Devonian miospore successions of Western Gondwana:update and correlation with

Southern Euramerican miospore zones

S. Loboziak a;1, J.H.G. Melo b;�

a U.S.T.L., UPRESA 8014 du CNRS, Sciences de la Terre, 59655 Villeneuve d’Ascq Cedex, Franceb PETROBRAS/CENPES/PDEP/BPA, Cid. Univ., I. Funda‹o, 21949-900 Rio de Janeiro, RJ, Brazil

Received 8 February 2001; accepted 24 July 2001

Abstract

The composition and main biohorizons of Devonian miospore successions of Western Gondwanan regions arereviewed and compared to corresponding miospore zones of Southern Euramerica. The stratigraphic distribution ofthe most significant taxa is recorded in the main investigated basins of North Africa, Saudi Arabia and Brazil.

With the exception of certain forms geographically restricted to Western Gondwana, most of the stratigraphicallyuseful taxa are common to both Southern Euramerica and Western Gondwana. Successions of Western European-defined zones can be recognized and correlated in Western Gondwana regions on the basis of these common indexspecies. Many Devonian miospore biozones defined in Western Europe are well represented in Western Gondwana,whereas some remain unrecorded or poorly characterized. The biozones may alternate with intervals with poorbiostratigraphic resolution which can be attributed to various causative agents. 2 2001 Elsevier Science B.V. Allrights reserved.

Keywords: miospores; biostratigraphy; Devonian; Western Gondwana

1. Introduction

This paper reappraises the Devonian miosporesuccessions of Western Gondwana and correlatesthem with Southern Euramerican biozonations onthe basis of common zonal species. It also high-lights the stratigraphic signi¢cance of associatedendemic Western Gondwanan species recorded

in various areas of North Africa (Hammadah Ba-sin of Tunisia^western Libya and Cyrenaica ofnortheastern Libya), Arabian Gulf region (north-ern and central Saudi Arabia) and South America(Amazon and Parna|¤ba Basins in northern Brazil,and Parana¤ Basin in southern Brazil). In additionto miospore information obtained from thoseareas by the same team of authors, other discretebiostratigraphic data are also taken into account.These concern the Grand Erg occidental and IlliziBasin of the Algerian Sahara, the Al Kufrah Ba-sin of southeastern Libya, the Eastern Elburz ofIran, and the Cordillera Oriental of Bolivia(Fig. 1).

0034-6667 / 01 / $ ^ see front matter 2 2001 Elsevier Science B.V. All rights reserved.PII: S 0 0 3 4 - 6 6 6 7 ( 0 1 ) 0 0 0 9 8 - 7

1 Deceased.* Corresponding author.

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2. Devonian miospore biozonations: a summary

Publications about the stratigraphic distribu-tion of Devonian miospores e¡ectively startedwith the classic work of Naumova (1953) on theRussian Platform in the northeast of the Eur-american Continent. Subsequent studies of vari-ous parts of the Devonian System have beenundertaken in di¡erent parts of the world butespecially in Euramerica.

Two broad zonal concepts have been adoptedin the study of Devonian miospore successions.One of them is the assemblage zone concept,mainly used by Richardson and McGregor

(1986) in their miospore zonation of the OldRed Sandstone Continent and adjacent regions.These authors described assemblage zones whichare de¢ned by a combination of di¡erent criteria,such as the co-occurrence of characteristic taxa,the ¢rst appearance of selected species, and theappearance of selected morphological features.This concept was proposed as the only one whichwould allow correlation on a global scale. Indeed,the Old Red Sandstone Continent and adjacentregions obviously belong to di¡erent phytogeo-graphic provinces, and therefore in order to ac-commodate a maximum of available data it wasnecessary to apply such a comprehensive zonal

Fig. 1. Occurrences of Devonian palyno£oras discussed in the text. Paleogeographic reconstruction after Heckel and Witzke(1979).

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concept. Note that in Eastern Europe, authorsstill favor a concept of Assemblage Zone basedon, and generally named after, the acme of char-acteristic species, whereas the true stratigraphicinceptions of taxa are seldom taken into account.This kind of biozonation is still largely applied toa Devonian phytogeographic area in the easternpart of northern Euramerica.

The other concept is the interval zone, de¢nedon the basis of ¢rst and/or last occurrences ofselected taxa and adopted by Streel et al. (1987)in their biozonation of the type marine Devonianstrata of the Ardenne^Rhenish regions in WesternEurope. Streel et al.’s miospore zonal schemecomprises a series of interval zones entirelyerected within the limits of a single phytogeo-graphic province, to the contrary of the Richard-son and McGregor assemblage zones. The advan-tage of the interval zone concept is that it allowsunequivocal correlation with interval zones basedon varied fossil groups.

