Upper Palaeozoic floras of SE Asia 73 Upper Palaeozoic floras of SE Asia

J F Rigby School of Natural Resource Sciences, Queensland University of Technology, Box 2434, GPO Brisbane, 4001, Australia

Key words: SE Asia, floras, Upper Palaeozoic, Cathaysialand, Gondwanaland

Abstract Edwards (1948) reported Lepidodendron and Stigmaria from Pahang, West Malaysia (Fig1, Small Carboniferous floras of limited significance occur in near locality 6) Edwards (1926) reported Thailand and West Malaysia They are related to the Pecopteris sp cf P+ cyathea (Schlotheim) pantropical Cathaysian and/or Laurasian floras Extensive Permian floras are known from Thailand, Laos, West Malay- Brongniart 1833 and Cordaites sp from the sia, Sumatra and Irian Jaya and these floras were substan- Raub Series of Kelantan (NE state of West tially related to Cathaysian floras However, some floras Malaysia) from Irian Jaya and Thailand include one or more species of Asama (1973) described two Lower Carbonif- the Gondwanaland genus Glossopteris When the distribu- erous (Visean) floral localities from Pahang, also tions are plotted on a reconstruction for SE Asia prepared by Carey, requirements are met for proximity to centres for ra- within the Raub Series, which he named the diation of both Cathaysian and Gondwanan floras Kuantan flora His Locality A at Panching (NW of Kuantan, Fig1, loc 6) yielded only lycopods: Lepidodendron acuminatum Goeppert 1847; Introduction Bergeria sp; Lepidodendropsis vandergrachtii Jongmans, Gothan and Darrah 1937; Stigmaria Upper Palaeozoic floras have been described sp His Locality B at Gambang (SW of Kuantan) from SE Asia The variety and distribution of yielded fern-like foliage and a seed: Rhodea these floras suggest that more occurrences may hsianghsiangensis Sze 1953, ?Adiantites sp, be found in the future as localities are rare, and ?Neuropteris sp, Carpolithus sp spread over a very large area which has not Asama et al (1975) included some additional been fully explored geologically Permian floras specimens from Trengganu (state adjoining have palaeogeographical significance for places Pahang to the north) and Pahang referring the outside the area because of the association of identifications to Franks, but omitting the refer- Cathaysian and Gondwanan species and the ence: Lepidodendron sp, Rhodea sp, Spheno- presence of endemic species phyllum sp They compared the Kuantan flora to the floras of the Kaolishan Formation of Jiangsu and the Tseshui Formation of Hunan, Carboniferous floras both in the Visean of southern China, in Wu (1995) I consider the data to be somewhat too Despite the presence of extensive Carbonifer- generalized for precise correlation, however the ous floras in neighbouring Yunnan and Guangxi, Early Carboniferous age is without doubt China, they are not well represented in SE Asia Laveine et al (1993) described some plants This may reflect the smaller number of geologi- from the Na Duang coal mine area of NE Thai- cal investigations undertaken in parts of SE Asia land (Fig1, loc 2) which they considered as

Biogeography and Geological Evolution of SE Asia, pp 73-82 Edited by Robert Hall and Jeremy D Holloway © 1998 Backhuys Publishers, Leiden, The Netherlands 74 J+ F+ Rigby

Fig1 The distribution of localities throughout SE Asia 1 Northern Laos: Nam-Ou (northerly), Phong-Saly (central), Bun-Tai (southern) The position of other localities in Laos is not known 2 Na Duang 3 Loei 4 Phetchabun 5 Jengka 6 Kuantan 7 Linggiu 8 Jambi 9 Rigby’s 1997 localities, Irian Jaya 10 Jongmans’ 1940 localities, Irian Jaya The quality of the locality data is variable, often based on data supplied by field workers

