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Paleoclimatic implications of Lycophyta in the Gondwana of Southern Brazil

Margot Guerra-Sommer, Miriam Cazzulo-Klepzig, Marleni Marques-Toigo Pesquisas em Geociências, 22 (1/2): 21-31, set./dez., 1995.

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Data de publicação - set./dez., 1995. Instituto de Geociências, Universidade Federal do Rio Grande do Sul, Porto Alegre, RS, Brasil PESQUISAS. 22 tl·2):21·11. I99S 1.. 1i,"",

Palaeoclimatic implications of Lycophyta in the Gondwana of Southern Brazil

lriSlitulO de Geociencias. Universidade Federal do Rio Grande do SuI, Caixa Postal 15052. CEP 91501·970. Porto Alegre. RS. Brasil.

(Recebido em 06,1)3195. Aceito para publica~Ao em 23/12195.)

Abstraci _ The Lycophyta played an importruu role in Ihe noral associations developed during times in the Gondwana area of South Brazil (State of Rio Grande do Sui). The Iycophyte affinities of certain palynomorphs (Lulldbladispora. Vllllarisporiles. Crisllllisporiles). and the impressions of compressed molds of lycophytes. suggeSlthat the climax of Lycophyta look place under coal·measure conditions in a warm. seasonal, climate. At some levels, dense thickets of arborescent forms have been found "in situ": thcir cormose TOOting systems are non·sligmarioid. Thechangcs in climatic and palaeoecological conditions that occurred during Late Pennian times are renected by a mpid decrease of pteridophylic spores. particularly those related to Lycophyta and the concommilant increase in gymnospenn saccate polien. During Ihe uppennost Lmc Permian Ihe nora became sirikingly impoverished. glossoplerids occurred as complementary elemcms nod lycophytes became dominants. These palaeonlological dma which are COrroboraled by facies analysis. indicalc a very warm, semiarid climate 10 Ihe sOUlhem portion of Paran~ Basin.

Resurno • As LycophYla desempenhamm importallie papel para 0 desenvolvimcmo das associilti6es norislicas durame 0 Permiano no Gondwana do sui do Brasil (Esmdo do Rio Grande do SuI). As afinidades botanicas de palinomorios como LWldbllldispor(l. V(ll/lllisporiIU. CriSUJlisporilu) e impressUcs e compressUcs de Lycophyta sugerem que 0 clima~ deslc grupoocolTCu duralliC adeposi~ao das camadas de earvao. em dima quellie c sazonal. Em alguns niveis. densas associiltiOes de fomlas arborescentes foram encontradas "in situ". Seu sistema radicular cormoso ~ do tipo nao eSligmari6ide. As modifica~Oes nas cond i~ Oes dim~ticas e paleoecot6gicas que ocorreram no Pcnniano Superior s;\.o evidenciadas por urn r~pido dedfnio de csporos de ptcrid6filas. particulamlentc os rcferidos a Lycophyta. e aumcnto concomitanle de p6lens sacados de gimnospermas. No IOpo do Permiano Superior a composi~ao norislica sofreu urn r:ipido cmpobrecimcnto; as glossoplcrfdcas ocorrern como clcmemos complcmentares e as Lycophyta vol lam a dominar. Esles dados palcomol6gicos, corroborados por dados de an~lises faciol6gicas, indicam clima muito quente. semi·:llido. Os resuhados oblidos nCSIC trabalho referem·se e~clusivamerlte i\ parte sui da Bacia do Paran~. Estado do Rio Grande do Sui.

INTRODUCTION

The intracratonic Parana Basin, with a total area of 1.700.00 km2(1.IOO.OOO kml of which correspond to Brazil) is situated in east-central South America. It contains Palaeozoic, Mesozoic and, locally, also Caenozoic sediments. The present study is based on data from Pennian sedimems in the southern part of this basin in Rio Grande do Sui slale. Brazil (Fig. I), comprising, in agreement with Schneider et al. (1974), the following lithostratigraphic units: (Fig. 2).

C3 c-",,,, I!ZJ

=EO]

Figure I • Localion of the gondwana rocks in Rio Gnmde do Sui (Southern Figure 2 • Stratigraphic column of the Paran~ Basin (Brazil). (Modif. Bra1';1). Schneider el (Jt., 1974). 22