As regards the Devonian of Western Gondwa-na, the earliest and most signi¢cant miospore bio-zonations date from the 1970s^1980s. These in-volve the Illizi (Fort-Polignac) Basin of theAlgerian Sahara (Jardine¤ and Yapaudjian, 1968),the Hammadah (Rhadame's) Basin of westernLibya (Massa and Moreau-Benoit, 1976), the Bra-zilian Amazon, Parna|¤ba and Parana¤ Basins(Daemon et al., 1967; Daemon and Contreiras,1971; Daemon, 1974, 1976) and, somewhat later,the Cordillera Oriental of Bolivia (McGregor,1984).

Subsequent miospore investigations have beencarried out in Libya (Paris et al., 1985; Streel etal., 1988; Loboziak and Streel, 1989), Tunisia(Loboziak et al., 1992a), and Brazil (Burjack etal., 1987; Loboziak et al., 1988, 1991, 1992b,1993, 1995). These Western Gondwanan studies,as previously pointed out by McGregor (1984) forBolivian sections, have revealed comparable suc-cessions of ¢rst occurrences of diagnostic speciesof both the Old Red Sandstone Continent andArdenne^Rhenish miospore zonal schemes (Rich-ardson and McGregor, 1986; Streel et al., 1987).Indeed, Loboziak et al. (1989) and Streel et al.(1990) demonstrated that Southern Euramericaand Western Gondwana constituted a single ma-

jor phytogeographic province during the Middleand Late Devonian. Although several endemicmiospore species are restricted to Western Gond-wanan areas (Loboziak et al., 1988; Loboziakand Streel, 1989, 1995a), many of them representuseful zonal markers for the regional biozonation,because their ranges can now be dated and cali-brated by means of associated miospore speciesalso present in Southern Euramerica.

During the past decade palynological joint re-search by Brazilian and Western European work-ers has been carried out in the Devonian of theAmazon and Parna|¤ba Basins of northern Brazil(Rodrigues et al., 1995; Loboziak et al., 1996,1997a,b, 2000a; Melo et al., 1996). During thesame period, palynostratigraphic results werealso obtained in northern and central Saudi Ara-bia (Loboziak and Streel, 1995b; Steemans, 1995;Loboziak, 2000) as part of a Saudi Aramco/CIMPproject.

3. Devonian miospore distribution in WesternGondwana

Fig. 2 summarizes the biozonal schemes pro-posed for the main investigated basins of NorthAfrica, Saudi Arabia and Brazil. Fig. 3 shows themost signi¢cant taxa recorded in these areas, andtheir stratigraphic ranges dated by correlationwith faunally calibrated miospore interval zonesof the Ardenne^Rhenish regions (Streel et al.,1987) and the miospore assemblage zones of theOld Red Sandstone Continent and adjacent re-gions (Richardson and McGregor, 1986). Withthe exception of forms geographically restrictedto Western Gondwana, most of these taxa arecommon to both Southern Euramerica and West-ern Gondwana.

During most of the Early Devonian, i.e. fromLochkovian to the early late Emsian, miosporeswere diverse and rather small-sized. These includeretusoid forms (Retusotriletes, Apiculiretusispora),spores with dominantly biform sculptural ele-ments (Dibolisporites), patinate forms withsmooth or variably sculptured exine (Archaeozo-notriletes, Chelinospora, Cymbosporites), sporeswith a proximo-equatorial crassitude (Aneuro-

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Fig. 2. Miospore zonal schemes proposed for the investigated main Western Gondwana regions and their relationships with regional lithostrati-graphic units, modi¢ed from Loboziak and Melo, 2000, ¢g. 2.

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Fig.3.

Stra

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Plate I. Slides from wells 1-AM-1-AM, 1-CM-1-PA, 1-CM-2-PA, 1-MS-4-AM, 1-NO-6-AM, 2-PC-1-AM, 1-RO-1-AM and CaimaPH-2 (Amazon Basin, Brazil), from outcrops of the Tocantins River Valley (Parna|¤ba Basin, Brazil) and from outcrops of theChapada dos Guimara‹es area (Parana¤ Basin, Brazil) are housed in the palynological slide collection of the Biostratigraphy andPaleoecology Section of Petrobras Research Centre (Cenpes/Divex/Sebipe), Rio de Janeiro, Brazil. Slides from wells A1-69 andMG-1 (Hammadah Basin, Lybia^Tunisia), 2-NA-1-PA (Amazon Basin, Brazil), 2-PM-1-MA (Parna|¤ba Basin, Brazil) and RSP-1(Parana¤ Basin, Brazil) are housed in the palynological collection of the Laboratory of Paleobotany, University of Lille, France.Miospore locations on slides are based on England Finder graticules. Magni¢cation of illustrated specimens: U475.

1. Zonotriletes sp. 3 in Jardine¤ and Yapaudjian, 1969Slide 287.CdG: F35, outcrop 11SD-16, pt 26, Chapada dos Guimara‹es, Parana¤ Basin, Brazil.

2. Dictyotriletes emsiensis (Allen) McGregor, 1973Slide 9305575: K43/4, well 1-AM-1-AM, core 35, Amazon Basin, Brazil.

3. Brochotriletes foveolatus Naumova, 1953Slide 287.CdG: M32/4, outcrop 11SD-16, pt 26, Chapada dos Guimara‹es, Parana¤ Basin, Brazil.