representative of the Middle Carboniferous Permian floras pantropical Paripteris flora Their Fig2 shows the flora’s distribution Their provisional species Floras from West Malaysia, Sumatra and Thailand list included: Stigmaria ficoides Brongniart, Lepidodendron sp, arborescent lepidophytes Extensive Permian floras occur in many parts of with large ulodendroid scars, ?Bothrodendron SE Asia Asama et al (1975, table 2) have assem- sp, Archaeocalamites sp, Rhodeopteridium sp bled distribution data from occurrences in Ma- cf R+ tenuis (Gothan) Zimmermann, ?Rhodeopte- laya, Sumatra and Thailand Table 1 is an edited ridium sp, Adiantites sp cf A+ spectabilis Read, version of some of their data which extend over Eusphenopteris sp cf E+ hollandica (Gothan 8 pages Their table includes distribution within and Jongmans) Novik, ?Cordaites sp This asso- the Cathaysian Permian and comparison with ciation confirms that northern Thailand lay extra-Chinese floras as well as the 66 species within the Carboniferous tropics occurring only at localities near Jambi (Djambi) Upper Palaeozoic floras of SE Asia 75 in Sumatra, which have been listed separately in identified Dadoxylon roviengense Vozenin-Serra Table 2, below Permian floras from Irian Jaya 1985 and D saxonicum (Goeppert) Frentzen 1931, (Western New Guinea) which are not closely both of which lacked pronounced growth rings related to the floras listed in Tables 1 and 2 are She noted that this characteristic was typical of discussed separately under ‘Permian floras of trees growing under a tropical or subtropical re- New Guinea’ gime

The Jambi flora Upper Permian floras from Thailand

Many fossil plant localities were discovered in Kon’no (1963) described a small flora from the Jambi district in central Sumatra, Indonesia Khlong Wang Ang, about 50 km SSW of during the Dutch administration An earlier suite Phetchabun (see Fig1, locality 12, in Iwai et al, of almost 200 slabs collected by A Tobler was 1966, for location data), identified from “five described by Jongmans and Gothan (1925) A small pieces of black shales” He commented second, very much larger suite of 487 slabs was that the specimens were “not very favourable in collected by O Posthumus, and also described preservation” His identifications included: by Jongmans and Gothan (1935) Table 1 lists Taeniopteris hallei Kawasaki 1931, T+ sp cf T+ the species identified from Jambi also occurring serrulata Kawasaki (non Halle) (both these spe- elsewhere in SE Asia, whereas Table 2 lists spe- cies are present in the approximately Kungurian cies identified only from Jambi A map showing Upper Jido Series in Korea); Sphenophyllum the positions for the collecting localities spread trapaefolium Stockmans and Matthieu 1957 over an area of 30 x 17 km was given by (Kon’no’s photo (Plate 8, Fig5) suggests the Jongmans and Gothan (1935, map 1) These au- specimen was distorted: it occurs in the approxi- thors have assembled the localities into six re- mately Kazanian Kobosan Series of Korea); gions, generally along stream valleys within the Palaeovittaria parvifolia Kon’no 1963; area, listing the species distribution Glossopteris sp cf G+ angustifolia Brongniart Most of these regions contributed floras of 20 1828 (the figured fragment is small, but has the or more species with most occurring in more distinctive glossopterid venation) The remain- than one region Even if one re-identifies speci- ing species were endemic: Bowmanites sp, mens into more broadly defined ‘species’, the Alethopteris thailandica Kon’no 1963, and total flora is still very large and varied The most Poacordaites phetchabunensis Kon’no 1963 striking features are the considerable variety of Asama (1966) subsequently collected from ferns and fern-like foliage, the variety of the same locality identifying specimens as Taeniopteris spp, and the number of Cordaites Sphenophyllum phetchabunensis Asama 1966, spp These indicate a Cathaysian affinity Asama Pecopteris sp, Compsopteris wongii (Halle) et al (1975) have also demonstrated a strong Zalessky 1934 (= Protoblechnum wongii), Laurasian affinity for the total flora The first col- Taeniopteris thailandica Asama 1966, T+ lection (Jongmans and Gothan, 1925) had affin- nystroemlii Halle 1927, T+ konnoi Asama 1966, ity only with Laurasia, lacking any Cathaysian T+ hallei Kawasaki 1934, T+ iwaii Asama 1966, species There was no suggestion of correlation Taeniopteris sp, Cordaites principalis (Germar) with coeval floras of Gondwanaland Geinitz 1855, Poacordaites linearis Grand’Eury Jongmans and Gothan dated the earlier col- 1877 (Asama included Kon’no’s P+ phetchabunensis lection (1925) as Carboniferous based on the as a synonym), Psygmophyllum komalarjunii flora’s apparent Laurasian aspect The much Asama 1966, Psygmophyllum? sp, Samaropsis sp larger, second collection (1935) also appeared Asama (1966) drew attention to aspects of the to be more Carboniferous than Permian, but a flora, including the variety of the Taeniopteris limestone underlying the two plant horizons species present, 3 large and 3 small leaved spe- yielded fusulinids dating the limestone as Mid- cies Taylor and Taylor (1993, p599) have ob- dle Asselian to Sakmarian, most probably Late served “In the Carboniferous (Pennsylvanian of Asselian (Vachard, 1989) Kansas), Taeniopteris foliage has been found Brouwer (1931) quoted Tobler as listing the with small seeds attached to the abaxial surface, occurrence of Dadoxylon above the plant-bear- suggesting affinities with the cycads In the ing horizons Vozenin-Serra (1985) studied Mesozoic, however, the taxon has been associ- some woods from Jambi that may have included ated with the bennettitalean Williamsoniella” It specimens from the earlier collections She is possible that the two groups of Taeniopteris 76 J+ F+ Rigby