Itarare Group. Guata Group ( Ri o Bonito and Palermo structures in diamictites indicate peri- and proglacial lake Fonnati ons) and Passa Dois Group (Irati. Estrada Nova and conditions (Itarm-e Group -/Artinskian). according Rio do Rasto Fonnations - Fig. 2). to Piccoli (1989). The macropalaeobotanical and palynological dalll howe In the associated rhythmic deposits the palynological made it possible to establish a climatic evol ution; the increase assemblage is composed primarily of po ll en grains belonging in Iycophyte remains up-sequence is considered to be a to Potollieisporites. Caheniasacciles, Camwfloropollis. parameter of wanning. Lithological characters have been PJicalipolielliles. Prolohapioxypilllls and Vitlalilla, and trilete used as supplementary data and have corroborated the spores such as Llllldbladispora. Vallatisporiles. infonnation suppl ied by micro and megafloral data. PUllctati!'poriles. GrllllUlalisporiles. etc ., as we ll as A comparison between the Late Palaeozoic floral palynomorphs of unknown affinilies (POl"taliles. Pilosporiles, elemenls of the Brazilian portion of the Parana Basi n and Telrllporilla). The high proportion of pollen grains indicates Ihose of other Gondwana areas reveals an overall similarit y that gymnospcmlS were the predominant floral elements in between NOlOafroamerican laphofloTlls with an important th is postion of the basin. Pteridophytes were mere accessories Iycophytecontribution. In the Auslralo/1ndian Province this (Cazzulo-Klepzig, el al., 1980). group is poorl y represented in the micro and megafloristic On the other hand. the megafloral remains recovered associations (Taylor & Taylor. 1989). from sandy sediments corresponding to the Facies Suspiro­ A review of Pennian lycopods recorded from southern (Piccoli, 1989) of the Itarare succession mainly belong 10 part of the Parana Basin shows high proportion ofspccimens the glossoplerids, e.g. Rllbidgea, GlllIgamopteri.~. with somc represented by isolated vegetative shoots preserved :IS Glossopteris (P I. I. Fig. 3). Relicts such as BOllychiopsis impressions. Unfortunmely, the poor preservation does not p/amialla (Carruthers) Archangelsky and AITOndo (PI. I. Fig. frequently offer an adequate basis for generic designations. 2) are still important components of th is taphoflora. which Fragments of lycopods are assigned in literature as represents an association similar to the present-day "taiga". Lycopodiopsis "sensu lalo" following a general tendency 10 suggesting cold climatic conditions (Retallack. 1980). attribute Pemlo- Gondwana lycophytes to this Pteridophytic megafossils, including Lycophyta, were not genus. registered in this periglacial palaeoflora (Cazzulo-Klepzig The first Lycophyta described from the Gondwana of & Guerra-Sommer, 1983). Rio Grande do Sui was identified as Flemi"gites petiroa"lIIn With the retrcat of the ice-caps, the glacigene by Carruthers (1869). Later. the type-material of this genus sedimentation was replaced by fluviodeltaic depositional has been redefined as Lepidodendron pedroollllm (Zei11er. systems, represented by a thick succession of sandstones. 1895. White. 1908. Lundqvist. 1919). Lycopodiopsis siltstones, and coal seams (about 60-70 m), belonging to the pedroallllln (Edwards, 1952) and. most recently. as Rio Bonito Fonnation (Paim el af.. 1983). The taphofloras BrasilOllendron petiroonllm (Chaloner el af. , 1979). Another of the basal sil tstones are chardcterized by the presence of genus described from south Brazilian strata is BOlrychiopsis p/alllialla and articulates (Phyllolheca iluJica Lycopodioph/oios (Krliusel, 1961), but this fonn probably Bumbury. PI. I , Fig. I), a hygrophile assemblage which corresponds to a different level of decortication of occurs in bands associated with glossopterids, G/OsSOIJteris. Lycopodiopsis or BrasilodendrOIl. Gangamopleris, Rubidgea. The association of Botrychiopsis More accurate observmion of some samples. hitherto IJfamialla with Pllyllotheca indica is considered to indicate undescribed, reveals a grem diversification among Pennian a climatic amelioration (Guerra-Sommer, 1979) Lycophyta Gondwana lycophytes in southern Brazil. However. a fonnal have not been recorded in these deposits. description of megafossil taxa is outside the scope of this Microfloral assemblages arc still dominated by saccate paper. pollen (Cahelliasaccites. POlOllieilporiles. Call1lflllOropolis. An abundance of lycophytes in the plant communities Plicatipollelliles. ProlOhap/o.l}piIlIlS, Villa/ilia). However. of Rio Grande do Sui is also apparent from the palynological there is a gradual increase in the proportion of Sporites, which analysis, taking into account the palaeobotanical affinities indicates that. with the retreat of the glaciers, the Pteridophyta of the spores. according to Azcuy (1978), Doubinger ( 1959). found favourable conditions, allowing them to prol ifera te. Gould & Dclevoryas ( 1977), Boureau & Doubinger ( 1975). Trilete Iycophyte spores are represented mainly by the genera Paleobotanical samples (Pb) and palynological slides Lllrulb/adispora. Crislalisporiles, Vallolisporiles and (M P-P) are registered and deposited at the Museum of Kraellselis/Jorites (Marques-Toigo, 1988). Paleontology, Instituto de Geociencias. UFRGS. The river system that drained the alluvial bas ins allowed Megafloristic data were based on Guerra-Sommer & peat fommlion to take place in subsiding backswamp areas, Cazzu lo- Klepzig (1993). marshes and lakes (coal-bearing strata - Piccoli el al.. 1986). The micro and macrofloral assemblages of the coal LYCOI'HYTE REMAINS AS EVIDENCE OF seams and associated strata show different vegetational PALAEOCLIMATIC EVOLUTION patterns in the different coalfields. However. the microfloras of the South BT'dZilian coal measures are mainly characterized The lower part of the Lale Palaeozoic Gondwana by a dominance of pteriodophytic spores. The principal trilete succession in the State of Rio Grande do Sui, southern Brazil spore genera are PlllICllIlisporiles . Leiotriletes. is characterized by glacial sediments compri sing mainly Calamospora. Grolllliarisporites. Cyclograllisporiles, ti ll ites and diamictites as a general rule. Sedimentary florriditrileles, Apicllialisporites. Allapicll/atisporiles.