4. Emphanisporites annulatus McGregor, 1961Slide 2483(1): Q35, well MG-1, cuttings sample 2483m, Hammadah Basin, Tunisia.

5, 6. Archaeozonotriletes chulus (Cramer) Richardson and Lister, 19695. Slide 2728(1): Q31, well MG-1, cuttings sample 2728m, Hammadah Basin, Tunisia.6. Slide 2527(1): J24/3, well MG-1, cuttings sample 2527m, Hammadah Basin, Tunisia.7. Synorisporites tripapillatus Richardson and Lister, 1969

Slide 2678(1): O29/4, well MG-1, cuttings sample 2678m, Hammadah Basin, Tunisia.8. Synorisporites libycus Richardson and Ioannides, 1973

Slide 2270(1): S28/2, well MG-1, cuttings sample 2270m, Hammadah Basin, Tunisia.9. Camarozonotriletes sextantii McGregor and Cam¢eld, 1976

Slide 2456(1): X28/2, well MG-1, cuttings sample 2456m, Hammadah Basin, Tunisia.10. Chelinospora ligurata Allen, 1965

Slide 2476(1): Q25/3, well MG-1, cuttings sample 2476m, Hammadah Basin, Tunisia.11. Chelinospora sp.

Slide 1995(1): O25/4, well MG-1, cuttings sample 1995m, Hammadah Basin, Tunisia.12. Chelinospora timanica (Naumova) Loboziak and Streel, 1989

Slide 9701258: X65, well 2-PC-1-AM, core 44/45, Amazon Basin, Brazil.13. Camarozonotriletes? concavus Loboziak and Streel, 1989

Slide 2285(1): U24, well MG-1, cuttings sample 2285m, Hammadah Basin, Tunisia.14. Samarisporites sp. E Streel and Loboziak, 1987

Slide 950645: N46/3, shallow well Caima-PH2, sample 346, Amazon Basin, Brazil.15. Samarisporites triangulatus Allen, 1965

Slide 950672: O64/2, shallow well Caima PH-2, sample 373, Amazon Basin, Brazil.16, 17. Geminospora lemurata Balme emend. Playford, 198316. Slide 950302: V62/4, shallow well Caima PH-2, sample 23, Amazon Basin, Brazil.17. Slide 9603108: Y48, outcrop sample RSO-5, Tocantins River Valley, Parna|¤ba Basin, Brazil.18. Geminospora piliformis Loboziak, Streel and Burjack, 1988

Slide 2178(1): R26, well MG-1, cuttings sample 2178m, Hammadah Basin, Tunisia.19. Teichertospora torquata (Higgs) McGregor and Playford, 1990

Slide 950301: U443, shallow well Caima PH-2, sample 22, Amazon Basin, Brazil.20. Tumulispora malevkensis (Kedo) Turnau, 1978

Slide 9305556: G52/2, well 1-CM-1-PA, core 25, Amazon Basin.21. Rugospora radiata (Jushko) Byvsheva, 1985

Slide 9505380: J59, shallow well Caima PH-2, sample 5a, Amazon Basin, Brazil.22. Rugospora bricei Loboziak and Streel, 1989

Slide A.B.99.25(2): H34/3, well 2-NA-1-PA, core 25, Amazon Basin, Brazil.23. Cymbosporites cyathus Allen, 1965

Slide 2178(3): P44, well MG-1, cuttings sample 2178m, Hammadah Basin, Tunisia.24. Cymbosporites catillus Allen, 1965

Slide 1995(1): S39/3, well MG-1, cuttings sample 1995m, Hammadah Basin, Tunisia.25, 26. Verrucosisporites bulliferus Richardson and McGregor, 198625. Slide 2178(1): V36/2, well MG-1, cuttings sample 2178m, Hammadah Basin, Tunisia.26. Slide 1995(1): R41/2, well MG-1, cuttings sample 1995m, Hammadah Basin, Tunisia.

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spora, Synorisporites), with proximal radial muri(Emphanisporites), and with a foveolate (Brocho-triletes) or reticulate (Dictyotriletes) sculpture(PLATE I).

Richardson and McGregor (1986, p. 10, ¢g. 2)also report prominent £imsy zonate taxa from thelower part of their Emphanisporites micrornatus^Streelispora newportensis Assemblage Zone withinthe early (but not earliest) Gedinnian (Lochko-vian). In the Ardenne^Rhenish regions (Streel etal., 1987, text ¢g. 2; Steemans, 1989, p. 174, text¢g. 220) the zonates, represented by the speciesCirratriradites diaphanus, are noted for the ¢rsttime (Streel et al., 1987, text ¢g. 3) in the equiv-alent MN Oppel Zone, more precisely in theupper part of the phylogenetic Zone E. microrna-tus var. sinuosus (Si), above the chitinozoan Zone29 of Paris (1981).

