Western floras of Japan are distinguished from Australian Australian taxa, and has been assigned an Early floras by the presence of, or greater representation Cretaceous (probable Hauterivian-Barremian) age of, bennettitalean taxa such as Zamites, Neozamites, (Archangelsky 1976; Baldoni 1979, 1980c; Baldoni Anomozamites, Dictyozamites, Ctenozamites, and Taylor 1983). cycadaleans (such as Ctenis), czekanowskialeans, Macrofloras from the South African Kirkwood, and conifers including Podozamites, Mngazana, and Makatini Formations range from Cupressinocladus, and Frenelopsis (Kimura and Berriasian to late Aptian in age and are represented Ohana 1987a,b,c, 1992, 1988, 1989; Kimura et al. by Marchantites, Ricciopsis, Cladophlebis, 1985, 1991, 1992). Malayan and Thai mid- to late Sphenopteris, Onychiopsis, Taeniopteris, Pseudoctenis, Mesozoic floras (Kon'no 1967, 1972; Smiley 1970; Zamites, Dictyozamites, Araucaria, and Podocarpus Kon'no and Asama 1975; Asama et al. 1981) and species (Anderson and Anderson 1985). Most of coeval Indochinese macrofloras (Serra 1963, 1966; these are readily distinguishable from Australian Vozenin-Serra 1977) have affinities with other forms, a few are shared with South American southern (Ryoseki-type) east Asian floras (Kimura assemblages but the majority appear to be endemic 1987, 1988) and have few if any elements in to South Africa suggesting an early floristic common with Australian assemblages. separation of this province from other Gondwanan regions. Palaeoenvironments and palaeoclimates The upper Fossil Bluff Formation flora The presence of high latitude forests and (Barremian-Albian) of Alexander Island, terrestrial vertebrate populations together with Antarctica, contains Hausmannia, Cladophlebis, oxygen isotope signatures from marine Taeniopteris, and Elatocladus species comparable to invertebrates suggest globally raised temperatures those in the Western Australian assemblages but during the Early and mid-Cretaceous (Frakes 1979; differs in its possession of ginkgophytes, Douglas and Williams 1982; Barron 1983; Spicer Pagiophyllum, Bellarinea, and Phlebopteris. The upper and Parish 1986; Rich and Rich 1989; Frakes and Fossil Bluff Formation floras correspond more Francis 1990). Various factors including reduced closely with the Victorian Zone C assemblages. The obliquity of the Earth's rotational axis, Hope and Botany Bay floras of the Antarctic palaeogeographic configurations of continents and Peninsula were originally regarded as Jurassic in ocean currents, higher atmospheric CO2 levels, and age (Halle 1913). Later studies suggested an Early increased levels of solar radiation have been Cretaceous age (Francis 1986; Gee 1989) based on suggested as possible causes for the apparently an abundance of sphenopterid, cladophleboid, high palaeote~peratures (Douglas and Williams ta-eniopterid, bennettitalean, and podocarpacean 1982; Creber and Chaloner 1985; Barron 1983, remains similar to other Gondwanan Early 1984). Palaeomagnetic evidence (Veevers et al. Cretaceous assemblages. However, these Antarctic 1991) indicates that the Early Cretaceous Western assemblages are characterized by the presence of Australian floras existed in middle latitudes (45°• possible sagenopterid pteridosperms and 55°) within presumably warmer climates than Goeppertella favouring an Early Jurassic age (Rees those experienced by the higher palaeolatitude 1993). Victorian and Antarctic floras (Douglas 1969, 1973; Jefferson 1983). However, fossil woods from the Comparisons with extra-Gondwanan floras Birdrong Sandstone show well-defined growth Many plant families have wide, often rings consistent with a distinctly seasonal climate. cosmopolitan, distributions in the Jurassic, Analysis of 48 permineralized conifer wood however, by Early Cretaceous times fragmentation samples from the Giralia Anticline, Carnarvon of Pangaea led to renewed floral provincialism Basin (McLoughlin et al. 1994), showed mean equivalent to that of the late Palaeozoic (MeyEm sensitivity values in the range of 0.148 to 0.673 [i.e. 1987). Although some families and genera complacent to strongly sensitive (Fritts 1976)] with maintained wide distributions in the Early an average mean sensitivity of 0.348 (moderately Cretaceous, Gondwanan floras are readily sensitive). These growth indices together with the distinguishable from those of other provinces at presence of sporadic false growth rings within the species (and often genus) level. Geographically, the woods suggest a pronounced seasonal climate but closest extra-Gondwanan landmasses to Australia with substantial inter-seasonal variability in during the Early Cretaceous were probably the environmental parameters. series of disjunct terranes presently constituting The presence of matted Taeniopteris daintreei South China, Indochina, and Malesia (Audley­ leaves in the Bullsbrook Formation suggests a Charles 1988; Metcalfe 1991). Although Mesozoic deciduous or semi-deciduous habit for these floras have been reported from these areas (Kimura pentoxylaleans consistent with evidence from the and Ohana 1992), many have not yet been fully Victorian Cretaceous (Douglas and Williams 1982; documented. Late Jurassic and Early Cretaceous Drinnan and Chambers 1986) which suggests that Early Cretaceous macrofloras 57 these plants were adapted to a seasonal climatic CONCLUSIONS regime. A deciduous habit for other plant groups Thirty-two fossil plant and fungi taxa have been in the Western Australian assemblages is less described from five Early Cretaceous obvious although the complete and sometimes lithostratigraphic units in Western Australia. Four matted fronds of bennettitaleans such as new species (viz., Phyllopteroides westralensis, Ptilophyllum cutchense in the Broome Sandstone Roebuckia spatulata, Elatocladus ginginensis, and may reflect deciduousness. Carpolithes bullsbrookensis) are defined while several The range of fern and lycophyte species in the others are described under open nomenclature collections suggests a humid microthermal to owing to insufficient or inadequately preserved megathermal climate based on the distributions of material prohibiting comparisons to established related extant taxa. However, most of the taxa. The fossil assemblages comprise lycophytes, fossiliferous units were deposited in lowland ferns, pteridosperms, pentoxylaleans, deltaic environments where high water table bennettitaleans, and araucarian and podocarpacean (rheotrophic) rather than necessarily high rainfall conifers. Bennettitaleans, pentoxylaleans, and (ombotrophic) conditions may have aided the conifers are