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Miospore events from late earlyto Late Devonian strata of Western Gondwana
M I O S P O R E E V E N T S F R O M LATE EARLY TO LATE D E V O N I A N STRATA OF W E S T E R N GONDWANA
STANISLASL O B O Z I A K & J o s e HENRIQUEG. MELO LOBOZIAK S. & MELO J.H.G. 2000. Miospore events from late Early to Late Devonian strata of Western Gondwana. [Stratigraphie ~v~nementielle des miospores du DSvonien ancien terminal au D~vonien r~cent du Gondwana occidental]. GEOBIOS, 33, 4: 399-407. Villeurbanne, le 31.08.2000. Manuscrit d~pos5 le 23.06.1999; accept~ d5finitivement le 22.11.1999. ABSTRACT - A miospore zonal scheme for late Early - Late Devonian strata of Western Gondwana is presented. It is based on the first occurrences of both distinctive zonal taxa or groups of taxa of the most significative biozonations of Southern Euramerica and endemic Western Gondwanan species. KEYWORDS: MIOSPORES, BIOSTRATIGRAPHY, DEVONIAN, WESTERN GONDWANA. RESUMt~ - Une zonation des miospores du D~vonien ancien terminal au D~vonien r~cent du Gondwana occidental est propos~e. Elle est basSe sur les apparitions ~ la fois de taxons ou groupes de taxons distincts des biozonations les plus significatives d'Euram~rique du Sud et d'esp~ces end~miques du Gondwana occidental. MOTS-CLI~S: MIOSPORES, BIOSTRATIGRAPHIE, DEVONIEN, GONDWANA OCCIDENTAL.
INTRODUCTION D u r i n g the l a s t decade, i n t e n s i v e r e s e a r c h on the Devonian miospore biozonation of W e s t e r n Gondw a n a h a s led to a considerable r e f i n e m e n t of its s t r a t i g r a p h i c subdivision a n d correlation. The a i m of this p a p e r is to p r e s e n t a zonal s c h e m e for late E a r l y to L a t e D e v o n i a n sections of t h a t province u s i n g t h e c r i t e r i a defined in S o u t h e r n E u r a m e r i c a n b i o z o n a t i o n s a n d the occurrence of W e s t e r n G o n d w a n a n species. T h e p r o p o s e d s c h e m e derives f r o m r e c e n t l y concluded a n d ongoing palynological i n v e s t i g a t i o n s c a r r i e d out by t h e s a m e group of w o r k e r s in v a r i o u s a r e a s of N o r t h Africa (Tunisia, Libya), Middle E a s t (Saudi Arabia) a n d S o u t h A m e r i c a (Brazil) (Fig. 1). FIGURE 1 - Occurrences of Devonian palynofloras discussed in
MIOSPORE INVESTIGATIONS IN WESTERN GONDWANA: A BRIEF REVIEW T h e earliest a n d m o s t significant m i o s p o r e synt h e s e s for the D e v o n i a n of W e s t e r n G o n d w a n a w e r e e s t a b l i s h e d d u r i n g the 70's. T h e y m a i n l y concern t h e Illizi (Fort-Polignac) B a s i n of A l g e r i a n Sahara (Jardin~ & Yapaudjian 1968), t h e H a m m a d a h ( R h a d a m ~ s ) B a s i n of w e s t e r n L i b y a (Massa & M o r e a u - B e n o i t 1976) a n d the B r a z i l i a n A m a z o n , P a r n a ~ a a n d P a r a n ~ B a s i n s ( D a e m o n et al. 1967; D a e m o n & C o n t r e i r a s 1971; D a e m o n 1974, 1976).
the text and respective data sources. 1. Amazon Basin, northern Brazil (Daemon & Contreiras 1971; Daemon 1974, 1976; Loboziak et al. 1991, 1993, 1996, 1997a, b; S. Loboziak & J.H.G. Melo, unpublished data). 2. Parna~a Basin, northern Brazil (Daemon 1974, 1976; Loboziak et al. 1992b, 1993; Rodrigues et al. 1995; S. Loboziak & J.H.G. Melo, unpublished data). 3. Paran~ Basin, southern Brazil (Daemon et al. 1967; Daemon 1976; Burjack et al. 1987; Loboziak et a1.,1988, 1995). 4. Grand Erg occidental, Algerian Sahara (Lanzoni & Magloire 1969). 5. Illizi (Fort-Polignac), Algerian Sahara (Jardin~ & Yapaudjian 1968; Attar et al. 1980; Boumendjel et al. 1988). 6. Hammadah (Rhadam~s-Ghadamis) Basin, Tunisia-Libya (Massa & MoreauBenoit 1976; Massa et al. 1980; Loboziak & Streel 1989; Loboziak et al. 1992a). 7. Cyrenaica, Libya (Streel et al. 1988). 8. Al Kufrah Basin, Libya (Grignani et al. 1991). 9. Northern Arabia, Saudi Arabia (Loboziak & Streel 1995; Loboziak in press). 10. Eastern Elburz, Iran (Coquel et al. 1977).