Small non-spinose zonates have been recordedin miospore assemblages of several WesternGondwana areas, as for example:

^in North Africa, from the Palynozone 1 be-longing to the lower Tadrart Formation of Ham-

madah Basin in western Libya (Massa and Mo-reau-Benoit, 1976, p. 297, text ¢g. 5); from ZonesVc-d in the uppermost Oued Ti¢st Formation ofthe Algerian Illizi Basin (Jardine¤ and Yapaudjian,1968, p. 453, table 1); and from Palynozone 1recorded in thin shaly layers within the TadrartFormation section of well A1-NC 43 between1492 m and 1435 m, in the Al Kufrah Basin ofsoutheastern Libya (Grignani et al., 1991,p. 1172);

^in Brazil, from cuttings at 738^741 m of well2-CA-1-AM, in the topmost part of the Manaca-puru Formation in the Amazon Basin (S. Lobo-ziak and J.H.G. Melo, unpublished data).

Another signi¢cant feature in Lower Devonianpalyno£oras is the ¢rst occurrence of the reticu-late index species Dictyotriletes emsiensis togetherwith the occurrence of Verrucosisporites polygona-lis and Brochotriletes foveolatus as well as the pro-liferation of Dibolisporites spp. (Richardson andMcGregor, 1986, p. 12). Dictyotriletes emsiensis isan eponym of Richardson and McGregor’s (1986)Dictyotriletes emsiensis^Verrucosisporites polygo-

Plate II.

1. Verrucosisporites nitidus Playford, 1964Slide P.B.99.34(1): W42/3, well 2-PM-1-MA, core 34, Parna|¤ba Basin, Brazil.

2. Vallatisporites verrucosus Hacquebard, 1957Slide 9301847: E52/2, 1-CM-2-PA, core 19, Amazon Basin, Brazil.

3. Vallatisporites vallatus Hacquebard, 1957Slide 9701152: S67/2, 2-PC-1-AM, core 33, Amazon Basin, Brazil.

4. Vallatisporites hystricosus (Winslow) Byvsheva, 1985Slide A.B.99.25(1): U31, well 2-NA-1-PA, core 25, Amazon Basin, Brazil.

5. Craspedispora ghadamisensis Loboziak and Streel, 1989Slide 1700(1): O28, well A1-69, cuttings sample 1700m, Hammadah Basin, Lybia.

6. Craspedispora paranaensis Loboziak, Streel and Burjack, 1988Slide 315,30(1): G40/2, well RSP-1, core 315.30, Parana¤ Basin, Brazil.

7. Indotriradites explanatus (Luber) Playford, 1991Slide 9602207: M50/2, well 1-NO-6-AM, core 28, Amazon Basin, Brazil.

8. Grandispora cornuta Higgs, 1975Slide 9606623A: E44, well 1-MS-4-AM, core 27, Amazon Basin, Brazil.

9. Retispora lepidophyta (Kedo) Playford, 1976Slide 9507442: N63/4, well 1-RO-1-AM, core 54, Amazon Basin, Brazil.

10. Grandispora permulta Loboziak, Streel and Melo, 1999Slide 358,60(1): X37, well RSP-1, core 358.30, Parana¤ Basin, Brazil.

11. Grandispora protea (Naumova) Moreau-Benoit, 1980Slide 2638(1): S28, well MG-1, cuttings sample 2638m, Hammadah Basin, Tunisia.

12. Grandispora megaformis (Richardson) McGregor, 1973Slide 1870(1): N38/3, well A1-69, cuttings sample 1870m, Hammadah Basin, Lybia.

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nalis Assemblage Zone, which is an equivalent ofthe Verrucosisporites polygonalis^Dibolisporiteswetteldorfensis (PoW) Oppel Zone de¢ned in theArdenne (Streel et al., 1987, text ¢g. 13). In thislatter region, the ¢rst occurrence of D. emsiensis(Streel et al., 1987, text ¢g. 2; Steemans, 1989,p. 180, text ¢g. 220) characterizes the base ofthe Dictyotriletes emsiensis (E) Interval Zone,the youngest subdivision of the Breconisporitesbreconensis^Emphanisporites zavallatus (BZ) Op-pel Zone which underlies the PoW Biozone. Ac-cording to Streel et al. (2000, p. 18, text ¢g. 8), thebase of Interval Zone E is close to the lowerboundary of the Pragian Stage.

The ¢rst occurrence of Dictyotriletes emsiensishas been pointed out in assemblages of severalWestern Gondwanan areas, as for example:

^in North Africa, in Palynozone 2 of the upperTadrart and lower Ouan Kasa Formations of theHammadah Basin (Massa and Moreau-Benoit,1976, p. 299, text ¢g. 5); in Zone VI of the lowerHassi Tabankorf Formation of the Illizi Basin(Jardine¤ and Yapaudjian, 1968, p. 446, table 1);from Palynozone 2 of wells A1-NC 43 and B1-NC43 in the Al Kufrah Basin (Grignani et al., 1991,p. 1172);

^in Saudi Arabia, in the upper Tawil Forma-tion section of well DMMM-45 (Steemans, 1995,p. 92, text ¢g. 2);

^in Brazil, in the basal Jatapu Formation sec-tion of various wells in the Amazon Basin (S.Loboziak, J.H.G. Melo and P. Steemans, unpub-lished data); in core 23 of well 2-CN-1-SC, withinthe uppermost Furnas Formation of the Parana¤Basin (Loboziak et al., 1995, p. 391);

^in Bolivia, in the younger part of the lowerSanta Rosa Formation of the Tarabuco andLaurani sections of Cordillera Oriental (McGre-gor, 1984, p. 11, text ¢gs. 2 and 3).