Table 1 Distribution of Permian floras in Malaya, Sumatra and Thailand modified and edited from Table 2 of Asama et al (1975) Additional species occurring only at Jambi (Djambi), Sumatra are listed on Table 2 I consider there are taxonomic differences between the Cathaysian and Gondwanan specimens currently identified as Trizygia speciosa *

Jambi Loei Phetchabun Jengka Linggiu flora flora flora flora flora Sumatra Thailand Thailand W Malaysia W Malaysia

LYCOPHYTA Lepidodendron sp cf L chosenense Kawasaki + NOEGGERATHIOPHYTA Tingia subcarbonica Kon’no and Asama + SPHENOPHYTA Sphenophyllum trapaefolium Stockmans and Mathieu + Sphenophyllum sp + Parasphenophyllum phetchabunense (Asama) Asama + Trizygia oblongifolia (Germar and Kaulfuss) Asama + + T sinocoreana (Yabe) Asama + *T speciosa Royle + Paratrizygia glossopteroides forma minor (Kawasaki) Asama + P koboensis (Kobatake) Asama + + Calamites suckowii Brongniart + cf Annularia shirakii Kawasaki + + Lobatannularia fujiyamae Kon’no and Asama + L johorensis Kon’no and Asama + + L johorensis subsp minor Kon’no and Asama + L suntharalingamii Kon’no and Asama + FERNS AND FERN-LIKE FOLIAGE Ptychocarpus malaianus Kon’no + Ptychocarpus sp + Rajahia (Pecopteris) bifurcata Kon’no + R linggiuensis Kon’no + R pseudohemitenoides Kon’no + R rajahii Kon’no + R sengensis Kon’no + Shirakiopteris loeiensis Asama + Pecopteris arcuata Halle + + + P hemitenoides Brongniart + + P lativenosa Halle + P yinii Kon’no and Asama + Pecopteris sp + + CIadophlebis ozakii Yabe and Oishi + Fascipteris sinensis (Stockmans and Mathieu) Gu and Zhi + Aphlebia sp + + Neuropteris sp ++ Neuropteridium yokoyamae Kon’no and Asama + + Alethopteris thailandica Kon’no + Alethopteris sp + Protoblechnum wongii Halle + + cf Cathaysiopteris whitei (Halle) Koidzumi cf Taeniopteris crassicaulis Jongmans and Gothan + + T hallei Kawasaki + + + T iwaii Asama ++ T konnoi Asama + T latecostata Halle + + T multinervis Weiss cf + cf T nystroemii Halle + + + + T shansiensis Halle + T taiyuanensis Halle + + T thailandica Asama + cf Taeniopteris sp cf T serrulata Kawasaki + Taeniopteris sp ++ Gigantonoclea lagrehi (Halle) Koidzumi + + Bicoemplectopteris hallei Asama + + + Tricoemplectopteris taiyuanensis Asama + Bicoemplectopteridium longifolium (Kodaira) Asama + + Gigantopteris nicotianaefolia Schenk + + Upper Palaeozoic floras of SE Asia 77