Plate I

Figure I· PhyllQlheCII im/iea.· Ph 28t5 (Rio Booilo Formation. QuilEria outcrop) oUlcrop) Figure 4· S"II4'IIo{lI4'fI£ cf ;sc!wIIO\'ellS;s , Pb 2530 (Rio Bonito FOmlalion. Figure 2· Bo/ryclriIJP£i£ JI/ulHiana . • Pb 21 12 ( Rio Bonito Fonnation, Faxinat Coalmine) Papaleo OUtcrop) Figure 5· GinkgophylOJlsis kids/onii· Pb 25 (trJlf Fo:nn.ation.. PanlanoGrande Figure 3· GIIJs$opltris sp .• Pb 2858 (Rio Bonito Formation. QuitEria regim). 23

M. Guerra· Sommer et 01. Plate 1 24

LUI/db/adispora, Vallarisporites and Crislatisporires. The rare Corfl/ophyton ",aWl/elise Pigg and Taylor. Meyen (1987) monolele spores are represented by Lnevigatosporiles and suggcsled that Gondwana lycophytes, with cormose bases Pllllctatosporites. Saccate gymnospenn pollen are repre­ should be ranged with (he Chaloneriaceae rather than the sented mainly by Shellringipol/enites. Vesicaspora. Lepidocarpaceae. Cahel1iasaccires, CUllllolloropolis and POlOllieisporiles. The " In situ" casts of lycophyte trunk bases, lacking mOSI common fo rms of Sirialili are the genera sti gmarian axes, were described by Cuneo & Andreis (1983) Protohaplox),piflllS and Vitta/ina. Algae and/or related from Permian strata of the Central Patagonian Basin. elements occur in several coal seams: these areBotryococcus, Argentina. The charncters of this association are quile similar Terraporino, Portaliles, Pifasporires, Quadrisporites and to those referred to in the present paper. According to Cuneo Bro;i/ea (Marques-Taiga, 1988). & Andreis (op. cit.) these fragments were remains of an Taking into account the quamitative analysis and the arborescent lycophyte forest that developed in a foodplain botanical affinities of the diagnostic of environment under frost-free climatic conditions. The dense Lundbladispora, Cristalisporiles. and Vallatisporires, thickets of arborescent lycophytes with cormose bases. Marques-Toigo & Correa da Silva (19&4) suggested that the occurring in the roof of coal seams, are found associated with peat-fonning vegetation of the South Brazilian coals was fernlike frond s, The predominance of herbaceous aniculates composed mainly of herbaceous plants and shrubs related (0 in hygrophilous environments suggests a change from cool Lycophyt:l: "LlIndbladispora. PlInctatisporires · Portalires temperatures to a milder season climate. The dominance of Associarioll" and "Cristatisporites - Vallatisporites Iycophyte spores in these Simla reinforces the suggestion of Association". a climatic improvement. The ScliellrillgipollelliteslCahelliasacci tes Associal ion In the upper part of the Rio Bonilo Fonnation a gradual and other mono and bisaccate fonns would represent a change from fluvial to coastal sedimentation may be inferred gym nos perm vegetation. mainly belonging to from the presence of fine-grained sandstone sequences with Glossopteridopliyta, Coniferophyta and Cordaitophyra. In hummocky, wavy and parallel lamination, representing this association lycoph yte spores are poorly represented. lagoonal washover fall. tidal flat, tidal inlet. foreshore, and The megaflora I assemblages of coal-bearing strata upper-shore fac ies (Palcnno Formation, according to Lavina reveal an important compositional change, as is evident from el al.. 1985). the presence of filicoid fronds, i.e. Sphenopteris cf. The upper part of this succession is composed of ishollOI'ellsis Zalessky (PI. I. Fi g. 4) Rodhea sp. and siltstones and sandstones, laid down as stonn deposits, Ellcopreris meridiollalis, Oolianiti which is of pteridophytic Lycophyte remai ns, preserved as molds and casts (PI. 3, Fig. or pteridospenn affinity (Guerra·Sommer, 1989). These I). are the most megaflora I elements found in these fossils correspond most likely to herbaceous plants that environments (Cazzulo-K lepzig & Guerra-Sommer, 1985). fonned the understorey of arborescent gymnospenn stands. Different lycophyte morphotypes can be recogni zed in The latter are mainly glossopterids. and, to a lesser extent. the samples: the most common one of these corresponds to also cordaitaleans. On the other hand, herbaceous articulates s tem and/or branch fragments with a lepidodendroid (Phyllotheca) dominate in the hygrophilous assemblages. phyl1otaxy, and distinctly elongated leaf cushions without (Guerra-Sommer et al., 1991) proper leaf scars (PI. 2, Figs, 3, 6. 7, 8). The leaves are Recent studies on the coal-bearing strata of Rio Grande supposed to have been attached to the topmost pan of the do Sui (Ri o Bonito Fonnation - Aninsk.ian/Kungurian) have leaf cushion, This type is similar to Lepidodendrop.\·is revealed the presence of "in situ" connose lycophyte bases Krausel and 8rasilodel/dron Chaloncr, Leistikow & Hill. forming dense thickets above coal seams (Piccoli et al., Another type, which is less abundantly represented. 1991). These lycophyte bases are in organic connection with shows leaf cushions of hexagonal shape, isodiametric 10 radial appendices which functioned as roots (PI. 3, Fig, 2). elongate with very narrow intereareas. Evidence of leaf scars These radial appendices, which are arranged can be observed ilt some levels (PI. 2, Figs. I, 4). These perpendicularly 10 the trunk, show spin y surface excrescences characters are re miniscent of Lycopodiopsis Krausel and (Pl. 3, Fig. 5). In the upper part of Ihese bases, the cushions Allgarodendroll Meyen. ImpressionsofGlossopleris, and of become elongate rhomboidal (PI. 3. Figs, 3, 4). The connose Paracalamites are fo und associated. The composition of the bases CUi across differenllithologies, ranging from mudstones associated palynoflora shows changes in percentages between to sandstones. Filicoid fronds ("Rhodea" sp.) and fragments the trilete spores and saccate Striatiti in re lation with those of glossopterids and Cordaites were identified from these of the coal seams. The increase and diversity of Striatiti in sediments. Sedimentological and palaeontological data from this assemblage may be considered as marking the begining these plant beds suggest an evolution from swampy of important floral and climatic changes, The palynoflora conditions to an alluvial plain environment with periodic appears to reflect contributions from a gymnosperm­ flooding (Guerra-Sommer, 1989), dominated "upland" flora producing disaccate, monosaccate Connose Iycophyte bases have been described from the and Striatiti pollen, mixed in with trilete spores from a more Palaeozoic sediments of Euroamerica (e.g. Chaloneria localised flora of pteridophytes. mainly Lycophyta cormosa. Pigg and Rothwell, ProfOsligmaria eggertiana (Llllldbladisporaand Crislatisporites). which grew in an area Gennings, Selagillella jraipollfii Schlanker and Leism:ln, marginal 10 the site of deposition (Cazzulo-Klepzig el al ..