400 Since then several additional studies have been carried out in Libya (Streel et al. 1988; Loboziak & Streel 1989), then in Tunisia (Loboziak et al. 1992a) and Brazil (Burjack et al. 1987; Loboziak et al. 1988, 1991, 1992b, 1993, 1995). These studies have revealed some similarity in the succession of first occurrences of many of the characteristic species used to erect two miospore zonations, viz., those from the Old Red Sandstone Continent and adjacent regions (Richardson & McGregor 1986) and particularly from the Ardenne-Rhenish regions (Streel et al. 1987). They have also demonstrated the potential of recognizing Western European standard miospore zones in the regional Devonian of Western Gondwana. During the past few years, palynological research has been intensively carried out in the Amazon and Parnafba Basins of northern Brazil in the form of a PETROBRAS/U.S.T.L. joint project, the main goal of which was to contribute to a greater refinement of the Devonian and Lower Carboniferous palynozonation in these basins. Some important results obtained from these investigations were the subject of several publications (Rodrigues et al. 1995; Loboziak et al. 1996, 1997a, b).
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During the same period a palynostratigraphic analysis of late Early to Middle Devonian sequences in Northern Saudi Arabia was carried out as a joint Saudi Aramco/CIMP project (Loboziak & Streel 1995b; Loboziak in press). This work has demonstrated that the miospore zonal scheme defined in Western Europe, which has been successfully extended to several parts of Western Gondwana, appears equally applicable to this Middle East region. The biozonal schemes obtained from these miospore investigations in the various Western Gondwanan regions are summarized in figure 2.
SOUTHERN EURAMERICA - WESTERN GONDWANA: A SINGLE PHYTOGEOGRAPHIC PROVINCE IN M I D D L E A N D LATE D E V O N I A N T I M E S
A quantitative approach based on the relative abundance of miospore taxa, first within Western Gondwana Loboziak et al. 1989) then between
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FIGURE 2 - Late Early-Late Devonian miospore biozonal schemes in the main investigated sections of Western Gondwanan regions. Amazon Basin after Loboziak et al. (1991, 1993, 1996, 1997a, b); S. Loboziak & J.H.G. Melo, unpublished data. Parnafba Basin after Loboziak et al. (1992b, 1993); Rodrigues et al. (1995); S. Loboziak & J.H.G. Melo, unpublished data. Paran~ Basin after Burjack et al. (1987); Loboziak et al. (1988, 1995). Northern Hammadah Basin after Loboziak et al. (1992a). Southern Hammadah Basin after Loboziak & Streel (1989). Northern Saudi Arabia after Loboziak & Streel (1995); Loboziak (in press).
401 Western Gendwana and Southern Euramerica (Streel et al. 1990), indicates a remarkable uniformity of land plant vegetation, and therefore of climate, from paleotropical to paleopolar realms during at least the Givetian and Frasnian. Although certain species are restricted to Western Gondwana (Loboziak & Streel 1995a), the forementioned studies demonstrated that Southern Euramerica and Western Gondwana constituted a single phytogeographic province during the Middle and Late Devonian. Several of the endemic species, in the light of biostratigraphic criteria defined in "Western Europe and mainly in the Ardenne-Rhenish regions (Streel et al. 1987), represent good zonal markers in Western Gondwanan areas.