the numerically dominant elements of growth of pteridophytes. Modem cycadophytes the flora. Macrofossil suites from the Broome typically grow in mesothermal to megathermal Sandstone (Canning Basin), Nanutarra Formation conditions and most forms probably occupied and Birdrong Sandstone (Carnarvon Basin), and similar climates in the past (Douglas and Williams Leederville and Bullsbrook Formations (Perth 1982). The abundance of bennettitalean foliage in Basin) are correlated with Douglas' (1969) the studied assemblages favours interpretation of a Ptilophyllum-Pachypteris austropapillosa Zone (Zone relatively warm (though seasonal) climate. Some of B) of Victoria and indicate a Neocomian­ the bennettitaleans have comparatively small Barremian age. The Nanutarra Formation is a fronds (e.g., Ptilophyllum boolensis) which may have diachronous marginal marine to paralic unit and been a foliar response to local water stress. its age may locally extend to the Aptian based on Differential representation of plant groups invertebrate assemblages (Cox 1961). The Western between the various Western Australian Australian Early Cretaceous floras are most closely Neocomian-Barremian assemblages is relatively comparable with coeval assemblages from South minor and probably reflects local differences Australia and the Northern Territory. Lesser between depositional settings (e.g., alluvial valley similarities are evident with eastern Australian and versus coastal environments). The Nanutarra Indian floras and fewer species are shared with Formation yields a small macroflora mixed with a South America, Antarctica, and South Africa. Few, marine invertebrate fauna (Cox 1961) and was if any, species are shared with extra-Gondwanan probably deposited within paralic to shallow assemblages. The representation of plant groups, marine conditions. The coastal to nearshore together with wood growth indices, sediments of the Birdrong Sandstone lack leaf palaeomagnetic data, and oxygen isotope remains but contain abundant mollusc-bored fossil signatures, suggests that the Western Australian wood, ammonoids, and marine vertebrates marginal basins experienced humid seasonal (McLoughlin et al. 1994). Whilst some of the plant mesothermal climates in mid-latitudes during the remains in the Broome Sandstone are clearly Neocomian-Barremian. Local variations between allochthonous, autochthonous root beds and assemblages are probably attributable to Nathorstianella corms are evident in exposures at differences in depositional settings, seasonal Garttheaume Point, Broome. This coarse-grained effects, and taphonomic factors rather than unit also contains diverse dinosaurian trackways significant climatic or stratigraphic anomalies (Long 1990) and sparse bivalves (Yeates et al. 1984) between localities and host formations. .and was proQ.ably deposited within a deltaic setting. The plant-bearing Callawa Formation and Cronin Sandstone, although not examined during ACKNOWLEDGEMENTS this study, lack invertebrate fossils and were Thanks are due to the numerous museum and probably deposited within proximal fluvial university officers and members of the public who, (alluvial valley/alluvial plain) settings. The over the past three decades, have contributed Leederville Formation consists of interbedded Cretaceous plant fossils from various localities to sandstones, siltstones, and claystones and was the Western Australian Museum and the deposited within paralic to shallow marine University of Western Australia. Dr Ken environments (Cockbain 1990). The correlative McNamara and Dr John Long of the Western Bullsbrook Formation is dominantly composed of Australian Museum are thanked for arranging the sandstone but contains local plant-bearing loan of the museum's specimens. Dr David Haig overbank shale lenses and was deposited within (University of Western Australia) and Mr Glynn fluvial environments (Cockbain 1990). Ellis (University of Lausanne) assisted with I -- -- 58 s. McLoughlin

collecting in the Carnarvon Basin. This research southern Thailand. Geology and Palaeontology of was carried out while the author held an Southeast Asia 22: 35-47. Australian Research Council Postdoctoral Ash, S.R. and Pigg, K.B. (1991). A new Jurassic Isoetites Fellowship at the University of Western Australia. (Isoetales) from the Wallowa Terrane in HelIs Canyon, Oregon and Idaho. American Journal of Botany 78: 1636-1642. Audley-Charles, M.G. (1988). Evolution of the southern REFERENCES margin of Tethys (north Australian region) from Early Permian to Late Cretaceous. In Gondwana and Anderson, J.M. and Anderson, H.M. (1985). Palaeoflora of southern AfrIca. Prodromus of South African Tethys M.G. Audley-Charles and A HalIam (eds), Geological Society Special Publication 37: 79-100. megafloras Devonian to Lower Cretaceous. A.A. Balkema, Rotterdam. 416 pp. Backhouse, J. (1988). Late Jurassie and Early Cretaceous palynology of the Perth Basin, Western Australia. Anderson, J.M. and Anderson, H.M. (1989). Palaeoflora of southern Africa. Molteno Formation (Triassic) Volume Geological Survey of Western Australia Bulletin 135: 1­ 233. 2 Gymnosperms (excluding Dicroidium). A.A. Balkema, Rotterdam. 567 pp. Backhouse, J., Godfrey, 1., and McLoughlin, S. (1995). Cretaceous fossil wood from Collie. Geological Survey Andrews, H.N. Jr (1970). Index of generie names of ofWestern Australia Annual Review 1994-1995: 51-54. fossil plants, 1820-1865. U.S. Geological Survey Bulletin 1300: 1-354. Baksi, S.K. (1968). Fossil plants from Raghavapuram Mudstone, West Godavari district, A.P., India. Arber, EAN. (1910). Some fossil plants from Western Palaeobotanist 16: 206-215. Australia. Geological Survey of Western Australia Baksi, S.K. and Naskar, P. (1981). Fossil plants from Bulletin 36: 25-28. Sarnu Hill Formation, Barmer Basin, Rajasthan. Arber, E.AN. (1917). The earlier Mesozoie floras of New Palaeobotanist 27: 107-111. Zealand. New Zealand Geological Survey Baldoni, AM. (1979). Nuevos elementos paleofloristicos Palaeontological Bulletin 6: 1-80. de la tafoflora de la Formaci6n Spring Hill, limite Archangelsky, S. (1963a). Notas sobre la flora f6sil de la Jurasieo-Cretacico subsuelo de Argentina y Chile zona de Tie6, Provineia de Santa Cruz. Ameghiniana austral. Ameghiniana 16: 103-119. 