Plate 2 Figures I. 4 _ Compressed stem: leaf cushion$ hexagonal shaped, leaf SCW'll absent - t'" 1229 (Patenno Formal ion, sao Scpe outcrop) isodiomctric to elongated: intereushioos areas narrow, leaf scatS evident - Figure 6· Pmgmenl with tepidodendroid phyllolaxyelongaled leaf cushions Pb t288 (Patenno Fonnalion, Sfto Sepe OUICrop) (mediwiline evide1ll) leaf scars ahse1ll - Pb 1104 (Palermo Fonn31ion, Siio Figures 2. 5 _ Frngment wilh lepidoderKIroid ph~Uolax y. broad intereushion Sept outcrop) areas· Pb 3292 (Eslruda Nova Fom18lion, Cerro OalO oU lcrop) Figures 7. 8 - Frngmem with lepidonderKIroid phyllolOxy.lcaf scars prcse1ll Figure 3 _ Fragment wihlltpidodcndroid phillOlaxyelongalcd leaf cushions, - Pb 1123 (Patenno Fonllation, SAo Sept outcrop). 25

M. Guerra - Sommer et al. Plate 2 26

1989). FINAL CONSIDERATIONS Overlying the Palermo Formation deposits a palaeoenvironmental change from shallow waler 10 restricted In evaluating the parolmeters supplied by mega- and marine conditions, associated with lagoonal, lacustrine and microflora associated with different sedimentary facies in coastal swamp areas, is marked by dark grey to black and the Permian sediments of the State of Rio Grande do Sui brown oi l shales, Iighl to dark grey shales (non-bituminous) southern part of the Paranil Basin (Brazil), it has become and carbonates (Irati FormUlion - Kazanianffatarian, evident that the palaeofloral evolution was related to climatic according to Lavina, 1982). Mudstones and silty sandstones, conditions. indicate shallow water platform facies, (Estrada Nova Since the palaeobotanical and palynological data are Fonnation - Tatarian. Fig. 2). insufficient on their own to produce a complete outline of The palynological assemblages from Irati Fonnation are the palaeoclimatic changes, lithological data were also taken dominated by Striatiti (PrOiohapioJ.}'Pinlls, Llleckisporilt!s, into account. Slriatopodocarpites, Villalina) and algal remains The homogeneous composition of Early Permian floral (BorryococCfls), whilst trilete lycophylC spores are Tare assemblages. in the southern part of the basin, which are (Cazzulo-Klepzigel al., op. cit.). According to Foster(1979), dominated by herbaceous to shrub-like plants considered to a high proportion of taeniae in the pollen grains among the be relicts from the rigorous climate of an Ice Age (e.g. saccates indicates a defense mechanism against desiccation. Botrychiopsis pIal/rial/a) suggests the persislance of a cold It is therefore suggested that the Stri atiti were adapted to drier climate. The dominance of Rubidgea and GOllgamopreri.f climatic conditions. leaves with a palmate venation, associated with glossopterids The megafloral assemblage of the Irati Formation is with penate venations seems to indicate a gradual warming. represented by varied wood remains corresponding mainly Diamictite-tillite sequences are indicative of glacigene to picnoxylic tree trunks of gymnosperm affinity, with a sedimentation, but there is no evidence of pennafrost in these diaphragmatic pith: Poiysoienox),loII, Scleromedlllloxyion, basal strata. Lycophyte megafossils are registered from these Arachnomedlllloxylon, Septomedllfloxylofl (pessoa. 1985). levels. In the microfloral assemblages Iycophyte spores are The growth rings in these wood remains are thick; the layers present only as accessory elements (Lundbladisporo, being very thick in relation to those of late wood. Detached Vallorisporires). and this suggests that the Lycophyta were roots of Vertebraria type are common in this flora, which not a flourishing group in cold climates. has been related to swampy-brackish environments under Upwards coal and coal-bearing strata (Coalfields warm climatic conditions (Mussaet al., 1980). Ginkgophyta Capanc, Candiota, trur. Pantano Grande, Faxinal, Leiio-Butia. (Gillkgopliytopsis kidstOflii) become commonc1ements in the Charqucadas-Sanla Rita. Chico-Loma e Santa Terezinha). megaflora (Backeuser el aJ., 1982) ilustnlted in Plate I, Figura 5. suggests improved climatic conditions. The palynological A gradual return to continental conditions is suggested assemblages of coals are dominated by Cingulicavati trilete by the presence of variegated sandstones, siltstones and spores. thus suggesting that the peat-forming mires were mudstones. deposited as coastal to fluvial sediments which generally populated by a lycophyte vegatation rather than cover carbonate deposits (Estrada Nova and Rio do Rasto gymnosperms. The low diversity of forms and the abundance Formations, Upper Permian - Lavina, 1992). of some genera (Lulldblodispora, Crisratisporites, The megafloras are dominated here by impressions and Kraellselisporites, Vollatisporires) among miospores of Rio casts of Iycophyte stems, which are abundant but poorly Bonito Formation in the southern part of the Parana Basin preserved (Lycopodiopsis), and detached lepidodendroid suggest a low-diversity swamp flora dominated by Lycophyta leaves. Rare glossopterid fmgments can be identified in these (Marques-Toigo & Correa da Silva. 1984). bands (Lima & Cunha, 1976). A common characteristic of In coal-bearing strata pinnate glossoperid relatcd to the Iycophyte fragments in this assemblage is a typically forms relate to Glossopteris are common; GOllgamoplcris lepidodendroid phyllotaxy, with broad interareas, and the and Rubidgea (palmate forms) are poorly represented. A absence of a ligule(PI. 2, Figs. 2. 5). In some remains, the sudden enrichment of articulates and filicoids is basal parts of acicular leaves are found in organic connection characteristic. Compressed trunks of lycophytes become with the cushions (PI. 2. Fig. I). Krausel (1961) described common elements (LycopodioJJsis, Brasilodendron). In the L)'copodioph!oios from these strata as a genus distinct from roof of some coal seams dense thickets of arborescent L)'copodiopsis. lycopods with cormose bases provide evidence that the The sudden impoverishment of Glossopteris and the climax of Lycophyta took place during the depositi on of this dominance of lycophytes can be attributed to the increasingly coal-bearing strata. Additionally, the presence of marls at warmer climatic conditions during the Permo- the top of the coal-bearing deposits is a lithological indicator transition. This agrees with sedimentary facies (red beds) of clim .. tic wamling (Guerra-Sommer et al.• 1991). indicating aridity. Palynological assemblages are not recorded Therefore, it appears that the cool seasonal climate of from these beds. earl y Pennian times changed into a moist seasonal interval The most representative genera of pollen and spores during the Artinskian-Kungurian. This climatic change was which occur throughout the studied succession are illustrated significant for the meso-hygrophile to hygrophile vegetation on Plates. 4 and 5. that originated the biomass of the Gondwana South Brazilian