Table 1 Continued

Jambi Loei Phetchabun Jengka Linggiu flora flora flora flora flora Sumatra Thailand Thailand W Malaysia W Malaysia

Glossopteris sp cf G angustifolia Brongniart + Palaeovittaria parvifolia Kon’no + Psygmophyllum komalarjunii Asama + Psygmophyllum sp ? Psaronius johorensis Ogura + CORDAITES Cordaites principalis (Germar) Geinitz + + C schenkii Halle + C simplicinervius Jongmans and Gothan + cf Poacordaites linearis Grand’Eury + Poacordaites sp + + Cordaianthus volkmannii (Ettingshausen) Zeiller cf Cordaicarpus cordai forma elongata Jongmans and Gothan + + CYCADOPHYTA Sphenozamites sp + GINKGOPHYTA Rhipidopsis baieroides Kawasaki and Kon’no + SEEDS Carpolithus sp + + Gigantospermum posthumii Jongmans and Gothan + + Samaropsis sp + + Trigonocarpus sp + +

Table 2 Permian plants whose only SE Asian occurrence is in the Jambi district of Sumatra The authorship of species designated as J and G was by Jongmans and Gothan 1935; most or all of these were endemic See Table 1 for species occurring both at Jambi and elsewhere in SE Asia

LYCOPHYTA Lepidodendron mesostigma J and G, L molle J and G, L posthumii J and G, Stigmaria asiatica J and G, S ficoides Brongniart, ?Lycopodites sp, Maroesia rhomboidea J and G

SPHENOPHYTA Sphenophyllum sp cf S emarginatum Brongniart, S verticillatum (Schlotheim) Brongniart, Parasphenophyllum thonii (Mahr) Asama, Sphenophyllostachys sp, Calamites jubatus Lindley and Hutton, Annularia stellata (Schlotheim) Wood, Annularia sp, Palaeostachya incrassata J and G

FERNS AND FERN-LIKE FOLIAGE Asterotheca sp, Monocarpia posthumii J and G, Sphenopteris zwierzycki J and G, S sp cf S gothanii Halle, S sp cf S grabaui Halle, S sp cf S matheti Zeiller, S sp cf S tingii Halle, Sphenopteris sp, Pecopteris arborescens Schlotheim, P candolleana Brongniart, P densifolia Goeppert, P djambiensis J and G, P mengkarangensis J and G, P oreopteridia Schlotheim, P polymorpha Brongniart, P unita Brongniart, P unitaeformis J and G, P verbeekii J and G, P sp cf P cistii Brongniart, P sp cf P daubreei Zeiller, Aphlebia ?acanthoides Zeiller, A dimorpha J and G, A minor J and G, A sp cf A crispa Gutbier, Neuropteris sp cf N gleichenoides (Stur) Sterzel, Nemejcopteris feminaeformis (Schlotheim) Barthel, Cyclopteris sp, Alethopteris strictinervis J and G, Macralethopteris hallei J and G, Callipteridium mengkarangense J and G, C sumatranum J and G, Dictyocallipteridium sundiacum J and G, Asterophyllites sp, Palaeogoniopteris mengkarangensis (J and G) Koidzumi, Taeniopteris camptoneura J and G, T densissima Halle, T incrassata J and G, T sp cf T norinii Halle, Gothanopteris bosschana (J and G) Koidzumi

CORDAITES Cordaites lingulatus Grand’Eury, Cordaianthus sp, Cordaicarpus cordai Geinitz, Cordaicarpus ovalis J and G, Cordaicladus sp