3: 57-62. Baldoni, AM. (1980a). Tafoflora Jurasica de la zona de Archangelsky, S. (1963b). Notas sobre la flora f6sil de la PieUn Leufu, Provincia del Neuquen. Ameghiniana 17: zona de Tic6, Provineia de Santa Cruz 2. Tres nuevas 243-254. especies de Mesosingeria. Ameghiniana 3: 113-120. Baldoni, AM. (1980b). Plantas fosiles Jurasicas de una Archangelsky, S. (1963c). A new Mesozoie flora from nueva localidad en la provincia del Nequen. Tie6, Santa Cruz Province, Argentina. Bulletin of the Ameghiniana 17: 255-272. British Museum (Natural History), Geology 8(2): 1-92. Baldoni, AM. (1980c). Nota sobre una nueva espeeie de ''Archangelsky, S. (1964a). Notas sobre la flora f6sil de la Elatocladus (?) (Coniferae) de la Formaeion Springhill, zona de Tie6, Provineia de Santa Cruz 3. Rriflorinia limite Jurasieo-Cretacico, Provincia de Santa Cruz. pilifera n. sp., 4. Equisetites sp. Ameghiniana 3: 221­ Ameghiniana 7: 373-378. 226. Baldoni, AM. (1981). Tafoflora del Jurasieo Superior de Archangelsky, S. (1964b). Notas sobre la flora f6sil de la la zona de Lago Argentino Provineia de Santa Cruz, zona de Tie6, Provineia de Santa Cruz 5. Sphenopteris Republica Argentina. Ameghiniana 18: 97-102. cf. goepperti Dunker, 6. Cladophlebis sp. Ameghiniana Baldoni, A and de Vera, H. (1980). Plantas fosiles de la 3: 280-284. Formaci6n Apeleg (Cretaeico) en la zona del Lago Archangelsky, S. (1976). Vegetales fosiles de la Fontana, Provincia de Chubut. Ameghiniana 17: 289­ Formaei6n Springhill, Cretaeico, en el subsuelo de la 296. Cuenca Magallaniea, Chile. Ameghiniana 13: 141-158. Baldoni, A and Taylor, T.N. (1983). Plant remains from Archangelsky, S. and Baldoni, A (1972). Notas sobre la a new Cretaceous site in Santa Cruz, Argentina. flora f6sil de la zona de Tie6, Provineia de Santa Review of Palaeobotany and Palynology 39: 301-311. Cruz 10. Dos nuevos especies de Pseudoctenis Bancroft, N. (1913). On some Indian Jurassic (Cycadales). Ameghiniana 9: 241-257. gymnosperms. Transactions of the Linnean Society of Archangelsky, S., Baldoni, A, Gamerro, J.c. and Seiler, London 8: 69-86. J. (1984). Palinologia estratigrafiea del Cretaeico de Banerji, J. (1989). Some Mesozoie plant remains from Argentina austral. 3. Distribueion de las especies y Bhuj Formation with remarks on the depositional conclusiones. Ameghiniana 21: 15-33. environment of beds. Palaeobotanist 37: 159-168. Arrondo, O.G. and Petriella, B. (1980). Alicura, nueva Banerji, J. and Pal, P.K. (1986). Mesozoic plant remains localidad plantifera Liasieo de la provincia de from Sarnu, Barmer District, Rajasthan. Palaeobotanist Neuquen, Argentina. Ameghiniana 17: 200-215. 35: 141-145. Arrondo, O.G. and PetrielIa, B. (1982). Revision del Barron, E.J. (1983). A warm, equable Cretaceous: The genera Goeppertella Oishi et Yamashita emend. nature of the problem. Earth Science Reviews 19: 305­ (Goeppertelloideae - Dipteridaceae). Ameghiniana 19: 338. 67-78. Barron, E.J. (1984). Climatic implications of the variable Asama, K., Nakornsri, N., Hinthong, c., and Sinsakul, obliquity explanation of Cretaceous-Paleogene high­ S. (1981). Some younger Mesozoic plants from Trang, latitude floras. Geology 12: 595-598. Early Cretaceous macrofloras 59

Blanchette, R.A. (1980). Wood decomposition by Brongniart, A. (1849). Tableau des genres de vegetaux Phellinus (Fomes) pini: a scanning electron fossiles consideres sous le point de vue de leur microscopy study. Canadian Journal of Botany 58: classification botanique et de leur distribution 1496-1503. geologique. Dictionnaire Universal Histoire Naturelle Blaschke, P.M. and Grant-Mackie, J.A. (1976). Mesozoic 13: 1-127. leaf genus Taeniopteris at Port Waikato and Clent Brown, R.W. (1939). Some American fossil plants Hills, New Zealand. New Zealand Journal of Geology belonging to the Isoetales. Journal of the Washington and Geophysics 19: 933-941. Academy of Science 29: 261-269. Bock, W. (1962). A study on fossil Isoetes. Journal of Brown, RW. (1958). New occurrences of the fossil Paleontology 36: 53-59. quillworts called Isoetites. Journal of the Washington Bohra, D.R and Sharma, B.D. (1980). Araucarites mittrii Academy of Science 48: 358-361 sp. nov., a petrified megastrobilus from the Rajmahal Brunnschweiler, R.O. (1960). Marine fossils from the Hills, India. Ameghiniana 17: 3-9. Upper Jurassic and the Lower Cretaceous of Bose, M.N. (1953a). Bucklandia sahnii sp. novo from the Dampier Peninsula, Western Australia. Bureau of Jurassic of the Rajmahal Hills, Bihar. Palaeobotanist 2: Mineral Resources, Geology and Geophysics, Australia, 41-50. Bulletin 59: 1-52. Bose, M.N. (1953b). On some fossil cycadean stems from Cantrill, D.J. and Webb, J.A. (1987). A reappraisal of the Rajmahal Hills, Bihar. Palaeobotanist 2: 71-74. Phyllopteroides Medwell (Osmundaceae) and its Bose, M.N. (1958). Some fragmentary plant fossils from stratigraphic significance in the Lower Cretaceous of Narsinghpur district, Madhya Pradesh, India. eastern Australia. Alcheringa 11: 59-85. Palaeobotanist 6: 49-50. Chapman, F. (1904). On a collection of Upper Palaeozoic Bose, M.N. (1974). The genus Otozamites Braun from the and Mesozoic fossils from Western Australia and Mesozoic rocks of India. Palaeontographica 147B: 100­ Queensland, in the National Museum, Melbourne. 106. Proceedings of the Royal Society of Victoria 16: 306-335. Bose, M.N. and Banerji, J. (1984). The fossil floras of Chapman, F. (1908). Report on Jurassic plants. Geological Kachchh. 1 - Mesozoic megafossils. Palaeobotanist 33: Survey of Victoria Record 2: 212-220. 1-189. Chapman, F. (1909). Jurassic plant-remains from Bose, M.N. and Jana, RN. (1979). Dictyophyllum and Gippsland. Geological Survey of Victoria Record 3: 103­ Hausmannia from the Lower Cretaceous of 111. Saurashtra, India. Palaeobotanist 26: 180-184. Chapman, F. (1914). Miscellaneous reports on fossils. Bose, M.N. and Kasat, M.L. (1972). The genus Geological Survey of Victoria Record 3: 317-321. Ptilophyllum in India. Palaeobotanist 19: 115-145. Clifford, H.T. and Constantine, J. (1980). Ferns, Fern Bose, M.N. and Roy, S.K. (1964). Studies on the Upper Allies and Conifers of Australia. University of Gondwana of Kutch - 2. Isoetaceae. Palaeobotanist 12: Queensland Press, St Lucia. 