Distribution, evolution and extinction of global Early Carboniferous flora
Shaila Chandra & Sun Keqin
Chandra S & Sun Keqin 1996. Distribution, evolution and extinction of global Early Carboniferous nora. Paiaeoboianisl 43(2) : 88-97
The uniform Lepidodendropsis nora ofthe Early Carboniferous is uniformly and widely distributed in the Cathaysia, Laurasia, Kazakhstania and Gondwana continents. This is characterized by Lepidodendropsis, Sublepidodendron, Archaeosigtllaria, Archaeoc;alamiles, Sphenopleridium, Cardiopleridlum, Rhodeoplendium, Rhac:opleris, Trlphyllopteris and Adlanlites, etc. The global climatic differentiation was not very obvious during the Early Carboniferous, therefore, all Early Carboniferous plant assemblages belonged (0 the same phytogeographic province. II is known Ihal some Iypical genera of the Lepidodendropsis nora of the Early Carboniferous, such as Lepidodendropsis, Lepidosigillaria and Archaeoslgillaria are recorded from the Lower Devonian in Libya, Africa (Lejal-Nicol, 1975) and some elements of this nora are also recorded from the Late Devonian in China and other parts of the world. It is considered that this Lepidodendropsis nora gradually started appearing in the Early Devonian, began to diversify in the Late Devonian and reached its maximum development in the Early Carboniferous. The nora tended to decline in the late Early Carboniferous and resulted in its extinction in the end of the Early Carboniferous.
Although some plant elements in Cathaysia, Euramerica, Angara and Gondwana had already appeared in the Early Carboniferous, especially in lhe late Early Carboniferous, the occurrence of the new taxa was regarded as a result of gmdually increasing climatic differentiation and plant evolution. During the tr,msition period from Ihe Early Carboniferous to the Late Carboniferous, lhe differenliations of climatic conditions, tectonic movements, continental posilions, oceanic currenlS and glaciation were quite obvious, resulting in the extinclion of numerous lypical elemenls of the Early Carboniferous nora and the appearance ofsome forerunners of the Lale Carboniferous nOrdS in the world. The Cathaysia, Euramerica, Angara and Gondwana noras derived from the identical Lepidodendropsis nora of the Early Carboniferous, bUI developed and nourished in different environmental conditions.
Key-words--Lepidodendropsis nora, Distribution, Evolution, Extinction, Early Carboniferous
Shaila Chandra, Birbal SahnllnsllfUie of Palaeobolany. 53 Unlverslfy Road, Lucknow 226 007, India.
Sun Keqin, China Universlfy of Geosciences, Beijing 100 083, China. mmt ~ ~, ~ ~ (111"' .. _')" ~~lJ:Ci~~ ~ ~~, ~'Wll1, ~ ~ 3fIM ~~~, 1{{IARCflI, WTm-q:ci~~~~~'CflTfcfCflm~Cfll.,f;jlq,nl~itit ~ ~ ~ ~ ~ ~3TI~ ~ ~ ~,~ ~ 'CflTfcfCflm -q:ciqtmit fcfCflm it 31-ffimCfll.,f;jlq,Hl ~ ~ ~ "Wm '*'1~lllcncrtffi:~,fqqdRCfl "l'lfufcn~,I4W;J4111~, ~ q;H:qfd'iild~:i.rCfl~ ~~. -q:ci~~ it ~it ~ V:~ lh<:ifq{<'q Cfll.,f;jlq,n"l ~~it~(f~~~~~;JFl1ft1<:n1~PT(1~f ~ ~3TI~~ ~~ ~ it m~~, 1{{IARCflI, WTm -q:ci q'1fqfd'iiIJi 'CflTfcfCflm QllTq{uRll it fcrcf;fuo SQ:~I THE term Lepidodendropsis flora was proposed by (. Rhodea)Tnphy/iopteris + + + Jongmans 0952, 1954), which dealt with a cos Rhacoptens + + + mopolitan flora of world-wide extent of the Early Card/optens + + Carboniferous. Lutz (933) established Lepidoden (- Fryopsis) dropsis as a new genus from the Lower Carboniferous Diplupleridium + of Geigen near Hof, Bavaria of Germany. P:.ygmophy/lum + lepidodendropsisis characterized by having long nar Archaeopterldium + row fusiform leaf cushions arranged in close spirals, Ad/anlites + + + + forming pseudowhorls bearing false leaf scars, lack Ane/m/les + + ing leaf abscission and a ligule pit (Lutz, 1933; Lacey, Angaropteridium + 1962; Mensah & Chaloner, 1971). Lepidodendropsisis known as the representative genus of lycopods and Cathaysia: Chang Shanchen (195(i), Deng Bao (1978), Zhang Shanzhen et al. (1980), WU Xiuyuan & Zhao Xiuhu is widely distributed in many parts of the world. Other (981), Feng Shaonan et al. (1982), Wu Xiuyuan important genera of the Early Carboniferous