SEEDS Artisia sp, Carpolithus coffeoides J and G, C granulosus J and G, C multigranosus J and G, Rhynchogonium permocarbonicum J and G, Schuetzia sp, Tobleria bicuspis J and G 78 J+ F+ Rigby spp at Phetchabun each belong to different gleichenoides from Jambi (in Jongmans and groups of plants Alternatively, the two groups Gothan, 1935), namely one pinnule from of species may represent variants from two Jengka, and a number from Linggiu being natural species as it is known from elsewhere smaller and having a difference in venation that Taeniopteris species can be quite variable which they think may eventually prove to spe- Psygmophyllum is present commonly here, and cifically significant also in many parts of Cathaysialand Neuropteridium yokoyamae — Linggiu is the Compsopteris wongii occurs only in Cathaysia- type locality; Jengka, one specimen identified land where it ranges from Kungurian to with reservation Kazanian Pecopteris arcuata — all specimens were identi- Asama (1966, pp194-196, Plate 1, Figs4-6) fied positively named some leaves as Sphenophyllum phetcha- Taeniopteris multinervis — the only specimen bunense Asama 1966 to which he suggested as- from Linggiu is “too incomplete” to add any- signment of leaves previously included in thing to a correct identification with this Zamiopteris glossopteroides forma minor by Laurasian species Kawasaki (1939) who had used this name for Bicoemplectopteris hallei — was abundant at some isolated leaves having a pronounced Linggiu, but rare at Jengka It is one of the index midrib One of Asama’s specimens has three species for the Cathaysian flora leaves attached to a stem I consider these Kon’no and Asama (1970) and Kon’no et al+ leaves to be gymnospermous conforming to the (1970) conclude both floras equate to Stage P 1 2 now understood, tufted habit of many Late of China, however the Linggiu flora was slightly Palaeozoic gymnosperms He also discussed the older The Jengka flora came from an interbed- presence of Glossopteris in Thailand suggesting ded marine/non-marine sequence dated as Late that the most probable explanation was that the Permian, having an age supported by faunas genus arose by parallelism An unfortunate con- The Linggiu flora was found in arenites associ- fusion has arisen from calling floras including ated with shallow seas Full data have been species from more than one phytological prov- given by Rajah (in Kon’no et al+, 1970) The en- ince ‘mixed floras’ In this case, a plant having vironmental difference is sufficient to account leaves of Glossopteris has grown alongside typi- for the difference in floral composition cal Cathaysian species, implying a mixture of the Cathaysian flora with the Gondwanan flora This could only occur if there were a land connection Permian floras of Laos between Cathaysialand and Gondwanaland across New Guinea in the Permian, also if the Many of the original references are not available climate at the time favoured the growth of the to me To overcome this, data have been taken tropical Cathaysian flora in the same place as the directly from Vozenin-Serra (1979) She identi- temperate Gondwanaland flora This would occur fied and figured specimens in the columns near the subtropical zone headed Sap-Pong and Laos Other references have not been seen by me, but citations have been extracted from Vozenin-Serra (1979) for Permian floras of West Malaysia completeness

Two floras from the Permian of West Malaysia were described within a few months of each Permian floras of New Guinea other in 1970, namely the Linggiu flora from Johore (Kon’no et al+, 1970) and the Jengka flora Permian floras occur in Irian Jaya (Western New from Pahang (Kon’no and Asama, 1970) The Guinea) along the southern or Australasian plate species occurring are listed on Table 1 Consid- side of the main suture zone All occurrences for ering the total number of species occurring in which I have data were found in the Lower each flora (Jengka = 20, Linggiu = 40) it is signifi- Permian Aiduna Formation The earliest pub- cant how few species they have in common (5) lished account of the fossil floras was by These authors’ discussion of the five species Jongmans (1940) who reported Sphenophyllum shows that even these may not have many fea- verticillatum, Pecopteris sp cf P+ arcuata, P sp tures in common cf P+ paucinervis, P+ sp cf P+ orientalis, P+ Neuropteris sp — Specimens in each collection unita, Taeniopteris sp cf T+ multinervis, T sp were compared with Neuropteris sp cf N+ cf T+ taiyuanensis and Vertebraria Other than Upper Palaeozoic floras of SE Asia 79