150 pp. 226-228. Cockbain, A.E. (1990). Perth Basin. In, Geology and ··''Bbse, MN. and Sah, S.C.D. (1968). Some-pteridophytic Mineral Resources of Western Australia, Geological remains from the Rajmahal Hills, Bihar. Survey of Western Australia Memoir 3: 495-524. Palaeobotanist 16: 12-28. Colbert, E.H. and Merrilees, D. (1967). Cretaceous Bose, MN. and Sukh Dev (1960). Studies on the fossil dinosaur footprints from Western Australia. Journal flora of the Jabalpur Series from the South Rewa of the Royal Society ofWestern Australia 50: 21-25. Gondwana Basin - 1. Cycadopteris, Nipaniophyllum Corsin, P. and Waterlot, M. (1979). Palaeobiogeography and Ginkgoites. Palaeobotanist 7: 143-154. of the Dipteridaceae and Matoniaceae of the Bose, M.N. and Sukh-Dev (1961). Studies on the fossil Mesozoic. Proceedings of the Fourth International flora of the Jabalpur series from the south Rewa Gondwana Symposium. Hindustan Publishing Gondwana Basin - 2. Onychiopsis paradoxus n. sp. Corporation (India), Delhi 1: 51-70. Palaeobotanist 8: 57-64. Cox, L.R (1961). The molluscan fauna and probable Bose, M.N., Kutty, T.s. and Maheshwari, H.K. (1982). Lower Cretaceous age of the Nanutarra Formation of Plant fossils from the Gangapur Formation. Western Australia. Bureau of Mineral Resources, Palaeobotanist 30: 121-142. Geology and Geophysics, Australia, Bulletin 61: 1-53. Brongniart, A. (1822). Sur la classification et la Creber, G.T. and Chaloner, W.G. (1985). Tree growth in distribution des vegetaux fossiles en general, et sur the Mesozoic and Early Tertiary and the ceux des terrains de sediment superieur en reconstructions of palaeoclimates. Palaeogeography, particulier. Museum Natlional Histoire Naturelle Palaeoclimatology, Palaeoecology 52: 35-60. (Paris) Memoire 8: 203-348. Crookshank, H. (1936). Geology of the northern slopes Brongniart, A. (1825). Observations sur les vegetaux of the Satpura between the Morand and the Sher fossiles renfermes dans les Gres de Hoer en Scanie. Rivers. Geological Survelj of India Memoir 66: 173-381. Annales des Sciences Naturelle, ser. 1, 4: 200-219. Del Fueyo, G.M. (1991). Una nueva Araucariaceae Brongniart, A. (1828). Prodrome d'une histoire des vegetaux Cretacica de Patagonia, Argentina. Ameghiniana 28: fossiles. Paris. 223 pp. 149-161. Brongniart, A. (1832). Histoire des vegetaux fossiles ou Del Fueyo, G., Archangelsky, S. and Taylor, T.N. (1990). recherches botaniques et geologiques sur les vegetaux Una nueva podocarpacea fertil (coniferal) del renfermes dans les diverses couches du globe. G. Dufour Cretacico inferior de Patagonia, Argentina. and E. D'Ocagne, Paris 1: 265-288. Ameghiniana 27: 63-73. 60 s. McLoughlin

Dettmann, M.E. (1963). Upper Mesozoic microfloras Feistmantel, O. (1877c). Flora of the Jabalpur Group from southeastern Australia. Proceedings of the Royal (upper Gondwami), in the Son-Narbada region. Society of Victoria 77: 1-148. Geological Survey of India. Palaeontologia Indica, Dettmann, M.E. and Douglas, J.G. (1976). Palaeontology. Memoir, ser. 11,2(2): 1-25. In, Geology of Victoria J.G. Douglas and J.A Ferguson Feistmantel, O. (1879). The fossil flora of the Upper (eds). Geological Society of Australia, Victorian Gondwanas. OutHers on the Madras Coast. Geological Division, Melbourne. pp. 164-176. Survey of India, Palaeontologia Indica, Memoir, ser. 2, Dettmann, M.E. and Playford, G. (1969). Palynology of 1(4): 191-224. the Australian Cretaceous: a review. In, Stratigraphy Feldtman, F.R. (1963). Some pelycypods from the and Palaeontology. Essays in honour of Dorothy Hill, Cretaceous Gingin Chalk, Western Australia, K.S.W. Campbell (ed.), Australian National together with descriptions of the principal chalk University Press, Canberra. pp. 174-210. exposures. Journal of the Royal Society of Western Dettmann, M.E., Molnar, RE., Douglas, J.G., Burger, D., Australia 46: 101-125. Fielding, e., Clifford, HT., Francis, J., Jell, P., Rich, Feruglio, E. (1937). Dos nuevos especies de Hausmannia T., Wade, M., Rich, P.V., Pledge, N., Kemp, A and de la Patagonia. Notas Museo de La PIa ta, 2, Rozefelds, A (1992). Australian Cretaceous terrestrial Paleontologia 9: 125-136. faunas and floras: biostratigraphic and biogeographic Frakes, L.A (1979). Climates throughout geologic time. implications. Cretaceous Research 13: 207-262. Elsevier, Amsterdam. 310 pp. Douglas, J.G. (1969). The.Mesozoic floras of Victoria, Frakes, L.A. and Francis, J.E., (1990). Cretaceous Parts 1 and 2. Geological Survey of Victoria Memoir 28: palaeoclimates. In, Cretaceous Resources, Events and 1-310. Rhythms R.N. Ginsburg and B. Beaudoin (eds), Douglas, J.G. (1971). Biostratigraphical subdivision of Kluwer Academic Publishers, Dordrecht. pp. 273­ Otway Basin Lower Cretaceous sediments. In, The 287. Otway Basin ofsoutheastern Australia, H. Wopfner and Francis, J.E. and Coffin, M.F. (1992). Cretaceous fossil J.G. Douglas (eds), Special Bulletin of the Geological wood from the Raggatt Basin, southern Kerguelen Surveys of South Australia and Victoria. pp. 187-19l. Plateau (Site 750). In, Wise, S.W., Schlich, R, et al. Douglas, J.G. (1973). The Mesozoic floras of Victoria, Proceedings of the Ocean Drilling Program, Scientific Part 3. Geological Survey of Victoria Memoir 29: 1-185. Results, 120. College Station, Texas (Ocean Drilling Douglas, J.G. and Williams, G.E. (1982). Southern polar Program). pp. 273-280. forests: the Early Cretaceous floras of Victoria and Francis, J.E. (1986). Growth rings in Cretaceous and their palaeoclimatic significance. Palaeogeography, Tertiary wood from Antarctica and their Palaeoclimatology, Palaeoecology 40: 199-212. palaeoclimatic implications. Palaeontology 29: 665­ Drinnan, A.N. and Chambers, T.e. (1985). A 684. reassessment of Taeniopteris daintreei from the Frenguelli, J. (1947). El genero Cladophlebis y sus Victorian Early Cretaceous: a member of the representantes en la Argentina. Anales del Museo de Pentoxylales and a significant Gondwanaland plant. La Plata (n.s.) Paleontologia B, 2: 1-74. Australian Journal ofBotany 33: 89-100. Fritts, H.C. (1976). Tree rings and climate. Academic Press, Drinnan, AN. and Chambers, T.e. (1986). Flora of the London. 576 pp. Lower Cretaceous Koonwarra fossil bed Gee, e.T. (1989). Revision of the Late Jurassic/Early (Korumburra Group), South Gippsland, Victoria. Cretaceous flora from Hope Bay, Antarctica. Association of Australasian Palaeontologists Memoir 3: Palaeontographica 213B: 149-214. 1-77. Glaessner, M.F. and Rao, V.R (1955). Lower Cretaceous Dun, W.S. (1898). Notes on the Australian plant remains from the vicinity of Mount Babbage, Taeniopteridae. Report of the Australian and New South Australia. Transactions of the Royal Society of Zealand Association for the Advancement of Science 7: South Australia 78: 134-140. 384-400. Goeppert, H.R. (1836). Die fossilen Farrenkrauter Dunker, W. (1846). Monographie der Norddeutschen (Systema filicum fossilium). Nova Acta Leopoldina 17: Wealdenbildung. Braunschweig. 83 pp. 1-486. Ettingshausen, e. von.