are Sub (992), Chen Fen et al. (1994), Chen Fen et al. (1995) lepidodendron, Archaeosigillaria, Archaeo Euramerica: Lutz (1933),Jongmans et al. (1937), Read & Mamay calamites, Sphenopteridium, Cardiopteridium, (19(i4), Lacey 09(i2), Cleal & Thomas (1991). Rhodeopteridium, Triphyllopteris, Rhacopteris, Angdra : Vakhrameev et al. (1970), Chaloner & Lacey (1973), Adiantitesand Aneimites, etc. and together constitute Chaloner & Meyen (973), Cleal & Thomas (1991) Lepidodendropsis flora (see Table 1). Gondwana: Jongmans (954), Hoeg et al. (1955), Pal & Chaloner (1979), Singh et al. (J 982), Cleal & Thomas (1991), Table 1 -Distribution of some important genera of Early Car Pant & Srivastava (995). boniferous flora Genera Cathaysia Euramerica Angard Gondwana DISTRIBUTION OF GLOBAL EARLY (China) CARBONIFEROUS FLORA Sublepidodendron + + + + Eolepldodendron + + + + The Lepidodendropsis flora of Early Car Siberlodendron + boniferous is known from China, Malaysia, the Lophlodendron + United States of America (Arkansas, Utah, Illinois, Pseudobumbudendron + + Pennsylvania, Virginia), Nova Scotia of Canada, Spondylodendron + Bavaria of Germany, France, North Wales of Great Archaeoslgillarla + + Britain, Spain, Poland, Ireland, Greenland, Donetz, Lepidosigtllarla + Kazakhstan, Siberia, Mangolia, Syria, Brazil, Ghana, Lepidophloios + Morocco, Niger, Libya, Sahara, Egypt, Kashmir, Peru, Cyclostigma + Australia and Argentina, etc. (Text-figure 1). The ArchaeocalamItes + + + Early Carboniferous flora of the world can be recog Sphenopteridlum + + + nized as belonging to four broader areas namely the Cardloplerldlum + + + Cathaysia, Angara, Euramerica and Gondwana areas Rhudeoplerid/um + + + on the basis of floral similarity. 90 THE PALAEOBOTANIST .c ,.. G~u P~ /1~ O' ~~ , IP;.. ~ <0>' '0. • • j" "i". '_' ". • • I q#, The Cathaysia area sp., etc. Lepidodendron, Lepidostrobophyllum and Lepidostrobusare also common genera of the Lower The Early Carboniferous flora has been recorded Carboniferous in the Euramerica area. Zhang from a number of localities in China and belong to Shanzhen et al. (980) considered that the flora the typical Lepidodendropsis flora. Chang Shanchen showed a close relationship with the Visean or Culm (956) described Tnphyllopteris collumbiana flora of West Europe. Wu Xiuyuan and Zhao Xiuhu Schimper, Cardiopteridium spitsbergense Nathorst (981) studied fossil plants from the Koalishan For and Sphenopteris (Rhodea?) sp. from the Lower Car mation of Lower Carboniferous in ]urong, southern boniferous in -::astern Kansu CGansu), China. Triphyl ]iangsu of China, including Sublepidodendron lopteris collumbiana and Cardiopteridium spitsber mirabile CNath.) Hirmer, S. d. mirabile CNath.) gense are the characteristic elements of the Hirmer, Eolepidodendron jurongense Wu & Zhao, Lepidodendropsis flora and the species of world Eolepidodendron sp., Lepidodendrongaolishanense wide occurrence. Deng Bao (978) recorded fossil Wu & Zhao, Stigmaria ficoides CSternberg) plants from the Lower CarboniferolIs in Shanyang, Brongniart, S. rugulosa Gothan, Rhodeoptericl.iumcf. southern Shanxi CShaanxi) as Archaeocalamites hsianghsiangense CSze), Rhodeopteridium sp., scrobiculatusCSchlotheim) Seward, Cardiopteridium Telangium sp. and Hamatophyton verticillatum Gu spetsbergense Nathorst, Triphyllopteris collumbiana & Zhi, etc. The flora indicates Late Tournaisian to Schimper, Cardiopteris? spp. and Lepidodendron Early Visean and represents a typical Lepidodendrop shanyangenseWu & He, etc. Zhang Shanzhen et at. sis flora. Wu Xiuyuan (992) described fossil plants (980) described the Early Carboniferous flora from from the Yangshan Formation of Lower Car the Tseshui Series in Shuangfeng of Hunan compris boniferous in Gushi, Henan of China, such as ing Cardiopteridiumspitsbergense Nathorst, Triphyl Lepidodendronaff. aolungphyllukenseSze, L cf. wor lopten's collumbiana Schimper, Adiantites gothani thenii Lesquereux, 1. d. shanyangense Wu & He, 1. (Sze), Rhodeopteridium d. hsiangh'siangense CSze), dabieshanense Wu, 1. ?gushiense Wu, Archaeocalamites prolixus Zhang, Zhao & Wu, Lepidodendron sp., Cathaysiodendrori? sp., Lepidodendron d. robertii Nathrost, Lepidodendron Bothrodendron yangshanense Wu, B. Jlabellatum spp., Lepidostrobophyllum spp. and Lepidostrobus Wu, Archaeocalamites scrobiculatus CSchlotheim) CHANDRA & SUN-DISTRIBUTION, EVOLUTION AND EXTINCTION OF EARLY CAR130NIFEROUS FLORA 91 Seward, Cardiopteridium podozamioides (Sze), Tri Bunbury, R. semicircularis Lutz, Archaeopten'dium phyllopteris gushiensis Wu, Triphyllopteris sp., dawsoni Stur, Sphenopteridium pac.hYIThachis Rhodeopteridium hsianghsiangense (Sze), Adian Goeppert, S. dissectum Goeppert, SphenopteriS tites d. gothanii (Sze), Eusphenopteris d. scribanii foliolata Stur, Adiantites tenuifolius Goeppert, Amerom and Neuropterissp., etc. The age of the flora Calathiops d. plauensis Gothan, Telangium sp., is regarded as Visean to Early Namurian. The above Rhodea patentissima En., R. lemayi Bertrand & mentioned elements show that some forerunners of Broussier and R. d. hochsteteri Srur, etc. Obviously, the Cathaysia flora had already appeared in the late this flora contains some important elements of Early Early Carboniferous, belonging to a number of orien Carboniferous flora and hence regarded as a typical tal species oflycopods. Chen Fen et al. (995) studied Lepidodendropsis flora. Lutz (933) established the fossil plants from the Carboniferous strata of Lepidodendropsis as a new genus with only one Ningxia and adjacent regions, China. Among the fos species, namely, Lepidodendropsis hirmeri Lutz. sil plants, some elements from Early Namurian strata Later, a number of species of Lepidodendropsis were (Namurian A) were recorded as Lepidodendron described from the Lower Carboniferous Pocono For ninghsianense Sze & Lee, L. subrhombicum Gu & mation in Pennsylvania and Virginia Oongmans et al., Zhi, Mesocalamites cistiformis Stur, Tingia trilobata 1937). Lacey and Eggert (964) described fossil plants Stockmans & Mathieu, Cardiopteris ningxiaensis from the Chester Series (Upper Mississippian) of Chen, Sun & Zhou, Rhodeopteridiumsp., Eusphenop teris d. obtusiloba (Brongniart), Paripteris gigantea southern Illinois, namely Archaeocalamites radiatus (Sternberg) Gothan, P. otozamioides(Sze & Lee) and (Brongniart) Stur, Lepidodendron volkmannianum Linopteris simplexGu & Zhi, etc. The genus Tingia is Sternberg, L. cf. veltheimii Sternberg, Stigmaria regarded as an important element of the Cathaysia ficoides (Sternberg) Brongniart, Rhynchogonium flora which first appeared in the lower part of the jayettevillenseWhite and Trigonocarpussp. Read and Namurian (Namurian A). Mamay (964) recognized three floral zones from the In China, the studies of the Early Carboniferous Mississippian (Lower Carboniferous) in the United flora have been further developed by Feng Shaonan States, namely, zones of Adiantites spp., Triphyllop et al. (982), Zhao Xiuhu et al. (982) and MiJiarong teris spp. and Fryopsis spp. which contain Adiantites et al. (990). spectabilis Read, Lepidodendropsis scobiniformis Asama (973) described Lower Carboniferous (Meek) Read, Sphenopteridium brooksi Read, fossil plants from Panching, West Malaysia, namely Rhodea vespertina Read, Rhacopteris latifolia (Ar Lepidodendron acuminatum Goeppert, nold) Read, Triphyllopteris lescuriana (Meek) Lepidodendropsis vandergrchtiJongmans, Gothan & Lesquereux, T. varineruis Read and Fryopsis abdpnsis Darrah, Bergeria sp., Stigmaria sp., Rhodea (Read) Wolf, etc. This sequence is probably incom hsianghsiangensis Sze, .?Adiantites sp., .?Neuropteris plete, because the American Mississippian is pre sp. and Carpolithus sp. dominantly marine (Read & Mamay, 1964). They The Early Carboniferous flora of the Cathaysia have pointed out that the Upper Devonian flora was area is mainly distributed in China and Malaysia and not only mere diverse but was more similar to the consists of some typical elements of the Mississippian flora that follows than the Lower Lepidodendropsis flora. Some forerunners of the Devonian flora. Cathaysia flora are known to occur in late Early Lacey (962) described Early Carboniferous fossil