MIOSPORE BIOHORIZONS The main characteristic miospore features of late Early to Late Devonian sections of Western Gondwana are summarized in figure 3 which shows successive miospore-defined events characterized by the first occurrence of one or more well known
taxa. These events are dated by correlation with faunally calibrated interval zones of the ArdenneRhenish regions (Streel et al. 1987) and the interregional assemblage zones of the Old Red Sandstone Continent and adjacent regions (Richardson & McGregor 1986). During most of the Early Devonian, i.e. from Lochkovian to the early late Emsian, miospores were diverse and rather small-sized. The base of Emsian, as recently defined (Yolkin et al. 1997) by the appearance of the conodont P. k i t a b i c u s , has not yet been characterized in terms of miospore data. However it is considered to be somewhere below the first occurrence of E m p h a n i s p o r i t e s a n n u l a t u s and C a m a r o z o n o t r i l e t e s s e x t a n t i i . Their inceptions define the base of the a n n u l a t u s - sext a n t i i Assemblage Zone of Richardson & McGregor's zonation which, according to Streel et al. (1987, fig. 13), corresponds to the E. a n n u l a t u s - B. b e l l a t u l u s (AB) and E. f o v e o l a t u s - V. d u b i a (FD) Oppel Zones of the Streel et al.'s zonation. C. sext a n t i i has not yet been recorded in Brazil, but it is present in North Africa. It has been reported from the Illizi Basin as ' A p i c u l a t i s p o r i t e s sp. I; forme zone interradiale' by Jardin~ & Yapaudjian (1968, pl. 1, fig. 15, 18), from the Hammadah Basin (Lobo-
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FIGURE 3 - Late Early to Late Devonian main miospore events of Western Gondwana.
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FIGURE 4 - 1. Grandispora protea (NAuMOVA)MOREAU~BENOIT,1980. Slide 2639(1): $28, well MG-1, cuttings sample at 2639 m, Hammadah Basin, Tunisia. 2, Grandispora megaformis (RICHARDSON)McGREGOR,1973. Slide 1870(1): N38/3, well A1-69, cuttings sample at 1870 ft, Hammadah Basin, Libya. 3. Grandispora douglastownense McGREGOR,1973. Slide 1550(2): J35/3, well A1-69, cuttings sample at 1550 ft, Hammadah Basin, Libya. 4. Grandispora incognita (KEDo) McGREGOR& CAMFIELD, 1976. Slide 2285(1): 030/4, well MG-1, cuttings sample at 2285 m, Hammadah Basin, Tunisia. 5. Grandispora gabesensis LOBOZL~K& STREEL, 1989. Slide C6(2): V26/3, well MG-1, core 6 at 2161.8 m, Hammadah Basin, Tunisia. 6. Grandispora permulta (DAEMON)LOBOZIAK,STREEL & MELO, 1999. Slide 315,30(1): J40/1, well RSP-1, core at 315.30 m, Paran~ Basin, Brazil. 7. Chelinospora ligurata ALLEN,1965. Slide 2476(1): Q25/3, well MG-1, cuttings sample at 2476 m, Hammadah Basin, Tunisia. 8,9. Chelinospora sp. 8. Slide 1995(1): 025/4, well MG-1, cuttings sample at 1995 m, Hammadah Basin, Tunisia. 9. Slide C5(1): L43/2, well MG-1, core 5 at 2160.6 m, Hammadah Basin, Tunisia. 10. Camarozonotriletes ? concavus LOEOZIAK& STREEL, 1989. Slide 2285(1): U24, well MG-1, cuttings sample at 2285 m, Hammadah Basin, Tunisia. 11. Tumulispora malevkensis (NAuMOVA)TURNAU, 1978. Slide P.B.99.38-40(1): L24, well 2-PM-1-MA, core 38-40 at 1195.6/1198.6 - 1253.4/1256.4 m, Parna~a Basin, Brazil. 12. Verrucosisporites nitidus (NAuMOVA)PLAYFORD,1964. Slide P.B.99.34(1): W42/3, well 2-PM-1-MA, core 34 at 1034.4/1037.4 m, Parnaiba Basin, Brazil. 13. Verrucosisporites bulliferus RICHARDSON• McGREGOR,1986. Slide C6(1): R41/2, well MG-1, core 6 at 2161.8 m, Hammadah Basin, Tunisia. Slides from wells A1-69, MG-1, 2-NA-1-PA, 2-PM-1-MA and RSP-1 are housed in the palynological collection of the Laboratoire de Pal~obotanique, Universit~ de Lille, France. Slides from wells 1-AM-I-AM, 1-AM-7-AM, 1-MS-3-AM and Caima PH2 are housed in the palynological slide collection of the Biostratigraphy and Paleoecology Section of Petrobras Research Centre (Cenpes/Divex/Sebipe), Rio de Janeiro, Brazil. Miospore locations on slides are based on England Finder graticules. Magnification of illustrated specimens: × 500.