Plate 3

Figure I _ Mold of Lycopllyt3 trunck paniolly enclosed in the sediments· Figures 3,4 • Elongated cusllions from the tOp of the cormose basis· Pb Pb 1185 (Palermo Formation. S!\o ScpE OUtcrop). 3088. Figure 2 . Lycophyta cormose basis· Pb 3088 (Rio Bonito Fommtion. Figure S - Details ofllle radial apendices sllowing occute ucrescences· Pb Quit~ria outcrop). 3088. 27

M. Guerra· Sommer el at. PlateJ 28 coals. Thus, the dominance of plcridophylic spores in the _-.--,-,-----::-: Piccoli. A.E.M. & Marques-Toigo. M. 1989. Reco­ peat-fonning assemblages is not only due to ecological nhccimcmode urn man:ot'$lrntignifico com significado paleoecol6gko conditions but also re fl ects an adaptation to ciimalic !lO Pernliano da Bacia do ParonA (Grupos Gual~ e Pass.a Dois). "'"1115. improvement. II' Congresso Brasi/tiro de P/lltoll/ologia. Curiliba.. f>aran1. 2:972- Changes in palaeoenvironmental conditions can be .. , Chaloner. W.G.; LciS'likow. K.V. & Hill. A. 1979. 8ra5i1od~ndrOll 8en. inferred on palaeobotanical and palynological grounds in the !lOv. and B. {WlirocJIlunr (Carruthers) comb. nov .. a Permian lycopod succession whic h overlies the coal-bearing rocks. Conditions from Brnzll. Rtl·. Pa/tobol. Po/yllOl., 28: 117-136. pI. 1-4. leading 10 peal fo rm mion disappeared; the major flo ral Cuneo. R. & Andreis. R.R. 1983. EslUdiode un bosque de lic6filas de la changes indicates a shift in climati c regime as shown by the formaci6n Nueva Lubecka. Permico de Chubul. Argentina. increase of gymnospermic plants adapted to a seasonal Implicaciones paleoclimalieas y paleogeograficas. Ameghinionu. cl imate. 20(1-2): 132-40. The microfloral succession also re fl ects important Doubinger. J. ]959. Palynologie el PalWbolanique. Polltll tr Sporn. changes re lated 10 sudden increase in saccale gymnosperm 1(2):279-309. pollen. Edwards, W.N. 1952. Lycopodiopsis. a southern hemisphere lepidophylc. A dramati c change in cl imate conditions occurred during The PoloeQb%llisl. Luck now. I : 159-64. FOSler. C.B. 1979. Permian plant microfossils of Ihe Btair Athol Coal the uppermost Permi an. The flora became strikingly Measures. Bnralab.1 Coal Measures and Basal Formation of Queensland. im poverished. glossoplerids became accessory elemenls, G~ol. S" n'e}" QIII:cIIsllllld. Brisbane. 372(45): 1-154. 41 cst.. 4 Figs., JO lycoph Yles (both arboresccnl and shrub-like forms) are ,ab. domi nanl (Lycopodiopsis. Lycopodiophloios). These Gould. R.E. & Delevoryos J. ]977. The biology ofGlossopieris: evidence palaeobolan ical records suggesl a wanner. semiarid interval, from pelrified seed-bearing and pollen-bearing organs. A/cherillgia. and this is apparentl y conlinned by the presence of re d 1:387-399. sandsloncs indicalive of long, dry seasons altcrnating wi th Guerra-Sommer. M. 1989. Megaflorns ocorrentes em horizoniCS associlldo5 more rainy seasons wil h a moderate precipitation. a carvlkl no Rio Grande do Sui. ]V Simp. Sui-bras. de Geologia. Porto Alegre. Brosi!. ACIII Geologico uopo/dnuiu. 29(2):93-122. Aknowledgements - We express our graleful appreciation _-,-....,_-=_ &Cal.lulo Klepzig M . 1993 - Biostratigraphy of lhe to Dr. Robert Wagner (Jardim Bolanico de C6rdoba. Spain) Soulhern Br.uilian Neopalaeoloic Gondwana sequence: a preliminary palaeobotanical approach. 