Carboniferous strata (Sun Keqin, 1996). plants from the Lower Brown Limestone in the Vale The Euramerica area of Clwyd, North Wales, namely Archaeocalamites radiatus (Brongniart) Srur, Lepidodendron pelfo Lutz (1933) described fossil plants from the Lower ratum Lacey, Lepidostrobophyllum timbriatum Carboniferous of Geigen near Hof, Bavaria of Ger (Kidston), Lepidodendropsis jonesi Lacey, L. recur many, namely Lepidodendropsis hirmeri Lutz, Lepidostrobus d. faudelii Schimper, Asterocalamites vi/olia Lacey, Stigmariasp., Archaeosigillaria stobbsi scrobiculatus Schlorheim, Sphenophyllum Lacey, Clwydia decussata Lacey, Rhacopteris sub saxifragaefolioides Leyh, S. geigense Lutz, Cardiop cuneata Kidston, R. d. subcuneata Kidston, R. weis teris frondosa Goeppert, Rhacopteris lindseaeformis siiWalton, R. d. weissiiWalton, R. d. geikiei Kidston 92 THE PALAEOBOTANIST and Calathiops dyserthensis Lacey, etc. The flora is known from Spiti, N.W. Himalaya (Gothan & Sahni, probably of the Middle to Late Visean (Lacey, 1962). 1937; Hoeg et al., 1955) and Kashmir Himalaya (Pal, The above mentioned fossil plants represent 1978; Pal & Chalone 1', 1979; Singh et aI., 1982; Pant & some of the best documented assemblages from the Srivastava, 1995). Gothan and Sahni (937) described Lower Carboniferous in the Euramerica area and fossil plants from the Po Series of Spiti, namely exhibit a remarkable floral uniformity. Rhacopteris ovata (McCoy) Walkom, Spheno phyllum?furcillatum Ludwing and Sphenopteris sp. The Angara area Another collection of fossil plants was made from the Po Series of Spiti, northwest Himalaya (Hoeg et aI., The Early Carboniferous flora of the Angara area 1955) and contained Rhacopteris ovata (McCoy) is characterized by the presence of Lepidodendropsis, Walkom, R. inaequilatera Goeppert, R. cf. circularis Sublepidodendron, Lophiodendron, Ar Walton, Rhacopten's sp. a, Rhacopteris sp. b, chaeocalamites, Chaccassopteris and Angaropte Sphenopteridium sp. a, Sphenopteridium sp. b, ridium(Chaloner & Lacey, 1973). Cleal and Thomas Rhodea sp. a, ?Rhodea sp. b, Sphenopteris sp., (991) listed some typical genera of the ?Asterophyllites sp., ?Adiantites sp. a and ?Adiantites Lepidodendropsis flora from Kuznetsk of the Angara sp. b. Pal and ChaloneI' (979) recorded fossil plants area, such as Lepidodendropsis, Archaeocalamites, from the Gund Formation of Lower Carboniferous in Cardiopten'dium and Adiantites. They also listed a Kashmir, namely, Lepidodendropsis d. sigillarioides number of characteristic forms of the Lepidodendrop Jongmans, Gothan & Darrah, L. cf. fenestrata sis flora namely, Lepidodendropsis, Sub Jongmans, Lepidodendropsis sp., Lepidosigillaria cf. lepidodendron, Archaeocalamites, Sphenopte quadrata Danze-Corsin, Archaeosigillaria sp, ridium, Cardiopteridium and Adiantites from 'Cyclostigma' ungeri Jongmans, Gothan & Darrah, Kazakhstan. These reports show that the Archaeocalamites radiatus (Brongniart) Stur, Lepidodendropsis flora of Early Carboniferous had Rhodeopteridium tenuis Gothan and Rhacopteris d. already existed in the Angara area although some circularis Walton. The flora contains several genera endemic elements of the area appeared in the late which link it with typical Early Carboniferous flora Early Carboniferous. from other parts of the world and shows the close In addition, some genera of the Angara area were similarity with the Lepidodendropsis flora. Singh et cited by Cleal and Thomas (991), like al. (982) described fossil plants from the Lower Lophiodendron, Siberiodendron, CfJacassopterisand Carboniferous in Kashmir Himalaya, namely, Angaropteridium, etc. Some elements appeared in Archaeosigillaria minuta Lejal, Lepidosigillaria cf. the late Early Carboniferous. Among these elements, quadrata Danze-Corsin, Lepidodendropsis a number of Angara genera, such as Tomiodendron, fenestrata Jongmans, L. d. peruviana (Gothan) Abacodendron, Caenodendron and Ursodendron Jongmans, Cyclostigma d. pacifica (Steinmann) have also been found in Africa. Tomiodendron, Jongmans, Rhacopteris ovata (McCoy) Walkom, Abacodendron and Caenodendron