403 ziak & Streel 1989; Loboziak et al. 1992a) and also as Craspedispora craspeda by Massa & MoreauBenoit (1976, pl. 2, fig. 15). It is also represented in the Middle East (Loboziak & Streel 1995b). A renewal of the microflora occurred in the late Emsian, mainly due to a dramatic increase in size of the specimens. This event is well documented by the appearance of large, prominently-spined zonates and pseudosaccates belonging to the Samarisporites/Grandispora complex and the disappearance of several smaller forms, particularly reticulate elements well known in the Early Devonian. The large spinose zonates-pseudosaccates first occur and proliferate in the transition between the annulatus - sextantii and the overlying douglastownense - eurypterota Zones of Richardson & Mc Gregor (1986, p. 13, fig. 2). This same interval of latest Emsian age correlates with the transition between the E. foveolatus - V. dubia (FD) and the succeeding A. apiculatus - G. protea (AP) Zones of the Ardenne-Rhenish regions (Streel et al. 1987, fig. 13). This group of miospores is abundant throughout the Eifelian, attains its epibole in the Givetian and becomes gradually rarer in t h e Frasnian. Some specimens, mainly belonging to Grandispora incognita, still persist into palynophase 'IV', which is the highest miospore subdivision of the Frasnian according to the Streel et al. (1987) zonal scheme. Therefore, rich assemblages of large zonates-pseudosaccates are useful in identifying the above mentioned stratigraphic interval. However, they still occur, though quite sparsely, as part of reworked Middle Devonian palynofloras within the latest Famennian marine or glacio-marine sections of Gondwanan regions bearing Retispora lepidophyta. Several successive miospore-defined events are recorded within the in situ range of the large zonates-pseudosaccates. In ascending order, the first event corresponds to the appearance of several endemic zonates and pseudosaccates in Western Gondwana areas, such as Grandispora permulta, G. gabesensis and Craspedispora ghadamisensis. This event occurs within a stratigraphic interval equivalent to the Ardenne-Rhenish AP/A. acanthomammillatus - D. devonicus (AD) transition of late early Eifelian age. Of all these species, G. permulta is the most representative pseudosaccate form in the Middle Devonian microfloras of Western Gondwana. The second event, at the base of the G. lemurata (lem) Interval Zone within the earliest Givetian, corresponds to the first occurrence of Geminospora lemurata, a species widely dispersed all over the world. It is usually well represented in Western Gondwanan regions, and its entry is coeval with the proliferation of highly verrucate and baculate patihate forms of the genus Chelinospora. The next horizon, at the base of the S. triangulatus -A. ancyrea (TA) Zone of late early Givetian age, corresponds to the first occurrence of Samarisporites triangulatus and allied congeneric forms which become widely distributed in the later Middle Devonian and slightly younger strata.
The two next events, of earlier and late early Frasnian ages, correspond to the appearance of miospore taxa bearing tabulate sculpture: Verrucosisporites bulliferus at the base of the V. bulliferus - C.jekhovskyi (BJ) Zone, and Geminospora piliformis, an endemic Western Gondwanan species, within a stratigraphic interval equivalent to the BJ/K bulliferus - L. media (BM) zonal transition of the Ardenne-Rhenish zonation. The succeeding event, of late Frasnian age, occurs at the base of the palynophase 'IV' and corresponds to the first occurrence ofRugospora bricei, a smallsized pseudosaccate species with small/fine rugulate sculpture. A return to palynofloras dominated by smallersized miospores occurs close to, but below, the Frasnian/Famennian boundary and was probably related to new strategies in plant reproduction. From this level to the upper Famennian, miospore assemblages in the Ardenne-Rhenish regions include various small-sized, spinose pseudosaccate species of the genus Grandispora, whose successive first occurrences characterize the bases of most