· (1852). Die Steinkohl~nflora van Goeppert, H.R (1841). Les genres des' plantes fossiles> Stradonitz in B6hmen. Kaiserlich-Koenigliche Bonn. pp. 1-70. Geologische Reichsanstalt, Abhandlungen, and c. I, Gould, RE. (1971). Report on plant fossils from Durikai, pt.3(4): 1-18. southeastern Queensland. Appendix 3. In, The post­ Feistmantel, O. (1876). Jurassic (Oolitic) flora of Kach. Palae020ic rocks of the Warwick 1:250 000 Sheet area, Geological Survey of India. Palaeontologia Indica, Queensland and New South Wales, N.F. Exon, RF. Memoir, ser. 11,2(1): 1-80. Reiser, D.J. Casey and RL. Brunker. Bureau ofMineral Feistmantel, O. (1877a). Jurassic (Liassic) flora of the Resources, Geology and Geophysics, Australia, Report Rajmahal Group in the Rajmahal Hills. Memoirs of 140: 63-64. the Geological Survey of India. Palaeontologia Indica, ser. Gould, R.E. (1975). The succession of Australian pre­ 2,1(2): 1-110. Tertiary megafossil floras. Botanical Review 41: 453­ Feistmantel, O. (1877b). Jurassic (Liassic) flora of the 483. Rajmahal Group from Golapili (near Ellore), South Gould, RE. (1978). Report on plant fossils from the Godavari district. Geological Survey of India. Tarlton Range, Tobermory 1: 250 000 Sheet area, Palaeontologia Indica, Memoir, ser. 2, 1 (3): 1-28. Northern Territory. In, Geology of the Eromanga Basin, Early Cretaceous macrofloras 61

B.R Senior, A Mond and P.L. Harrison. Bureau of Hind, M.C. and Helby, RJ. (1969). The Great Artesian Mineral Resources, Geology and Geophysics, Australia, Basin in New South Wales. In, Geology of New South Bulletin 167: 87-88. Wales. G.H. Packham (ed.). Journal of the Geological Gould, R.E. (1980). The coal-forming flora of the Society ofAustralia 16: 481-497. Walloon Coal Measures. Australian Coal Geology 1: Hocking, RM. (1990). Carnarvon Basin. In, Geological 83-105. Survey of Western Australia, Geology and Mineral Greguss, P. (1955). Identification of living gymnosperms on Resources of Western Australia. Geological Survey of the basis of xylotomy. AkademiaiKaido, Budapest. 263 Western Australia, Memoir 3: 457 - 495. pp. Hocking, RM., Moors, H.T., and van de Graaff, W.J.E. Gupta, KM. (1955). Notes on some Jurassic plants from (1987). Geology of the Camarvon Basin. Geological the Rajmahal hills, Bihar, India. Palaeobotanist 3: 1­ Survey of Western Australia Bulletin 133: 1-289. 25. Hooker, W.J. (1843). Figure and description of a new Halle, T.G. (1913). The Mesozoic flora of Graham Land. species of Araucaria, from Moreton Bay, New Wissenschaftliche Ergebnisse der Schwedischen Siidpolar­ Holland, detected by J.T. Bidwill, Esq. Hooker's expedition 1901-1903 3: 1-123. London Journal of Botany 2: 498-506. Harris, T.M. (1962). The occurrence of the fructification Jack, RL. and Etheridge, R Jr. (1892). The geology and Carnoconites in New Zealand. Transactions of the Royal plaeqntology of Queensland and New Guinea. Society ofNew Zealand (Geology) 1: 17-27. Geological Survey of Queensland Publication 72: 1-766. Harris, T.M. (1964). The Yorkshire Jurassic flora n. ­ Jain, KP. (1968). Some plant remains from the Upper Caytoniales, Cycadales and Pteridosperms. British Gondwana of east coast, India. Palaeobotanist 16: 151­ Museum (Natural History), London. 191 pp. 155. Harris, T.M. (1969). The Yorkshire Jurassic Flora rn. Jefferson, T.H. (1983). Palaeoc1imatic significance of Bennettitales. British Museum (Natural History), some Mesozoic Antarctic fossil forests. In, Antarctic London. 186 pp. Earth Science, RL. Oliver, P.R James and J.B. Jago Harris, T.M. (1979). The Yorkshire Jurassic Flora, IV. (eds). Australian Academy of Science, Canberra. pp. Coniferales. British Museum (Natural History), 593-598. London. 166 pp. Jefferson, T.H. (1987). The preservation of conifer wood: Helby, R, Morgan, R and Partridge, AD. (1987). A examples from the Lower Cretaceous of Antarctica. palynological zonation of the Australian Mesozoic. Palaeontology 30: 233-249. Association of Australasian Palaeontologists Memoir 4: Jussieu, AL. de (1789). Genera plantarum secundum 1-94. ordines naturales disposita, juxta methodum in horto Herbst, H. (1960). Descripcion de ejemplares fertiles de regio Parisiensi exaratum, anno 1774. Paris. 498 pp. «Hausmannia (Protorhipis) papilio» Feruglio, Kaulfuss, G.F. (1824). Enumeratio Filicum, quas in itinere procedentes de los estratos de Baquero (Santa Cruz). circa terram legit. A. de Chamisso adjectis in omnia Acta Geologica Lilloana 3: 227-235. haram Plantarum genera permultasque species non satis Herbst,'R (1962a). Sobre las especies de Gleichenites de cognitas vel novas animadversionibus. Lipsiae. 300 pp. los sedimentos Baqueroenses de Santa Cruz, Karrfalt, E. (1986). New observations on the structure Patagonia. Ameghiniana 2: 141-147. and development of Nathorstianella. Review of Herbst, R. (1962b). Gleichenites juliensis n. sp. Del Palaeobotany and Palynology 47: 1-8. Junisico medio de Santa Cruz, Patagonia. Kasat, M.L. (1970). Some pteridophytic remains from the Ameghiniana 2: 187-191. Mesozoic rocks of India. Palaeobotanist 18: 212-214. Herbst, R. (1964). La flora Uasica de C. Meschio, Kimura, T. (1987). Recent knowledge of Jurassic and Provincia de Chubut, Patagonia. Ameghiniana 3: 227­ Early Cretaceous floras in Japan and 234. phytogeography of this time in east Asia. Tokyo Herbst, R (1966). Revision de la flora Liassica de Piedra Gakugei University, Sect. IV, Mathematics and Natural Pintada, prov. Neuquen, Argentina. Revista del Museo Sciences Bulletin 39: 87-115. de La Plata (n.s.) Paleontologia 5: 27-53. Kimura, T. (1988). Jurassic macrofloras in Japan and Herbst, R (1971). Palaeophytologia Kurtziana. Ill. 7. palaeophytogeography in east Asia. Tokyo Gakugei Revision de las especies Argentinas del Genero University, Sect. IV, Mathematics and Natural Sciences Cladophlebis. Ameghiniana 8: 265-281. Bulletin 40: 147-164. Herbst, R. (1975). Consideraciones sobre las Kimura, T. and Ohana, T. (1987a). Some Early Dipteridaceae (Filices), del Gondwana. Actas I Cretaceous plants from the Outer Zone of Japan. Congreso Argentino Paleontologia y Bioestratigrafia, National Science Museum, Tokyo, ser. C (Geology and Tucuman 1: 525-535. Paleontology) Bulletin 13: 13-27. Herbst, R. (1979). Review of the Australian Kimura, T. and Ohana, T. (1987b). Middle Jurassic and Dipteridaceae. Proceedings of the Linnean Society of some late Liassic plants from the Toyora Group, New South Wales 103: 7-21. southwest Japan (I). National Science Museum, Tokyo, Hill, D., Playford, G. and Woods, J.T. (1966). Jurassic ser. C (Geology and Paleontology) Bulletin 13: 41-76. fossils of Queensland, Queensland Palaeontographical Kimura, T. and Ohana, T. (1987c). Middle Jurassic and Society, Brisbane. 32 pp. some late Liassic plants from the Toyora Group, Hill, R.S. (1987). Tertiary Isoetes from Tasmania. southwest Japan [I1]. National Science Museum, Tokyo, Alcheringa 12: 157-162. ser. C (Geology and Paleontology) Bulletin 13: 115-148. 62 s. McLoughlin