are described Triphyllopteris lescuriana (Meek) Lesquereux, from the Lower Devonian in Libya (Lejal-Nicol, 1975) Rhodea d. subpetiolata (Potonie) Gothan and and Ursodendron and Tomiodendron are recorded Palmatopteris d.furcata Potonie. Recently, Pant and from the Lower Carboniferous in Niger (Rouvre, Srivastava (995) described the Early Carboniferous 1984). In the late Early Carboniferous, the fossil plants from Wallarma Spur of Panjab-Kashmir Lepidodendropsis flora was on the decline, while the Himalaya and established two new genera, viz., Pseu Angara flora began to develop. We believe that the dobumbudendron Pant & Srivastava and Spondylo Angara flora later gradually separated from the dendron Pant & Srivastava and four new species. The Lepidodendropsis flora of the Early Carboniferous. plant elements in this flora are as follows: Pseudo bumbudendron chaloneri Pant & Srivastava, P The Gondwana area meyeniiPant & Srivastava, P.fenestrata(jongmans & Koopmans) Pant & Srivastava, Spondylodendron The Kashmir region exposes one of the best wallaramensis Pant & Srivastava, Lepidodendropsis developed Lower Carboniferous sequences. The liddarensis Pant & Srivastava, Archaeosigillaria sub Lepidodendropsis flora of the Early Carboniferous is costata Danze-Corsin, Knorria Sternberg, Aspidiariu CHANDRA & SUN-DISTRIBUTION, EVOLUTION AND EXTINCTION OF EARLY CARBONIFEROUS FLORA 93 Presl, Triphyllopteris lescuriana (Meek) Lesquereux LepidodendropsisfenestrataJongmans, L. schw'man and Nothorhacopteris argentinica (Geinitz) Archan ni Jongmans, Lepidodendropsis sp., .?Sub gelsky. Early Carboniferous fossil plants from several lepidodendron fasciatum Jongmans, Cyclostigma localities in Kashmir Himalaya are now known to sinaica Jongmans, 'Cyclostigma' egyptiaca occur as a typical Lepidodendropsis flora. Jongmans, Asterocalamites cf. scrobiculatus The Lepidodendropsis flora of Early Car Schlotheim and Sphenopteris whitei (Berry) boniferous is also recorded from South America, Jongmans, etc. Lejal (969) described fossil plants Africa and Australia. Jongmans (954) recorded a from the Lower Carboniferous of Sahara, namely, typical Lepidodendropsis flora from Peru, including Protolepidodendropsis pulchra Hoeg, Lepidodendropsis de voogdiJongmans, L. cf. de voog Lepidodendropsis hirmeri Lutz, L. scobiniformis di Jongmans, L. peruviana (Gothan) Jongmans, L. (Meek) Read, L. sinaica Jongmans & Koopmans, L. steinmanni Jongmans, ?Lepidodendropsis africanum Lejal, Lepidodendropsis sp. Mensah and (Lepidodendron) lissoni (Steinmann) Jongmans, ChaloneI' (971) described two new species from the ?Lepidodendropsis sp., Cyclostigmaria pacifica Lower Carboniferous in Ghana, namely, Archaeosi (Steinmann) Jongmans, C. cf. pacifica (Steinmann) gillaria essiponensis Mensah & ChaloneI' and Jongmans, Rhacopterisovata(McCoy) Walkom, R. cf. Lepidodendropsis sekondiensis Mensah & Chaloner. circularis Walton, R. d. cuneata (Walkom), Tri Rouvre (984) described fossil plants, phyllopteris collombiana Schimper, T lescuriana Lepidodendropsis rhombi/ormis Rouvre, (Meek) Lesquereux, ?T peruviana Jongmans and Pseudolepidodendropsis nigeriensis Rouvre, Ur Sphenopteris whitei (Berry)]ongmans. Archa ngelsky sodendron w~jkianum(Heel') Radczenko and recorded fossil plants namely, Archaeosigil (990) Tomiodendron varium (Radczenko) Meyen from the laria conferta (Frenguelli) Menendez and Lepidoden Lower Carboniferous of Niger. The genera Ur dropsis sekondiensis Mensah & ChaloneI' from the sodendron and Tomiodendron are regarded as the Lower Carboniferous of Argentina. representative endemic elements of the Angara area, Rigby (969) compiled coaliferous plants but the two genera have also been found in Niger reported from Australia and recognized Calamites of Africa, therefore, they can not be regarded as peruvianus Gothan, Cyclostigma australe Feistman tel, Fryopsis sp., Lepidodendropsis sp., Rhacopteris the typical Angara elements. septentn'onalis Feistmantel, Rhacopteris intermedia As mentioned above, the Early Carboniferous Feistmantel, Rhacopteris ovata (Mc Coy) Walkom, flora of the Gondwana area