The next miospore event is the ¢rst occurrenceof Camarozonotriletes sextantii and Emphanispori-tes annulatus, a well-known radially ribbed specieswith a distal concentric ring. The inception ofthese two taxa de¢nes the base of the Emphanispo-rites annulatus^Camarozonotriletes sextantii As-semblage Zone of Richardson and McGregor’szonation, above the base of Emsian as now rede-¢ned (Yolkin et al., 1997) by the appearance of

the conodont P. kitabicus, close and probablyabove the base of the Dictyotriletes subgranifer(Su) miospore Zone (Streel et al., 2000, p. 16,text ¢g. 7). According to Streel et al. (1987, text¢g. 13), the Emphanisporites annulatus^Camarozo-notriletes sextantii Zone corresponds to the Em-phanisporites annulatus^Brochotriletes bellatulus(AB) and E. foveolatus^Verruciretusispora dubia(FD) Oppel Zones of the Ardenne^Rhenish re-gions.

Camarozonotriletes sextantii is present in NorthAfrica but has not yet been recorded in Brazil. Ithas been reported as ‘Apiculatisporites sp. I;forme a' zone interradiale’ from the Illizi Basinby Jardine¤ and Yapaudjian (1968, p. 447, tableI, pl. 1, ¢gs. 15, 18), in their Zone VIII withinthe lower Alrar Formation. Furthermore, C. sex-tantii was identi¢ed in the Hammadah Basin byLoboziak and Streel (1989, p. 175, text ¢g. 2, pl. I,¢gs. 11 and 12) and Loboziak et al. (1992a, text¢g. 2, pl. II, ¢g. 5), and as Craspedispora craspedaby Massa and Moreau-Benoit (1976, pl. 2, ¢g.15). It is also present in the Tawil and lower Ju-bah Formation sections of several Saudi Arabianwells (Loboziak and Streel, 1995b, p. 105, text ¢g.2; Steemans, 1995, p. 93, text ¢g. 2).

Emphanisporites annulatus is a species wellknown all over the world. It has been recordedin Western Gondwanan regions such as NorthAfrica (Jardine¤ and Yapaudjian, 1968, p. 447, ta-ble I, pl. 1, ¢gs. 4 and 5; Massa and Moreau-Benoit, 1976, p. 300, text ¢g. 5, pl. 3, ¢g. 1; Pariset al., 1985, p. 49, pl. 18, ¢g. 4; Grignani et al.,1991, p. 1173, pl. 10, ¢gs. 5^7; Loboziak et al.,1989, p. 179, text ¢g. 2, pl. I, ¢g. 2; Boumendjel etal., 1988, text ¢g. 2, pl. 1, ¢gs. 11 and 12; Lobo-ziak et al., 1992a, p. 196, text ¢g. 2, pl. I, ¢g. 6),the Middle East (Loboziak and Streel, 1995b,p. 105, text ¢g. 2, pl. I, ¢g. 3), and Bolivia(McGregor, 1984, p. 13, text ¢gs. 2 and 3, pl. 3,¢g. 1). In Brazil its accurate stratigraphic incep-tion is still uncertain because early Emsian strataare either missing or poorly characterized palyno-logically (Fig. 2). This is particularly true in theAmazon and Parna|¤ba Basins (S. Loboziak andJ.H.G. Melo, unpublished data), whereas in theParana¤ Basin the species was used by Daemonet al. (1967, p. 106, table 1, pl. 1 ¢g. 10), under

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the designation Emphanisporites cf. erraticus, tocharacterize their D2b and D3 Intervals attributedto the Emsian ^ Eifelian.

As pointed out by Loboziak and Melo (2000,p. 403), late Emsian micro£oras are distinguishedfrom older Devonian ones through a remarkableincrease in miospore size, and the disappearanceof several smaller forms, mainly reticulate ele-ments well known in the Lower Devonian. Partic-ularly signi¢cant is the appearance of large,prominently spinose zonates and pseudosaccatesbelonging to the Samarisporites/Grandispora com-plex.

The large spinose zonates^pseudosaccates, suchas Grandispora douglastownense, G. megaformis,G. protea and Samarisporites eximius, have theirstratigraphic inception in the transition betweenthe Emphanisporites annulatus^Camarozonotriletessextantii and the overlying G. douglastownense^Ancyrospora eurypterota Assemblages Zones ofRichardson and McGregor (1986, p. 13, ¢g. 2).In the Ardenne^Rhenish regions this same inter-val correlates with the transition between the Em-phanisporites foveolatus^Verruciretusispora dubia(FD) and the overlying Acinosporites apiculatus^Grandispora protea (AP) Oppel Zones of latestEmsian age (Streel et al., 1987, ¢g. 13).