Kimura, T. and Ohana, T. (1988). Late Jurassic plants de la Provincia de Santa Cruz. Ameghiniana 21: 305­ from the Tochikubo Formation (Oxfordian), 315. Somanakamura Group, in the Outer Zone of McClelland, J. (1850). Report of the Geological Survey of northeast Japan. National Science Museum, Tokyo, ser. India for the Season 1848-49. Calcutta. 53 pp. C (Geology and Paleontology) Bulletin 14: 151-185. McCoy, F. (1874). Prodromus of the Palaeontology of Kimura, T. and Ohana, T. (1989). Late Jurassic plants Victoria. Geological Survey of Victoria, Melbourne. from the Oginohama Formation, Oshika Group in 37 pp. Outer Zone of northeast Japan (Il). National Science McDougall, I. and McElhinny, M.W. (1970). The Museum, Tokyo, ser. C (Geology and Paleontology) Rajmahal Traps of India - K-Ar ages and Bulletin 15: 53-70. palaeomagnetism. Earth and Planetary Science Letters Kimura, T. and Ohana, T. (1992). Cretaceous 9: 371-378. palaeobotany and phytogeography in eastern McLoughlin, S. (1993). Plant fossil distributions in some Eurasia. Paleontological Society of Korea, Special Australian Permian non-marine sediments. Publication 1: 27-34. Sedimentary Geology 85: 601-619. Kimura, T., Saiki, K, and Arai, T. (1985). Frenelopsis McLoughlin, S. and Drinnan, AN. (1995). A Middle choshiensi sp. nov., a cheirolepidaceous conifer from Jurassic flora from the Walloon Coal Measures, the Lower Cretaceous Chosi Group in the Outer Mutdapilly, Queensland, Australia. Memoirs of the Zone of Japan. Proceedings of the Japan Academy 61 Queensland Museum 38: 257-272. (ser. B): 426-429. McLoughlin, S. and Guppy, L. (1993). Western Kimura, T., Ohana, T., and Aiba, H (1990). Late Jurassic Australia's Cretaceous floras. Fossil Collector Bulletin plants from the Shishiori Group, in the Outer Zone 39: 11-2l. of northeast Japan (1). National Science Museum, McLoughlin, S. and Hill, RS. (in press). The succession Tokyo, ser. C, (Geology and Paleontology) Bulletin 16: of Western Australian Phanerozoic floras. In, 127-153. Gondwanan Heritage. Systematics, Evolution and Kimura, T., Ohana, T., and Aiba, H. (1991). Late Jurassic Conservation of the Western Australian Biota. Hopper, plants from the Shishiori Group, in the Outer Zone S. et al. (eds), Perth. pp. 000-000. of northeast Japan (Il). National Science Museum, Tokyo, McLoughlin, S., Haig. D.W., Backhouse, J, Holmes, ser. C (Geology and Paleontology) Bulletin 17: 21-40. M.A, Ellis, G., Long, J.A. and McNamara, KJ. Kimura, T., Lebedev, E.L., Markovich, E.M., and (1994). Oldest Cretaceous sequence, Giralia Anticline, Samylina, V.A (1992). Macroflora of eastern Asia Carnarvon Basin, Western Australia: late and other circum-Pacific areas. In, The Jurassic of the Hauterivian-Barremian. AGSO Journal of Australian Circum-Pacific. G.E.G. Westermann (ed.), Cambridge Geology and Geophysics 15: 445-468. University Press, Cambridge. pp. 311-316. McLoughlin, S., Drinnan, A.N., and Rozefelds, A.C Koch, K (1873). Dendrologie: Baeume, Straeucher, und (1995). Th~ Cenomanian flora of the Winton Halbstraeucher, welche in Mittel- und Nord-Europa im Formation, Eromanga Basin, Queensland, Australia. Freien Kultivirt werden / Kritisch beleuchtet von K.Koch. Memoirs of the Queensland Museum 38: 273-313.- Erlangen. 2(2): 206. McWhae, J.R.H., Playford, P.E., Undner, A.W., Kon'no, E. (1967). Some younger Mesozoic plants from Glenister, B.F., and Balme, B.E. (1958). The Malaya. Geology and Palaeontology of Southeast Asia 3: stratigraphy of Western Australia. Journal of the 135-164. Geological Society ofAustralia 4: 1-16l. Kon'no, E. (1972). Addition to some younger Mesozoic Maheshwari, HK. and Singh, N.P. (1976). On some plants from Malaya. Geology and Palaeontology of plant fossils from the Pariwar Formation, Jaisalmer Southeast Asia 4: 139-155. Basin, Rajasthan. Palaeobotanist 23: 116-123. Kon'no, E. and Asama, K (1975). Mesozoic plants from Medwell, L.M. (1954a). A review and revision of the Khorat, Thailand. Geology and Palaeontology of flora of the Victorian Lower Jurassic. Proceedings of Southeast Asia 12: 149-17l. the Royal Society ofVictoria 65: 63-111. Lele, KM. (1955). Plant fossils from Parsora in the south Medwell, L.M. (1954b). Fossil plants from Killara, near Rewa Gondwana Basin, India. Palaeobotanist 4: 23­ Casterton. Proceedings of the Royal Society of Victoria 34. 66: 17-23. Lindley, J. and Hutton, W. (1834). The fossil flora of Great Mehta, K.R and Sud, J.D. (1953). On some ginkgoalean Britain, or figures and descriptions of the vegetable leaf impressions from the Rajmahal Hills. remains found in a fossil state in this country. London. Palaeobotanist 2: 51-54. 2: 57-156. Metcalfe, I. (1991). Late Palaeozoic and Mesozoic Linnaeus, C von. (1753). Species plantarum, exhibentes palaeogeography of Southeast Asia. Palaeogeography, plantasrite cognitas, ad genera relatas, cum differentiis Palaeoclimatology, Palaeoecology 87: 211-22l. specificis, nominibus trivialibus, synonymis selectis, locis Meyen, S.V. (1987). Principles of Palaeobotany. Chapman natalibus, secundum systema sexuale digestas. L. Salvii, and Hall, London. 432 pp. Holm. 1200 pp. Middleton, M.F. (1990). Canning Basin. In, Geology and Long, J.A. (1990). Dinosaurs of Australia. Reed Books, Mineral Resources of Western Australia, Geological Balgowlah. 87 pp. Survey of Western Australia Memoir 3: 425 - 453. Longobucco, M.I., Azcuy, c.L. and Aguirre Urreta, B. Miller, CN. and Lapaska, CA (1985). Two species of (1985). Plantas de la Formaci6n Kachaike, Cretacico Elatocladus from the Early Cretaceous Potomac Early Cretaceous macrofloras 63

Group of Virginia. Review of Palaeobotany and Roy, S.K. (1968). Pteridophytic remains from Kutch and Palynology 44: 183-191. Kathiawar, India. Palaeobotanist 16: 108-114. Moore, P.S. (1986). Jurassic and Triassic stratigraphy and Sah, S.C.D. (1952). Occurrence of Ginkgoales in the hydrocarbon potential of the Poolowanna Trough Rajmahal Series of Behar. Citrrent Science 21: 129-130. (Simpson Desert region) northern South Australia. Sah, S.C.D. (1953). On some species of Ginkgoites from Geological Society of Australia Special Publication 12: the Jurassic of the Rajmahal Hills, Bihar. 39-51. Palaeobotanist 2: 55-58. Morris, J. (1840). Appendix. In, C.W. Grant. Memoir to Sah, S.C.D. (1965). Palaeobotanical evidence on the age illustrate a geological map of Cutch. Transactions of of the Khatangi Beds (? Dubrajpur), in the Rajmahal the Geological Society, 2nd ser. 5: 289-329. Hills, Bihar. Palaeobotanist 13: 218-221. Morris, J. (1843). A catalogue of British fossils, comprising Sah, S.C.D. and Jain, KP. (1965).' Ginkgoites rajmahalensis all the genera and species hitherto described, with sp. novo from the Rajmahal Hills, Bihar, India. references to their geological distribution and to the Palaeobotanist 13: 155-157. localities in which they have been found. London. 