contains numerous ele Sphenopteris clarkei Dun and Triphyllopteris sp. ments of the Lepidodendropsis flora (see Table 2), Jongmans and Heide (955) described fossil plants which is similar to those known from the other parts from the Lower Carboniferous in Egypt, namely of the world. Table 2 -Distribution of characteristic species of Early Carboniferous In Gondwanaland Species Kashmir Peru Australia Argentina Brazil Lepidodendropsis de voogdl Jongmans + Lepldodendropsisjenestrata Jongmans + Lepidodendropsis liddarensis Pant & Srivastava + Lepidodendropsispernviana (Gotha n) Jongma ns + LepidodendropsIS sekondiensls Mensah + & Chaloner Lepidodendropsis cf. sigillariaIdes + Jongmans, Gathan & Darrah Lepidodendropsis steinmannl Jangmans + Lepidodendropsis sp. + + + Pseudobumbudendron chalonerl Pant & Srivastava + 94 THE PALAEOI30TANIST Pseudobumbudendmn meyenii Pant & Srivaslava + Spondytodendron wal/aramensis Pant & Srivastava + Anhaeosigil/aria minuta Lejal Archaeosigil/an'a conferta (Frt::ngut::lli) + Menendez Anhaeoslgi/taha subcoslala Oanzt::-Corsin + Lepidosigil/aria cf. quadrata + Danze-Cmsin Cyclostigma austrate Feislmantel + CyclostIgma brasiliensis Oolianili Cyclostigma pacifica (Sleinmann) + Jongmans Archaeocatamiles radiatus (Brongniarl) SlUr + Rhodeoplmdium lenuis Gothan Thphyl/optern col/ombiana Schimper + Triphyl/opteris tescurtana (Meek) + Lesquereux Triphyl/opteris sp, Racopteris ouata (McCoy) Walkom + Rhacopteris cf. circutaris Walton + Shphenoptmdium sp, + + Sphenopteris whilei (l3erry) Jongmans Adianlilespentianus (Berry) Read + Adiantitesparacasica GOlhan + Cardiopleridium sp, + + Nothorhacopteris argenlinica (Geinilz) Archangelsby + + + + Fedekurtzia argentinica Archangelsky + + Kashmir Hoeg et at, (955), Pal & Chaloner (979), Singh et at, (982), Pant & Srivastava (995) Peru Jongmans (954) Australia see quotation in Rigby (969) EVOLUllON AND EXTINCTION OF GLOBAL EARLY Libya was the centre of origin for lycopods (Lejal CARBONllEROUSFLORA Nicol, 1975). In China, Sze (936) described So far, the evolution of Early Carboniferous flora Protolepidodendron ?arborescens from the Middle is not clear, owing to limited data and different views. Devonian in Hunan Province. Later, Sze (953) According to Lejal-Nicol (975), the elements of the placed this species with a question mark under the earliest Lepidodendropsis flora appeared in the genus Lepidodendropsis, namely Lepidodendropsis Lower Devonian of Libya, North Africa, namely, ?arborescens Sze (Sze). Lepidodendropsis? arbores Lepidodendropsis hirmeri Lutz, L. sinaica]ongmans cens was also recorded from the Upper Devonian in & Koopmans, L. africanum Lejal, Lepidosigillaria Hubei Province (Sze, 1952). Sze (956) reported fossil whitei Krausel & Weyland, L. massae Lejal-Nicol, plants from the Wutung Series of Kiangsu (Jiangsu), Archaeosigillaria minuta Lejal and Prelepido namely, Lepidodendropsis hirmeri Lutz and Sub dendron mourzoukenseLejal-Nicol, etc. It seems that lepidodendron wasihense (Sze) Sze. Chang Chien- CHANDRA & SUN-DISTRIBUTION, EVOLUTION AND EXTINCTION OF EARLY CARBONIFEROUS FLORA 9S shen (965) described Lepidodendropsis scobinifor 1952). So far some elements of the Lepidodendropsis mis (Meek) Read and Leptophloeum suzhousenese flora are on record from the Upper Devonian, there Chang from the Wutung Series of Kiangsu Qiangsu). fore it is possible that the Lepidodendropsis flora Leptophloeum suzhousense has been transferred to represents the floral appearance of the Late Devonian Leptophloeum rhombicum Dawson (Gu & Zhi, 1974). and the Early Carboniferous (Sun Keqin, 1996). In The Wutung Series (Wutung Formation) has now view of the above mentioned facts, the regarded as the Late Devonian in age (Wu been Lepidodendropsis flora gradually separated from the Chongzhang et al., 1982; Zhao Xiuhu & WU Xiuyuan, Early Devonian, started to diversify in the Late 1986). Zhao Xiuhu and Wu Xiuyuan (986) listed Devonian and reached its maximum development in fossil plants from the Wutung Formation, namely, Leptophloeum rhombicum Dawson, Sub- the beginning of Early Carboniferous and