In Western Gondwana similar palyno£oras areabundant in Eifelian and particularly Givetianrocks, and become gradually scarcer higher upin the Frasnian. However, a few taxa like Gran-dispora incognita are known to range well intopalynophase ‘IV’, which is the highest miosporesubdivision of the Frasnian in Western Europe(Streel et al., 1987). Because of their considerablyrestricted stratigraphic distribution, large zonates^pseudosaccates are generally useful in characteriz-ing the late Emsian^early late Frasnian interval.In Western Gondwanan basins they are occasion-ally found as part of reworked Middle Devonianpalyno£oras, mainly within latest Famennian ma-rine or glacio-marine sections bearing Retisporalepidophyta.

Useful miospore events in Western Gondwanacan be recognized on the basis of successive ¢rstoccurrences of characteristic or index specieswithin the in situ range of the large spinose zo-nates^pseudosaccates (Loboziak and Melo, 2000,

p. 403, text ¢g. 3). In ascending order these strati-graphic inceptions concern (Fig. 3):

^the endemic Western Gondwanan zonates^pseudosaccates Craspedispora ghadamisensis,Grandispora gabesensis and particularly G. per-multa, within an interval which in the Ardenne^Rhenish regions corresponds to the AP/Acinospo-rites acanthomammillatus^Densosporites devonicus(AD) transition of late early Eifelian age;

^Geminospora lemurata2, a species of worldwidedistribution, at the base of the Geminospora le-murata (lem) Interval Zone, only slightly abovethe base of the Givetian as de¢ned in WesternEurope by the ¢rst occurrence of the conodontP. hemiansatus ;

^Samarisporites triangulatus and allied conge-neric forms, often associated in Western Gondwa-na regions with the distinctive patinates Cymbo-sporites catillus and C. cyathus, at the base of theSamarisporites triangulatus^Ancyrospora ancyrea(TA) Zone of late early Givetian age;

^Verrucosisporites bulliferus at the base of theVerrucosisporites bulliferus^Cirratriradites jekhov-skyi (BJ) Zone of early Frasnian age;

^Geminospora piliformis, an endemic WesternGondwanan species, within a stratigraphic inter-val equivalent to the late early Frasnian BJ/Ver-rucosisporites bulliferus^Lophozonotriletes media(BM) zonal transition of Western Europe;

^and Rugospora bricei, a small-sized pseudosac-cate species with ¢ne rugulate sculpture, at thebase of the palynophase ‘IV’ of late Frasnian age.

A reappearance of palyno£oras dominated bysmaller-sized miospores is documented slightly be-low the Frasnian/Famennian boundary as de¢nedin Western Europe by the base of the P. triangu-laris conodont Zone. Latest Frasnian - late Fa-mennian miospore assemblages in the Ardenne-Rhenish areas include many small-sized, spinosepseudosaccate species of the genus Grandispora,the successive ¢rst occurrences of which charac-terize the bases of most of the miospore subdivi-sions between the ‘IVc’ palynophase and the

2 In Western Gondwanan basins the entry of G. lemuratacoincides with the proliferation of strongly verrucate and ba-culate patinate forms of the genus Chelinospora ( =Chelino-spora sp. in text ¢g. 3 and pl. 1, ¢g. 11).

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A. verrucosa^V. hystricosus (VH) Oppel Zone(Streel et al., 1987, pp. 220^222, text ¢g. 7, 9).

In Western Gondwanan basins, as noted byLoboziak and Melo (2000, p. 405, text ¢g. 3),the successive inceptions of only three diagnosticspecies are recorded in the aforementioned strati-graphic interval. These are:

^Teichertospora torquata (in the latest Fras-nian), an eponym of the Teichertospora torqua-ta^Grandispora gracilis Assemblage Zone of theOld Red Sandstone Continent biozonation whichis thus far unrecorded in Ardenne^Rhenish re-gions;

^Rugospora radiata, an eponym of the Rugo-spora £exuosa^Grandispora cornuta AssemblageZone of the Old Red Sandstone Continent, which¢rst occurs in the late Famennian at the baseof the Diducites versabilis^Grandispora cornuta(VCo) Oppel Zone in Ardenne^Rhenish regions;

^and Vallatisporites hystricosus, the strati-graphic inception of which, immediately beneaththe uppermost Famennian (or ‘Strunian’), charac-terizes the base of the Vallatisporites pusillites(sensu lato)^Apiculiretusispora fructicosa Assem-blage Zone of the Old Red Sandstone Continent(Richardson and Ahmed, 1988), as well as thebase of the Apiculiretusispora verrucosa^Vallati-sporites hystricosus (VH) Zone in Ardenne^Rhen-ish areas (Maziane et al., 1999).