222 Sah, S.C.D. and Sukh Dev. (1958). Thinnfeldia pp. chunakhalensis sp. novo from the Jurassic of the Morris, J. (1845). Fossil flora. In, P.E. Strzelecki Physical Rajmahal Hills, Bihar. Palaeobotanist 6: 22-24. description of New South Wales and Van Diemans Land. Sahni, B.- (1928). Revision of Indian fossil plants. Pt. 1. Longman, Brown, Green, and Longmans, London. Coniferales (a. Impressions and incrustations). 452 pp. Geological Survey of India, Palaeontographica Indica, MUnster, G.G. (1842-43). Beitrage zur Petrefacten-Kunde. Memoir, NS. 11: 1-49. Beyreuth. pt. 5: 1-131 (1842); pt 6: 1-100 (1843). Sahni, B. (1931). Revision of Indian fossil plants. Pt. n. Petriella, B. and Arrondo, O.G. (1984). La tafoflorula Coniferales (b. petrifactions). Geological Survey of Liasica de la Estancia la Juanita, Provincia de Santa India, Palaeontographica Indica, Memoir, NS. 11: 51­ Cruz, Argentina. Ameghiniana 21: 35-41. 124. Pigg, KB. (1992). Evolution of isoetalean lycopsids. Sahni, B. (1932). A petrified Williamsonia (W. Sewardiana Annals ofthe Missouri Botanic Gardens 79: 589-612. sp. nov.) from the Rajmahal Hills, India. Geological Piggott, A (1988). Ferns of Malaysia in colour. Tropical Survey of India, Palaeontographica Indica, Memoir, NS. Press, Kuala Lumpur. 458 pp. 20: 1-19. Playford, G. and Cornelius, KD. (1967). Palynological Sahni, B. (1948). The Pentoxyleae: a new group of and lithostratigraphic features of the Razorback Beds, Jurassic gymnosperms from the Rajmahal Hills of Mount Morgan District, Queensland. Papers of the India. Botanical Gazette 110: 47-80. Department of Geology, University of Queensland 6: 81­ Sahni, B. and Rao, AR. (1933). On some Jurassic plants 94. from the Rajmahal Hills. Journal and Proceedings, Playford, P.E., Cockbain, AE., and Low, G.H. (1976). Asiatic Society ofBengal (N.S.) 27: 183-208. Geology of the Perth Basin, Western Australia. Saporta, H. (1894). Flore fossile du Portugal. Academie Geological Survey of W.estern Australia Bulletin 124: 1­ Royale des..£ciences, Lisbon. 288 pp. 312. Schimper, W.P. (1869-70). Traite de paleontologie vegetale Rao, A.R. (1943). The stucture and affinities of ou la flora du monde primitif. J.B. Bailliere et fils., Paris Taeniopteris spatulata McCl. Proceedings of the National 1: 1-740 (1869); 2: 1-522 (1870). Academy of Science (India) 13: 333-355. Serra, C. (1963). Presence d'un Brachyoxylon rotnaensis Rao, A.R. (1981). The affinities of the Pentoxyleae. Mathiesen dans la flore Mesozoique du Bas-Laos. Palaeobotanist 28-29: 207-209. Compte Rendus du 88th Congres National des Societes Rees, P.McA (1993). Dipterid ferns from the Mesozoic Savantes 2: 469-482. of Antarctica and New Zealand and their Serra, C. (1966). Etude anatomique et paleogeographique stratigraphical significance. Palaeontology 36: 637-656. de quelques especies homoxylees du Sud Viet-Nam Retallack, G. (1977). Reconstructing Triassic vegetation et du Cambodge. Archives geologiques du Vi€t-Nam 8: of eastern Australasia: a new approach for the 59-131. biostratigraphy of Godwanaland. Alcheringa 1: 247­ Seward, AC. (1904). On a collection of Jurassic plants 277. from Victoria, Records of the Geological Survey of Retallack, G. (1980). Late Carboniferous to Middle Victoria 1: 155-211. Triassic megafossil floras from the Sydney Basin. Seward, A.C. (1917). Fossil Plants, 3. Pteridospermae, New South Wales Geological Survey Bulletin 26: 384­ Cycadofilicales, Cordaitales, Cycadophyta. Cambridge 430. University Press, Cambri.dge. 656 pp. Rich, T.H.V. and Rich, P.V. (1989). Polar dinosaurs and biotas of the Early Cretaceous of southeastern Seward, A.C. (1919). Fossil Plants, 4. Cambridge Australia. National Geographic Research 5: 15-53. University Press, Cambridge. 542 pp. Romerro, E.J. and Archangelsky, S. (1986). Early Seward, AC. and Sahni, B. (1920). Indian Gondwana Cretaceous angiosperm leaves from southern South plants: a revision. Geological Survey of India, America. Science 234: 1580-1582. Palaeontologia Indica, Memoir, n.s. 7(1): 1-55. Roy, S.K (1963). Studies on the Mesozoic flora of Kutch Shah, S.c. and Singh, G. (1964). Hausmannia crookshanki and Kathiawar - 1. Ptilophyllum horridum sp. novo sp. novo from Jabalpur Series of India. Current Science Proceedings of the 50th Indian Science Congress, Dehli 3: 33: 751-752. 396. Sharma, B.D. (1969a). Bucklandia dichotoma sp. novo from 64 s. McLoughlin

the Middle Jurassic of Rajmahal Hills, India. formations and their relative age. Palaeobotanist 36: Ameghiniana 6: 303-308. 161-167. Sharma, RD. (1969b). On a collection of fossil ferns from Sukh-Dev and Bose, M.N. (1974). On some conifer Dhokuti in the Rajmahal Hills, India. remains from Bansa, South Rewa Gondwana Basin. Palaeontographica 128B: 56-63. Palaeobotanist 21: 59-69. Sharma, B.D. (1971). Further studies on fossil Sukh-Dev and Zeba-Bano (1980). Some plant remains pteridophytic fronds collected from the Middle from Hoshangabad district, Madhya Pradesh, Jurassic rocks of Dhokuti in the Rajmahal Hills, Palaeobotanist 26: 206-213. India. Palaeontographica 133B: 61-71. Taylor, T.N. and Taylor, E.L. (1993). The Biology and Sharma, B.D. (1975): Additions to the Jurassic flora of Evolution of Fossil Plants. Prentice Hall, Englewood Dhokuti in the Rajmahal Hills, India. Acta Cliffs, New Jersey. 982 pp. Palaeobotanica 16: 83-100. Tenison-Woods, J.E. (1883). On the fossil flora of the Simpson, E.S. (1912). Unusual types of petrifaction from coal deposits of Australia. Proceedings of the Linnean Dandarragan. Journal of the Natural History and Society ofNew South Wales 8: 37-167. Science Society ofWestern Australia 4: 33-37. Towner, RR and Gibson, D.L. (1983). Geology of the Sitholey, R.V. (1984). An addition to knowledge of onshore Canning Basin, Western Australia. Bureau of Otozamites pecten Sahni and Sitholey. Palaeobotanist Mineral Resources, Geology and Geophysics, Australia, 32: 76-81. Bulletin 215: 1-51. Skwarko, S.K. (1968). Lower Cretaceous Trigoniidae Townrow, J.A. (1967). On Rissikia and Mataia from Stanwell, eastern Queensland. Bureau ofMineral podocarpaceous conifers from the lower Mesozoic of Resources, Geology and Geophysics, Australia, Bulletin southern lands. Papers and Proceedings of the Royal 80: 167-187. Society ofTasmania 101: 103-136. Skwarko, S.K. (1990). Jurassic fossils of Western Traves, D.M., Casey, J.N. and Wells, AT. (1956). The Australia. Geological Survey of Western Australia geology of the south-western Canning Basin, Palaeontological Report 1990116: 174 pp. Western Australia. Bureau of Mineral Resources, (unpublished). Geology and Geophysics, Australia, Report 38: 1-74. Smiley, CJ. (1970). Later Mesozoic flora from Maran, van de Graaff, W.J.E., Denman, P.D., Hocking, RM., Pahang, West Malaysia. Part 2: taxonomic and Baxter, J.L. (1980). Yanrey-Ningaloo, Western considerations. Bulletin of the Geological Society of Australia. Geological Survey ofWestern Australia, 1:250 Malaysia 3: 69-113. 