caused its lepidodendron mirabile (Nathorst) Hirmer, extinction in the end of the Early Carboniferous. Lepidodendropsis hirmeri Lutz, Lepidostrobus Seeing that some genera of the Angara area such grabaui Sze, Cyclostigma kiltorkense Haugh, as Ursodendron, Tomiodendronand Lophiodendron, Hamatophyton verticillatum Gu & Zhi, Ar etc. occurred in the South of former U.S.S.R., Tuva chaeocalamites aff. scrobiculatus (Schlotheim) and Mongolia. Vakharmeev et at. (970), ChaloneI' Seward, Sphenophyllum pseudotenerrimum Sze, S. and Lacey (973) and ChaloneI' and Meyen (973) lungtanense Gothan & Sze, Archaeopteris spp., recognized an Angara floral province of Early Car Sphenopteris taihuensis Sze and Platyphyllumd. wil boniferous in the region. However, the genera liamsoni(Nathorst) Hoeg, etc. Li Xingxue and Wang Tomiodendron, Abacodendron and Caenodendmn Hongfeng (982) described the Late Devonian fossil have been recorded from the Lower Devonian in plants from Mt. Longmenshan, North Sichuan, name Libya, Africa (Lezal-Nicol, 1975) and Ursodendmn ly Lepidodendropsis theodori (Zal.) Jongmans, L. d. and Tomiodendron were reported from the Lower theodori (Zal.)Jongmans, Lepidodendropsis? sp. (d. Carboniferous in Niger, Africa (Rouvre, 1984). There L. ?dzungariensis Sze), Lepidosigillaria ? si fore, the genera Ursodendron, Tomiodendron, chuanensis Li & Wang, Archaeocalamites longinter Abacodendron and Caenodendron in the Angara nodus Li & Wang, Archaeocalamites sp., etc. Zhao area could not be regarded as typical Angara floral Xiuhu and Wu Xiuyuan (986) described fossil plants elements. In this light the Early Carboniferous flora of from the Zhongning Formation of the Upper Devonian in southern Ningxia. These are Leptoph the Angara area also belonged to the Lepidoden loeum rhombicum Dawson, Sublepidodendron dropsis flora of the world. mirabile (Nathorst) Hirmer, Hamatophyton verticil As might be expected, a number of forerunners latum Gu & Zhi, Archaeocalamites sp., Sphenopteris of the Angara flora existed in the late Early Car taiuensis Sze, etc:. The flora of the Zhongning Forma boniferous, which indicated that the Lepidodendrop tion shows a striking resemblance to that of Wutung sis flora was on the decline, and the Angara flora Formation in southeastern China and is probably of started to develop. Sun Keqin (996) pointed out that the Devonian age (Zhao Xiuhu et al., 1986). In view owing to the climatic differences, some obvious of the above mentioned facts, the early Early Car changes in floral components of the Cathaysia, boniferous flora closely relates to the Late Devonian Euramerica, Angara and Gondwana areas occurred flora, while the Late Early Carboniferous flora during the transition from the Early Carboniferous to resembles the Namurian flora. Late Carboniferous, which resulted in extinctions of In the Donetz Basin, the Lepidodendropsis flora some typical plant genera, such as Lepidodendropsis, Sublepidodendron, Archaeocalamites, Triphyllop is said to occur in the Upper Devonian Qongmans, tern, Cardiopteridium, Rhacopteris, Fryopteris, 1954). "It is interesting to note that such plants, espe Sphenopteridium, Rhodeopteridium and Adiantites, cially Lepidodendropsis, also occur in different etc. of the Early Carboniferous. Moreover, a number localities and are considered as Upper Devonian" of forerunners of the Cathaysia, Euramerica, Angara Qongmans, 1954). "It is a well known fact that the and Gondwana floras had already appeared in their flora of the lower part of the Lower Carboniferous has respective areas during the late Early Carboniferous. relations with that of the Upper Devonian and of the During the late Early Carboniferous, most parts of the younger part with that of the Namurian" Qongmans, Gondwana area were controlled by glaciation, there- 96 THE PALAEOBOTANIST fore, the record of typical forerunners of the Chaloner WG & Lacey WS 1973. The dIstribution of Late PalaeozOIc floras. Gondwana flora is unclear in this age. But the genera In: Hughes NF (Editor)-~anismsand continents through tIme. Spec. Pap. Palaeont. 