Unfortunately, the lower and middle Famen-nian interval has not been well characterized paly-nologically in North Africa and the Arabian GulfRegion. Results from the Aouinet Ouenine IVFormation of the Hammadah Basin (Massa andMoreau-Benoit, 1976) and from the upper part ofthe Binem Formation (palynozones 9 and 10) ofthe Al Kufrah Basin (Grignani et al., 1991) arenot reliable because of the absence of diagnosticFamennian miospore species within the describedassemblages and the lack of any faunal control.By contrast, a late Famennian (Fa2c) palyno£orawas recorded in the upper Gazelle Formation ofthe Illizi Basin (Boumendjel et al., 1988) and Cy-renaica (Paris et al., 1985; Streel et al., 1988) onthe basis of Rugospora radiata, locally misidenti-¢ed as Rugospora £exuosa according to Byvsheva(1985).

In the northern Brazilian basins, the index spe-

cies of the latest Frasnian to middle Famennianpalynozones of Ardenne^Rhenish regions are rareor unknown, even along potentially coeval, care-fully investigated rock sections. Possible causes ofthis problem could include the presence of litho-facies unsuitable for miospore preservation (distalmarine black shales), biostratigraphic gaps relatedto sedimentary condensation, or the scarcity ofcontemporary vegetation cover in those areasdue to climatic or paleogeographic constraints.Therefore, the regional identi¢cation of WesternEuropean-de¢ned Famennian miospore zones re-mains troublesome in northern Brazil, at least asregards the interval ranging from ‘IVc’ (‘V’? inthe Parna|¤ba Basin) through the Grandispora gra-cilis^Grandispora famenensis (GF) Oppel Zone.On the other hand, stratigraphically signi¢cantmiospore species with late Frasnian and youngerage, such as Teichertospora torquata and Crassi-spora catenata (as well as Knoxisporites sp. a¡.dedaleus in the Parna|¤ba Basin), are occasionallyfound above the inception of Rugospora bricei.Equivalent sedimentary sections are still unknownin the Parana¤ Basin.

The youngest Devonian characteristic miosporeevent with real global extent is the ¢rst occurrenceof Retispora lepidophyta which characterizes thebase of the latest Famennian (or ‘Strunian’). Itstotal range within the ‘Strunian’ is currently sub-divided into three biozones (Maziane et al., 1999),the bases of which are de¢ned by the ¢rst occur-rences of diagnostic species, viz., Knoxisporitesliteratus (LL), Indotriradites explanatus (LE) andVerrucosisporites nitidus (LN) (Fig. 3, PLATE II).

‘Strunian’ miospore successions have been rela-tively well investigated in North Africa and theArabian Gulf Region giving rise to di¡erent re-gional biozonations. The Retispora lepidophyta^Knoxisporites literatus (LL) Interval Zone wasidenti¢ed by Streel et al. (1988, p. 113, ¢g. 2) inwell A1-37 of Cyrenaica, northeast Libya. NorthAfrican zonal schemes erected by Lanzoni andMagloire (1969), Attar et al. (1980) and Massaet al. (1980) were correlated by Loboziak et al.(2000b, p. 251, ¢g. 3) with biozones of northernBrazilian basins. Elsewhere in Libya, in the AlKufrah Basin (Grignani et al., 1991, p. 1175, ¢g.4), the joint occurrence of Verrucosisporites niti-

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dus, Retispora lepidophyta, Knoxisporites literatus,Indotriradites explanatus and other typical latestFamennian associates characterizes the youngestpart of the Binem Formation (palynozone 11),immediately above palynozones 9 and 10 of sup-posed early to late Famennian age. Based on itsdistinctive miospore assemblage, palynozone 11can be reliably correlated with the LN biozoneof Western Europe. This same assignment, or atleast an attribution to the undi¡erentiated LE^LN zonal span, also applies to the lowest assem-blage of Khosh-Yeilagh in the Ghosnavi-Tilabadcut of Eastern Elburz, northeast Iran (Coquel etal., 1977, pp. 62 and 63).

In northern Brazilian basins, latest Famennianmiospore assemblages are invariably characterizedby the joint occurrence of Retispora lepidophytaand Indotriradites explanatus, and often includealso Vallatisporites verrucosus, Vallatisporites val-latus and Tumulispora malevkensis. Therefore, theLL Zone has not been identi¢ed thus far in any ofthe investigated sections and may be entirely ab-sent in those regions. According to Loboziak andMelo (2000, p. 405) this could indicate a biostrati-graphic, and probably lithological, gap between atleast the VH Zone (or even older parts of theunderlying VCo Zone) and the base of the ‘Stru-nian’ section containing R. lepidophyta. Due tothe scarcity or local absence of Verrucosisporitesnitidus in most of the studied sections, the totalrange of R. lepidophyta cannot be accurately sub-divided in Brazil. Therefore a LE-LN zonal attri-bution is preferably proposed for that entire in-terval.