000 Geological Series, Explanatory Notes. 24 pp. Smith, J.E. (1793). Memoires de 1'Academie Royale des Veevers, J.J., Powell, CMcA and Roots, S.R (1991). Sciences, Turin 5: 419. Review of seafloor spreading around Australia. 1. Spicer, R.A. and Parish, J.T. (1986). Paleobotanical Synthesis of the patterns of spreading. Australian evidence for cool north polar climates in middle Journal of Earth Sciences 38: 373-389. Cretaceous (Albian-Cenomanian) time. Geology 14: Visshnu-Mittre (1953). A male flower of the Pentoxyleae 703-706. with remarks on the structure of the female cones of Srivastava, RP. (1946). Silicified plant remains from the the group. Palaeobotanist 2: 75-84. Rajmahal series of India. Proceedings of the National Visshnu-Mittre (1957). Studies on the fossil flora of Academy of Science (India) 15: 185-211. Nipania (Rajmahal Series) India - Pentoxyleae. Sternberg, G.K. (1825). Versuch einer geognostischen Palaeobotanist 6: 31-46. botanischen Darstellung der Flora der Vorwelt. Leipzig. Vozenin-Serra, C (1977). Sur une penetration insolite de 1(4): 1-48. gymnospermes australes dans le sud-est asiatique au Stirling, J. (1892). Notes on the fossil florula and faunae Mesozoique. Compte Rendu Acad.de l'Academie des of the Gippsland carbonaceous area. Victorian Sciences, Paris 285(D): 1419-1421. Department ofMines, Special Report Coalfields 1: 10-13. Walkom, AB. (1915). Mesozoic floras of Queensland. Stubblefield, S.P. and Taylor, T.N. (1986). Wood decay Part 1. The flora of the Ipswich and Walloon Series. in silicified gymnosperms from Antarctica. Botanical (a.) Introdution. (b.) Equisetales. Geological Survey of . Gazette U7: 116-125. Queensland Publication 252: 1-51 Stubblefield, S.P. and Taylor, T.N. (1988). Recent Walkom, AR (1917a). Mesozoic floras of Queensland. Advances in palaeomycology. New Phytology 108: 3­ Part I.-continued. The flora of the Ipswich and 25. WaUoon Series. (c.) Filicales, etc. Geological Survey of Stubblefield, S.P., Taylor, T.N., and Beck, CB. (1985). Queensland Publication 257: 1-65. Studies of Paleozoic fungi. V. Wood decay in the Walkom, AB. (1917b). Mesozoic floras of Queensland. Upper Devonian progymnosperm, Callixylon Part I.-continued. The flora of the Ipswich and newberryi. American Journal of Botany 72: 1765-1773. Walloon Series. (d.) Ginkgoales, (e.) Cycadophyta, Sukh-Dev (1972a). Ferns from the Cretaceous of Madhya (f.) Coniferales. Geological Survey of Queensland Pradesh - 2 (Cladophlebis medlicottiana (Oldham) Publication 259: 1-49. Pascoe, Sphenopteris sp.). Palaeobotanist 19: 277-280. Walkom, AB. (1918). Mesozoic floras of Queensland. Sukh-Dev (1972b). Ferns from the Cretaceous of Madhya Part 2. The flora of the Maryborough (Marine) Series. Pradesh - 3 (Dipteridaceae - Hausmannia pachyderma Geological Survey of Queensland Publication 262: 1-21. n. sp.). Palaeobotanist 19: 281-283. Walkom, AB. (1919a). Mesozoic floras of Queensland. Sukh-Dev (1987). Floristic zones in the Mesozoic Parts 3 and 4. The floras of the Burrum and Styx Early Cretaceous macrofloras 65

River Series. Geological Survey of Queensland White, M.E. (1966). Report on 1965 plant fossil Publication 263: 1-77. collections. Bureau of Mineral Resources, Geology and Walkom, AB. (1919b). On a collection of Jurassic plants Geophysics, Australia, Record 1966/111. 1-10. from Bexhill, near Lismore, N.5.W. Proceedings of the (unpublished). Linnean Society ofNew South Wales 44: 180-190. White, M.E. (1981). Revision of the Talbragar fish bed Walkom, AB. (1921a). On the occurrence of Otozamites flora Gurassic) of New South Wales. Records of the in Australia with descriptions of specimens from Australian Museum 33: 695-721. Western Australia. Proceedings of the Linnean Society White, M.E. (1986). The Greening of Gondwana. Reed ofNew South Wales 46: 147-153. Books, Frenchs Forrest, N.S.W. 256 pp. Walkom, AB. (1921b). Mesozoic floras of New South Wilde, S.A and Low, G.H. (1978). Explanatory Notes on Wales. Part 1 - Fossil plants from Cockabutta the Perth Geological Sheet. Geological Survey of Mountain and Talbragar. Geological Survey of New Western Australia, 1:250 000 Geological Series ­ South Wales, Palaeontology, Memoir 12: 1-21. explanatory notes. Perth, Western Australia. 36 pp. Walkom, A.B. (1928). Fossil plants from Plutoville, Cape Wiltshire, M.J. (1989). Mesozoic stratigraphy and York Peninsula. Proceedings of the Linnean Society of palaeogeography, eastern Australia. In, The Cooper New South Wales 53: 145-150. and Eromanga Basins, Australia. B.J. O'Neil (ed.). Walkom, AB. (1944). Fossil plants from Gingin, W.A. Petroleum Exploration Society of Australia, Society Journal of the Royal Society of Western Australia 28: of Petroleum Engineers, Australian Society of 201-207. Exploration Geophysicists, Adelaide. pp. 279-291. Yeates, A.N., Crowe, RW.A., Towner, RR, Wyborn, Weaver, L., McLoughlin, S., and Drinnan, AN. (in L.AI., and Passmore, V.L. (1984). Regional geology press). Fossil woods from the Upper Permian Bainmedart Coal Measures, northern Prince Charles of the on-shore Canning Basin, W.A In, The Canning Basin, W.A. P.G. Purcell (ed.), Geological Society of Mountains, East Antarctica. AGSO Journal of Australia and Petroleum Exploration Society of Australian Geology and Geophysics. Australia, Perth. pp. 23-55. White, M.E. (1961a). Plant fossils from the Canning Zeba-Bano (1980). Some pteridophytes from the Jabalpur Basin, Western Australia. Appendix 6, In, The geology Formation. Palaeobotanist 26: 237-247. of the Canning Basin, Western Australia. J.J. Veevers and A.T. Wells. Bureau of Mineral Resources, Geology Zeba-Bano, Maheshwari, H.K. and Bose, M.N. (1979). and Geophysics, Australia, Bulletin 60: 291-320. Some plant remains from Pathargama, Rajmahal Hills, Bihar. Palaeobotanist 26: 144-156. White, M.B. (1961b). Report on 1960 collections of Mesozoic plant fossils from the Northern Territory. Bureau of Mineral Resources, Geology and Geophysics, Manuscript received 21 August 1995; accepted 1 Februanj Australia, Record 19611146: 1-26. (unpublished). 1996.