Vertebraria which was typical of Gondwana- largely fern or fern-like fronds and resembled land, most other species were typical of Carbon- floras from Jambi, however species differed iferous floras of both Cathaysialand and Hopping and Wagner (1962), and Rigby (1997) Laurasia Lehrner et al+ (1955) reported plant reported predominantly species of Glossopteris fossils from a number of localities in the Aiduna with some ferny foliage and having a superficial Formation, however I cannot comment as the appearance to floras from the warmer parts of paper is not referred to in other publications, Gondwanaland such as India The most signifi- and I do not have the citation, and therefore it is cant aspect of these floras was the recognition omitted from the references by Rigby of seed plants linking the region to Hopping and Wagner (1962) figured and both Gondwanaland (Glossopteris) and Cathay- briefly described species from the Aiduna For- sialand (Gigantonoclea) as there was no animal mation from a number of localities identifying vector available to transport seeds across the Sphenophyllum sp cf S+ speciosum, Pecopteris Tethys Sea which had an opening of about 45o monyi, Cladophlebis sp cf C+ australis, based on the more popular plate tectonics pal- ‘Validopteris’ sp, Glossopteris sp cf G+ aeogeographical reconstructions However, us- browniana, G sp cf G+ indica, G+ sp aff G+ ing a reconstruction based on the earth expan- retifera , Vertebraria sp, Taeniopteris sp cf T+ sion theory, the Tethys was a narrow seaway, hallei These species include both Gondwana- and sometimes dry land (Dickins, 1996) This land and Cathaysialand representatives reconstruction would have allowed terrestrial Rigby (1997) revised previous identifications contact between Irian Jaya and Cathaysialand and identified new collections from the Aiduna along which seed plants could have migrated Formation He noticed that Jongmans’ figured Carey (1996) has proposed a reconstruction of Taeniopteris spp showed evidence of occasion- the Permian land masses forming the SE Asia of al cross-veins in the secondary venation and today It is used as a basis for the reconstruction therefore more correctly were Glossopteris spp, herein (Fig2) The data have also been plotted also proposing G+ iriani and the Cathaysian- on a recent plate tectonic reconstruction (Fig3) related Gigantonoclea iriani both occurring on by Metcalfe (1996) the same rock slab Together with revisions of The only reference to Permian floras in Papua previous workers’ identifications, the total flora New Guinea is by Mackay and Little (1911) who included Trizygia speciosa (Sphenophyllum reported the occurrence of Glossopteris in coal speciosum in Hopping and Wagner, above); measures from central Papua New Guinea At three Pecopteris spp, fern frond, Ptychocarpus present the coal measures are considered to be sp, Cladophlebis sp, Fascipteris aidunae Tertiary, and this record is considered a mis- (‘Validopteris’ sp of Hopping and Wagner); identification Glossopteris iriani, eight other Glossopteris spp; Vertebraria indica; Gigantonoclea iriani; and Koraua hartonoi (probably a gymnosperm) In Conclusions the case of Trizygia speciosa there are problems with both the specimens referred to the species Carboniferous floras and their generic attribution Specimens from Cathaysialand and Gondwanaland differ, as Plant remains from West Malaysia are nonde- pointed out by Li and Rigby (1995), which script, however, they do suggest that the region means they probably should belong in different formed part of the same continental block that species The other problem is whether the included southern China The only floras of this species belongs in Sphenophyllum or in the general age that includes specifically identifiable Gondwanaland genus Trizygia as the type plants occurs in the Middle Carboniferous at the species Further work is necessary Na Duang coal mine of NE Thailand The flora The wood, Planoxylon stopesii, from the was within the distribution of the pan-tropical Aimau Formation of Vogelkop, Irian Jaya, was Paripteris flora described as showing characters of both araucarian and abietinian wood (Prasad, 1981), common wood types from the Upper Palaeo- Permian floras zoic of Gondwanaland (Marguerier, 1973) The significance of these floras is their ende- Permian floras are all tropical Cathaysian al- mism, i+e, constrained to species not found though some from Thailand include the Gond- elsewhere Jongmans’ (1940) flora comprised wanan genus Glossopteris The floras from Irian 80 J+ F+ Rigby