12 Congn!s Imernational de la Stratigrnphie for making the English revision. el Geologie du Carbonifere et Permian. Buenos Aires, Comptt5 Rend"s XII. ]CCP.2:61-72. REFERENCES --,--,--:-cc.,-'; Marques-Toigo. M. & Corna da Silva. Z.C. 1991. Original Biomass and ooal deposition in Southern Bra1.il (Lower A1.cuy. C.L. 1978. Posible signir.cado paleoecol6gico de microfloras Permian. PlUllII' Basin). Bulletill Soc. Geol. Fronct. 162(2):227-37. Gondwanicas del Paleo1.6ico superior. e.speo::ialmente argentinas. Iiolz. M. 1987. L.eques aluviais - Faciologia e ooonfncia em sedimcmilos AmtglrilllllllU. IS (I - 2): 85-95 do Supergrupo Tubar:io (Eopermi300) da Bacia do Parami. no Rio Grande: Backeu~r. Y.: Silveira J.B.R. & Guerra-Sommer. M. 1984. Revis30 da do Sui. ACtu Gcologica uopo/deruia. 25:65 - 104. lafoflont do aflommemo Km 89-90 da Rodovia BR 290. Rio Gntnde Krliu.o;.el. R. ]961. L)"copodiopsis derbyi Renault und einige andere do Sui. Brasil. A'I/I;s. 33" Congrcsso Brusileiro de Gcologiu. Rio de Lycopodialcs aus den Gondwana schiehlen. Po/aconrologruphico B .. Janeiro. SBG. 2:1062·64. 109:62-92. pl. 32-41. Boureau. E. & Doubinger J. 1975. f>leridophylo. In: BOUl·cau. E. Truiti de L.avina. E.l... 1992. Geologia Scdimemar e Palrogcogntfia do Neopermiano Pa/iobollmic. 4 Fasc. I. pl. 1. 772 p. e Eoirilissico(lniervalo KI\7.aniano-Scyllillno)da Bacia do Pllram1.. Cursu Camuhcrs. W. 1869. On Ihe planl remains from Ihe Brn7jlian cool beds de P6s·gradu~lIoem Grociendas. Universidade Federal do Rio Grande with. remarks on Ihe genus Fiemingill'S. Geol. Mag. 6 (58): ]51-155. do SuI. Test de Doulorodo. I6Op. Ca1.1.ulo-Klep1.ig. M. 1979. Esludo do mfoflorn do Membro Morro Pe lado _-,--:-:-__: Nowal1.ld. C.H.: Amujodos Sanlos. M. & l..cao. H.Z. 1985. na sua localidade - lipo. PesquisllJ. 11 :225-303. Ambientes de sed i men l a~lio do Supcrgrupo Tubarao na regiilo de _-=-:--=_. & GuelTll-Sommer. M. 1983. Relationship belween the Cachocira do Sui. RS. ACIII GMlogico Lcopoldcruia. 21 :5-76. Taphoflora of lhe lI arnn! Group. Parnn~ Basin. Southern BTa2.il and the Limo e Cunha. M.C. ]976. Contribuj~ilo A palcontologia cslraligr.ifica do PernIO Carboniferous boundary. D i~i4!me Congn! Imernalional de Grupo Passa Dois no Rio Grande do Su!.Acra Gtologico Lropo/drllsia. Sirnligrnphie el Geologie du Carbonif4!re. Madrid. Espanha. Complt 1(1):1-48. Remlu. 4:395-402. L.undqvist. G. 19 19. Fossile Pflanzen derGlossopltris Flora aus Brasilien. _-,-_-,-,-" & . 1985. Pallleofloristic succession in lhc Kungl. SI·tnska VtUIISWpS Akadtmitru lIalllingar. 60(3):1-36. ParMi Basin. Rio Grande do SuI. Southern Brui!. MemQrius. 6~ Marqucs-Toigo. M. 1988. Palinologia. bioe.slrnligrafia e paleoecologia do COIIgresso wtill()lJmericano dt Gtologia. Bogool. Colombia. 1:177- neopalcoz6ico da Bacia do ParanA nos E. do R.G. Sui e Santa Catarina. 92, Ontsi]. CUT50 de P6s-grad.Iar;io em Geocifncias. Universidade Federnl ____, --::-,..,.--:--= & Marques-Toigo. M. 1980. E5tudo do Rio Gntnde do SuI. Te~ de Doulorado. 259 p. macro e microfloriS'lico do G",po haro~ (Bacia do ParanA). RS. Anais. __--,-,,--...,, &Conia da Silva. Z.C. 1984. On theorigin of gondwanic 31' Cl)ngresso Brasiltiro d~ G~ologia. Balnetrio de CamboriLi. SBG. south Bruilillfltoo! measures. Comun.Sev.GtoI. Portugol. 70(2): 151.00. 5:3027-34. Meyen. S.V. 1987. Fundumtll/olsofPaloeobotan),. Chapmann and Hall. 432 p.