In the Parana¤ Basin, reworked LE-LN mio-spore assemblages are recorded within Late Car-boniferous glaciogenic rock units containing sac-cate pollen grains. On the other hand, theRetispora lepidophyta palyno£ora reported by Lo-boziak et al. (1995) from diamictites apparentlydevoid of pollen grains (well 2-O-1-PR) coulddocument the ¢rst in situ occurrence of ‘Strunian’miospores in the Parana¤ Basin (Fig. 2).

4. Conclusion

The broad outlines of the successions of West-

ern European miospore zones can be recognizedand correlated in Western Gondwanan regions onthe basis of common index species. Further bio-stratigraphic re¢nement is possible by using co-occurring endemic species with well-controlled in-ceptions or ranges and widespread distributionthroughout those regions. Miospore results areparticularly accurate for parts of the Middle andUpper Devonian, whereas the Lower Devonianbiostratigraphy could still bene¢t from additionalinvestigations of coeval palyno£oras.

Data derived mainly from our Brazilian studies(plus more limited evidence from North Africa)indicate that not all Devonian miospore zonesde¢ned in Western Europe can be recognizedeverywhere in Western Gondwana. Instead, therecorded biozones alternate with intervals withpoor biostratigraphic resolution which can be at-tributed to various causative agents, such as un-suitable lithologies or depositional settings, con-densed sections, sedimentary gaps, or climaticconstraints on the coeval miospore-producingvegetation. The most conspicuous ‘missing’ paly-nozones concern the early Emsian, the Givetian^Frasnian transition, the latest Frasnian^middleFamennian interval, and the early ‘Strunian’.

5. Species listed

Acinosporites lindlarensis Riegel, 1968Archaeozonotriletes chulus (Cramer) Richardson

and Lister, 1969Archaeozonotriletes variabilis Naumova emend.

Allen, 1965Brochotriletes foveolatus Naumova, 1953Camarozonotriletes? concavus Loboziak and

Streel, 1989Camarozonotriletes sextantii McGregor and

Cam¢eld, 1976Cirratriradites diaphanus Steemans, 1989Chelinospora ligurata Allen, 1965Chelinospora timanica (Naumova) Loboziak and

Streel, 1989Cordylosporites marciae Playford and Sat-

terthwait, 1985Cordylosporites spathulatus (Winslow) Playford

and Satterthwait, 1985

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Craspedispora craspeda Allen, 1965Craspedispora ghadamisensis Loboziak and

Streel, 1989Craspedispora paranaensis Loboziak, Streel and

Burjack, 1988Crassispora catenata Higgs, 1975Cymbosporites catillus Allen, 1965Cymbosporites cyathus Allen, 1965Dictyotriletes emsiensis (Allen) McGregor, 1973Diducites mucronatus (Kedo) Van Veen, 1981Emphanisporites annulatus McGregor, 1961Emphanisporites erraticus (Eisenack) McGregor,

1961Geminospora lemurata Balme emend. Playford,

1983Geminospora punctata Owens, 1971Geminospora piliformis Loboziak, Streel and

Burjack, 1988Grandispora cornuta Higgs, 1975Grandispora douglastownense McGregor, 1973Grandispora facilis (Kedo) Avkhimovitch, 1988Grandispora gabesensis Loboziak and Streel,

1989Grandispora incognita (Kedo) McGregor and

Cam¢eld, 1976Grandispora megaformis (Richardson) McGregor,

1973Grandispora permulta Loboziak, Streel and

Melo, 1999Grandispora protea (Naumova) Moreau-Benoit,

1980Indotriradites explanatus (Luber) Playford, 1991Knoxisporites dedaleus (Naumova) Streel, 1977Knoxisporites hederatus (Ishchenko) Playford,

1963Knoxisporites literatus (Waltz) Playford, 1963Leiotriletes struniensis Moreau-Benoit, 1979Retispora lepidophyta (Kedo) Playford, 1976Rugospora bricei Loboziak and Streel, 1989Rugospora £exuosa (Jushko) Streel, 1974Rugospora radiata (Jushko) Byvsheva, 1985Samarisporites eximius (Allen) Loboziak and

Streel, 1989Samarisporites triangulatus Allen, 1965Samarisporites sp. E Streel and Loboziak,

1987Spelaeotriletes granulatus (Kedo) Moreau-Benoit,

1980

Synorisporites libycus Richardson and Ioannides,1973

Synorisporites tripapillatus Richardson and Lis-ter, 1969

Teichertospora torquata (Higgs) McGregor andPlayford, 1990

Tumulispora malevkensis (Kedo) Turnau, 1978Vallatisporites hystricosus (Winslow) Byvsheva,

1985Vallatisporites vallatus Hacquebard, 1957Vallatisporites verrucosus Hacquebard, 1957Verrucosisporites bulliferus Richardson and

McGregor, 1986Verrucosisporites nitidus Playford, 1964Verrucosisporites polygonalis Lanninger, 1968

Acknowledgements

J.H.G.M. publishes with the permision of PE-TROBRAS ^Petro¤ leo Brasileiro S.A.

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