Fig2 Permian plant localities indicated on a Triassic recontsruction by Carey (1996, Fig44) based on the earth expansion theory The Late Permian palaeogeography of SE Asia was almost identical with the Triassic restoration as no major earth movements took place over this period The bold letters signify: south of a a a, typical Gondwanaland floras Transitional floras at Hazro, Turkey by h, and in Irian Jaya by n Pantropical floras north of the line joining t t, with the Thai and Laotian floras near p The Jambi flora of Sumatra occurs at s

Jaya (West New Guinea) are predominantly although it had more than 30 species known Gondwanan but with the addition of two only from Jambi All floras included a number of Cathaysian genera each with one endemic spe- fern species, often Pecopteris The floras were cies, viz Fascipteris aidunae and Gigantonoclea tropical, but where the Gondwanan species also iriani The large and varied flora occurring occurred, they may have been sub-tropical or around Jambi, Sumatra, had Cathaysian affinities warm temperate Upper Palaeozoic floras of SE Asia 81

Fig3 Distribution of Permian floras on a Late Permian restoration from Metcalfe (1996, Fig13, lower left) based on the plate tectonics theory Letters on the map are: A: Arabian peninsula, Au: Australia, C: Cimmerian continent, I: Indo Chinese region, In: India, L: Lhasa block, NC: Northern China, P: Pangea, SC: Southern China, WB: Western Burma Pure pantropical floras of SE Asia occurred on block I, these floras were related to the floras on block SC and P Floras combining species from both the pantropical region and Gondwanaland (In and Au) occurred on the southern margin of block I and on New Guinea Transi- tional floras at Hazro, Turkey by h, and in Irian Jaya by n The Thai and Laotian floras were situated near p The Jambi flora of Sumatra occurs at s

Acknowledgements Bulletin of the National Science Museum 9(2): 171-211 pls 1-6 Asama, K 1973 Lower Carboniferous Kuantan flora, Professor S W Carey, Department of Geology, Pahang, West Malaysia Geology and Palaeontology of University of Tasmania, Australia, for the base Southeast Asia 9: 109-118, pls XIV-XVI map on which the floral distribution has been Asama, K, Hongnusonthi, A, Iwai, J, Kon’no, E, Rajah, S plotted Professor Li X-X, Nanjing Institute of and Veeraburus, M 1975 Summary of the Carboniferous Geology and Palaeontology, Academia Sinica, and Permian plants from Thailand Geology and Palae- ontology of Southeast Asia 15: 77-101 China, for discussion concerning aspects of the Brouwer, H A 1931 De stratigraphie van Nederlandsch Permian Cathaysian floras Professor R Hall, Oost-indië 18 Paleozoic Leidsche Geologische Royal Holloway University of London, and my Mededeelingen 5: 552-566 referees for improvements in the text Carey, S W 1996 Earth, Universe, Cosmos Geology De- partment, University of Tasmania, pp xii + 231 Dickins, J M 1996 The southern margin of Tethys Ninth International Gondwana Symposium 2: 1125-1134 References Edwards, W N 1926 Carboniferous plants from the Malay States Journal of the Malayan Branch of the Royal Asiatic Asama, K 1966 Permian plants from Phetchabun, Thailand Society 4: 171-172, pl 1 and problems of floral migration from Gondwanaland Edwards, W N 1948 Lepidodendroid remains from Malaya 82 J+ F+ Rigby