Plale4 (all figures x 700)

Figure I - Plillctalisporitcs grete'lSis forma minor Hart 1965 MP- P 343 Gamerro 1979 MP-P2573 Figure 2 - Calomo5pora sp MP-P 3573 Figure 8 - Crisllliis/wriles sp MP-P 2008 Figure 3 _ Leilri/eles "irtii Tiwari 1965 MP-P 343 Figure 9 - BlllryoclXCus brawlii KU I ~ing 1849 MP-P 3 ]4 Figure 4 - Gfllllu/misporites micronodosus Blllme& 1-bn:IIy 1956MP-P238 Figure 10 - TetrllporilJl/ hor%Sia (Slaplin) Bahne ]963 MP_ P 3402 Figure 5 _ LUlldb/adi!Jpora brazilicflJis (Pan l & Srivaslava) Marque.s-Toigo Figure II - Portaliltsgomlwallellsis Nahuys, Alpern & Ybert 1968 & Pons 1975 MP-P 343 MI' -P 343 Figore 6 - Kraeu51'lisporirn IIpiCllitlll15 Jansonius 1962 MP-P 2007 Figure 12 - Pilas/writes wlno/lis (BallOe & Hennelly) Tiwari & Navale. Figure 7 _ Valla/lsporius arCUll/lU (Marques-Toigo) Archangelsky & 1967 MP-P 340. M. Guerra - Sommer eI al. Plale4