12: 271-289. like Nothorhacopteris and Fedekurtzia may be Chaloner WG & Meyen SV 1973. Carboniferous and Permian floras of the regarded as endemic elements to Gondwana northern continents. In Hallam A (Editor)-Atlas oj Palaeo countries (Pant & Srivastava, 1995). Global climatic biogeography: 169-186. Elsevier, Amsterdam. differentiation was not very obvious during the Early Chen Fen, Sun Keqin & Zhou Hongrui 1994. Discovery and significance of the genus Cardlopteris. Acta Bot. sin. 36(7) : 552-556 (in Chinese Carboniferous. The lycopods, which were quite with English Abstract). abundant and dominant in the Lepidodendropsis Chen Fen, Zhou Hongrui, Sun Keqin, ]ia ]inhua, Zhang ]ianping & Wu flora, indicated warm and humid climatic conditions. higuo 1995. Carboniferous flora in Ningxia and adjacent regions. During the transition from the Early Carboniferous to Geoscl.]. ojGraduate School, China Unlv. oj Geosci. 9(1) : 1-10 (in the Late Carboniferous, the differentiation of obvious Chinese with English Abstrdct). Cleal C] & Thomas BA 1991. Carboniferous and Permian palaeoge climatic conditions, tectonic movements, continental ography. In: Cleal C] (Editor)-Plantjossils Ingeologlcallnvestiga positions, oceanic currents and glaciation might have tion: The Palaeozoic: 154-181. Ellis Horwood Limited, England. caused the extinction of the Lepidodendropsis flora. Deng Bao 1978. On the discovery of Early Carboniferous flora from The Cathaysia, Euramerica, Angara and Gondwana Shanyang, S. Shanxi and its stratigraphicsignificance. Acra.geol. sin. 52(1) : 15-21 (in Chinese with English Abstract). floras derived from the identical Lepidodendropsis Feng Shaonan, Hu Yufan & Zhu ]ianan 1982. Fossil plants and their flora of the Early Carboniferous, but developed in assemblages from the Early Carboniferqus in Guangdong. ACta bot. different environments respectively (Sun Keqin, sin. 24(4): 374-382 (in Chinese with English Abstract). 1995,1996). However, it is noteworthy that the origin Gothan W& Sahni B 1937 Fossil plants from the Po Series of Spiti (N.W and extinction stages of the Lepidodendropsis flora Himalayas). Rec. geot. Surv. India 12: 195-206. Gu & Zhi 1974. Palaeozoleplantsjrom China. Science Press, Beijing (in varied in four major areas with theirdifferent environ Chinese). ments and ecological variations. In the Early Car Hoeg OA, Bose MN & Shukla BN 1955. Some fossil plants from Po Series boniferous although the Cathaysia, Euramerica, An ofSpiti (NW. Himalayas). Palaeoboranist4: 10-13 gara and Gondwana areas contained some of their ]ongmans W] 1952. Some problems on Carboniferous stratigraphy. Compt. Rend. Congr. Avan. Dud. Strarlgr. Carbon. 3me, 1: 295 elements to minor ecological variations of dif due 306 ferent vast areas, they retained the character of the ]ongmans WS 1954. The corboniferous flora of Peru. Bull. Br Mus. original Lepidodendropsis flora. (Natural History) Geo/. 2 (5) : 191-223 ]ongmans W] & Heide S van der 1955. Flore et faune du Carbonifere inferieur de I Egypte. Meded. Geo/. Sticht. Nleuwe ser. 8: 59- 75. ACKNOWLEDGEMENTS ]ongmans W], Gothan W& Darrah WC 1937. Beitrage Zill Kenntnis der Flora der Pocono-Schichten aus Pennsylvanien 1Jnd Virginia. C.R. Sun Keqin expresses his deep sense of gratitude 2me Congr Avanc. Etudes Stratigr. Carbonifere (Heerlen, 1935) 1 to Dr R.S. Tiwari, Former Director, BSIP, Lucknow : 423-444. Lacey WS 1962. Welsh Lower Carboniferous plants. I. The flora of the who accepted his visit as a Visiting Scientist from Lower Brown Limestone in the Vale of Clwyd, North Wales. October 1995 to March 1996. He is thankful to all the Palaeontographica 1118: 126-160. staff members of the Birbal Sahni Institute of Lacey WS & Eggert D 1%4. A flora from the Chester Series (Upper Palaeobotany for helping him in various ways. He Mississippian) of southern Illinois. Amer.]. Bor 51(9) : 976- 985. Lejal A1969. Etude des Sublepidodendraceae du Djado (Sahara Oriental). also acknowledges the authorities of the Birbal Sahni Palaeobolanisl17: 137-151. Institute for awarding him Research Fellowship. 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