Appendix 9 in H M Muir-Wood, Malayan Lower Carbon- Li X-X and Rigby, J F 1995 Further contributions to the iferous fossils British Museum (Natural History) London, study of the Qubu flora from southern Xizang (Tibet) pp 78-81, pl 10 Palaeobotanist 44: 38- 47 Hopping, C A and Wagner, R H 1962 Photographs of fos- Mackay, D and Little, W S 1911 The Mackay-Little Expedi- sils Enclosure 17 In W A Visser and J J Hermes Geo- tion in southern New Guinea Geographical Journal 38: logical results of the exploration for oil in Netherlands 483-487, 1 map, 2 pls New Guinea Koninklijk Nederlands Geologisch Marguerier, J 1973 Paleoxylologie du Gondwana africain Mijnbouwkundig Genootschap Geologische Serie, Etude et affinités du genre Australoxylon speciaal nummer 20 Palaeontographica Africana 16: 37-58 Iwai, J Asama, K, Veeraburas, M and Hongnasonthi, A Metcalfe, I 1996 Pre-Cretaceous evolution of SE Asian 1966 Stratigraphy of the so-called Khorat Series and a terranes In Tectonic evolution of Southeast Asia pp 97- note on the fossil plant-bearing Palaeozoic strata in Thai- 122 Edited by R Hall and D J Blundell Geological Soci- land Japanese Journal of Geology and Geography 37 (1): ety of London Special Publication 106 21-38 Prasad, M N V 1981 New species of fossil wood Jongmans, W and Gothan, W 1925 Beiträge zur Kenntnis Planoxylon from Late Paleozoic of Irian Jaya, Indonesia der Flora des Oberkarbons von Sumatra Verhandelingen Bulletin of the Geological Research and Development koninklijke nederlandse Geologisch Mijnbouwkundig Centre 5: 37-40 Genootschap 8: 231-287, 5 pls Rigby, J F 1997 The significance of a Permian flora from Jongmans, W and Gothan, W 1935 Die Ergebnisse der Irian Jaya (West New Guinea) containing elements re- paläobotanischen Djambi-Expedition 1925 2 Die lated to coeval floras of Gondwanaland and paläobotanischen Ergebnisse Jaarboek van het Cathaysialand Palaeobotanist 45: 295-302 Mijnwezen in Nederlandsch-indië 50: 71-201, pls 1-58 Taylor, T N and Taylor, E L 1993 The biology and evolu- Jongmans, W 1940 Beiträge zur Kenntnis der Karbonflora tion of fossil plants Prentice-Hall, Englewood Cliffs xxii von niederländisch Neu-Guinea Mededeelingen + 982 pp Geologische Stichting 1938-1939: 263-274, pls 1-3 Vachard, D 1989 A rich algal microflora from the Lower Kawasaki, K 1939 Addition to the flora of the Heian Sys- Permian of Jambi Province In The Pre-Tertiary fossils of tem Bulletin of the Geological Survey of Tyôsen 6 (5): 1- Sumatra and their environments pp 59-69 pls 6-9 Ed- 39, pls 1-9 ited by H Fontaine and S Gafoer Committee for co- Kon’no, E 1963 Some Permian plants from Thailand Japa- ordination of joint prospecting for mineral resources in nese Journal of Geology and Geography 34 (2-4): 139- Asian offshore areas Coop Technical Section, Bangkok 159, pl 8 Vozenin-Serra, C 1979 Étude de quelques empreintes de Kon’no, E, and Asama, K 1970, Some Permian plants from végétaux fossiles du paléozoïque supérieur du Laos the Jengka Pass, Pahang, West Malaysia Geology and Comptes Rendus du 104 Congrès national des Sociétés Palaeontology of Southeast Asia 8: 97-132, pls 17-24 savantes 1: 155-174 Kon’no, E, Asama, K and Rajah, S S 1970 The Late Vozenin-Serra, C 1985 Bois homoxylés du permien Permian Linggiu flora from the Gunong Blumut area, inférieur de Sumatra Implications paléogógraphiques Johore, Malaysia Bulletin of the National Science Mu- Actes du 110 Congrès national des Sociétés savantes Sec- seum 13(3): 491-580, pls 1-17 tion des sciences 5: 55-63 Laveine, J-P, Ratanasthien, B and Sithirach, S 1993 The Wu X-Y 1995 Carboniferous floras In Fossil floras of Carboniferous flora of Northeast Thailand, its China through the geological ages (English edition) pp paleogeographic importance Comptes Rendus des 78-126 Edited by Li X-X and others Guangdong Sci- Académie des Sciences de Paris 317 (2): 279-285 ence and Technology Press, Guangzhou