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Mussa, D.: CarvulhQ, R.G. & Santos. P.R. 1980. Esludo eslrnlignilico e Porsch.er. C. 1991. Faciologia da seqUcocia sedimenlar nas folhas de palcotCol6gico em ocontncias fossilfferas da FOITlla~ao InuL Estado Quil~ria e V6n:ea do CapivarilP. R.S. Pesqllisos. 18([):311-43. de 550 Paulo. Brasil. Bolnlfll ICI. USP. 11:142-48. Relallack. G.1. [970. Lale Carboniferous to Midd[e Triassic megafOMil Paim. P.S.G.: Piccoli. A.F_M.: Sal'1uri. J.A.D.: Munaro. P. & Granitoff. W. noras from the Sidney B:l.~in. Geological Sun'ry of New South Wales 1983. Evolu~lIo paleogeogr:!lica do Supergrupo Tubarao na Mea de Bull. 26:384-430. Marillfla Pimenlcl·Fuinal. Gualba, RS. / " SimpOsio Sul-Brrui/eiro de Sch!leider, R.L.: MUhlm:mn. H.: Tommasi. E.: Medeiros. R.A.: Daemon. GeologlO,l'ofto Alcg~. Alus: 140-60. R.E & NoglJClra, A.A. 1974. Revisiocstnuigrtficada Bacia do Paron:i. Ptssoa, R.H.C. 1985. Madeiras gimoospErmicas no Irati do Estadodo Rio Anais. 28' CongrtsSO Brasileiro de GMlogia. Pono Alegre. Brasil. Grande do Sui: observat;Oes sobre a analomia. sistemAtica e SSG. 1:41·65. paleoecologia. Coletllnca de Trabalhos Paleomol6gicos. DNPM. 5lrii' Taylor, N.T. & Taylor E.L. 1989. Amanic Paleobio[ogy _ ilS role in tile Geologia 27. Pu/eolllologio e £Slroligrafla. 2:623-39. reconstruction ofGoodwana. Springer-Verlag cds .. New York. 26[ pp .. Piccoli. A.E.M. 1989. Rela¢es C:SIr:nigrificas emil: as Facies 8ud6 e Suspim 74 m. (Grupo h~) nas folhas de Vila Nova, Lagoa da Meia Lua e Suspim. While, I.e. 1908. Rclal6rio sobre lIS "roal ml'asures" e rochas associad3'l RS. Pesquisas. 12:45-52. do sui do Brusil. HI'I(I{Qrio FirI(Jl tla Comissiiode ESfimasdas Minosde ____ ; Lopes. R.C. & CaITlOl.1.alo. E. 1986. AnAlise paleoambiemal Can'OO /11' Ped", do Brasil. Imprensa National. Rio de Janeiro. p.l· das jazidas de carvlo do Eslado do Rio Gra!lde do SuI. Acta Geologica 300. uopoldensi/J. 22:35·54. Zciller.R. [895. Note sur [8 nore fossile des gisements houillers du Rio ____ : Menegul. R.: Guerra·Sommer. M.: Marques·Toigo. M. & Grande do Sui (Brtsil Meridional). 81111"lin Socieu! Gi-ologiqu .. ill! Fmltce,.1:602·29.lp11l. 8·10. seric 3

.' Iate 5 (all figures x 700)

Figure I . PO/oltieisporill'S sp MP·P 3844 Figure 5 . Pr%liuJI/oxYJlinus sp Mp·P 3844 Figure 2· Pliclllipol/,,"iles ma/obllreltsis (Poloni~ e Sah) FOSler 1975 Figure 6 · 1_U(~d:ispori/es "irkki"t (Poloni~ e Klaus) Klaus 1967 Mp·P 411 MP·P2A Figure 7 . Willi/ina sp Mp·P 343 Figure 3 - Ca/lCniasllcrilt's Ol'/J/IIS (Bose e Kar) Archangelsky e Gamerro Figure 8· Slrill/()pOC/OC(Jrpil~S sp MP·P 2344 1979MI>·P 1534 Figure 9 • f'lIsllco/pill:S sp Mp·P 317 Figure 4 . Proloitol'/Oxypillll$ sp Mp·P 3572 Figure 10· Scite"riIiSipo//t'lIiltS IIlluilllus (Hart) T iwari 1973 Mp· 324. 3 1

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