of the miospore subdivisions between the 'IV'c patyaophase and the A. verrucosa - V. hystricosus (VH) Oppel Zone. Unfortunately, the lower to upper Famennian has not been sufficiently documented by us in North Africa and the Middle East. A late Famennian miospore assemblage (Fa2c) was recorded in the Gazelle sup~rieure Formation of the Illizi Basin (Boumendjel et al. 1988). It is characterized by the presence ofRugospora flexuosa, a name incorrectly used, according to Byvsheva (1985), by Western European palynologists for R. radiata. Results from the Aouinet Ouenine IV Formation of the Hammadah Basin of western Libya (Massa & Moreau-Benoit 1976, p. 307-309, fig. 5) and from the upper part of the Binem Formation (palynozones 9 and 10) of the A1 Kufrah Basin in southeastern Libya (Grignani et al. 1991, p. 1174, fig. 4), are not considered reliable because of the absence of diagnostic Famennian species within the described assemblages and the lack of any faunal control. In the northern Brazilian basins, index species of latest Frasnian to middle Famennian palynozones from Ardenne-Rhenish regions are rare or even unknown. Possible causes of such problem may include palynologically unsuitable lithologies (distal marine black shales), biostratigraphic gaps related to sedimentary condensation, or the scarcity of contemporary vegetation cover in those areas due to climatic constraints. Therefore, no Western European-defined miospore zone between palynophase 'V' and Oppel Zone GF have been individualized on an indisputable basis, in spite of very erratic occurrences, above the inception of Rugospora bricei, of other significant miospore species with late Frasnian or younger age, such as Teichertospota torquata and Crassispora catenata, as well as Knoxisporites sp. aff. dedaleus (in the Parna~a Basin). Sedimentary sections of same age are seemingly unrepresented in the Paran~ Basin.
404
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FIGURE 5 - 1. Craspedispora ghadamisensis LOBOZIAK• STREEL, 1989. Slide 358,6(2): $37/4, well RSP-1, core at 358.6 m, Paran~i Basin, Brazil. 2. Craspedispora paranaensis LOBOZL~K,STREEL & BURJACK,1988. Slide 315,30(1): G40/2, well RSP-1, core at 315.30 m, Paran~i Basin, Brazil. 3. Indotriradites explanatus (LUBER)PLAYFORD,1991. Slide 940294: V39/2, well 1-AM-7-AM, core 21 at 1272.10 m, Amazon Basin, Brazil. 4. Retispora lepidophyta (KEDo) PLAYFORD,1976. Slide 9701042: P50, well 1-MS-3-AM, core 24 at 1184/1189 m, Amazon Basin, Brazil. 5. Geminospora punctata OWENS,1971. Slide 2270(1): G40/3, well MG-1, cuttings sample at 2270 m, H a m m a d a h Basin, Tunisia. 6. Geminospora lemurata BALMEemend. PLAYFORD,1983. Slide C6(1): W54/2, well MG-1, core at 2161.8 m, H a m m a d a h Basin, Tunisia. 7. Geminospora piliformis LOBOZIAK,STREEL & BURJACK,1988. Slide 2178(1): R26, well MG-1, cuttings sample at 2178 m, H a m m a d a h basin, Tunisia. 8, 9. Emphanisporites annulatus MCGREGOR, 1961. 8. Slide 2483(1): Q35, well MG-1, cuttings sample at 2483 m, H a m m a d a h Basin, Tunisia. 9. Slide C21(2): E38/1, well MG-1, core at 2450.6 m, H a m m a d a h Basin, Tunisia. 10. Teichertospora cf. torquata (Higgs) McGREGOR & PLAYFORD, 1990. Slide 950301: U44/3, shallow well Caima PH-2, sample 22(3) at 31.12/31.18 m, Amazon Basin, Brazil. 11. Rugospora radiata (JusHKO) BYVSHEVA, 1985. Slide A.B.99.25(2): 026/3, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 12. Rugospora bricei LOSOZlAK& STREEL,1989. Slide A.B.99.25(2): H34/3, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 13. VaUatisporites vallatus HACQUEBARD,1957. Slide 8502263: K49, well 1-AM-I-AM, core 9 at 1236.66/1239.56 m, Amazon Basin, Brazil. 14. Vallatisporites hystricosus (WINSLOW)BYVSHEVA,1985. Slide A.B.99.25(1): U31, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 15. Vallatisporites v e r r u c o s u s HACQUEBARD,1957. Slide P.B.99.38-40(1): H38, well 2-PM-1-MA, core 38-40 at 1195.6/1198.6 - 1253.4/1256.4 m, P a r n a ~ a Basin, Brazil. 16. Camarozonotriletes sextantii MCGREGOR & CAMFIELD,1976. Slide
405 The successive first appearances of only three diagnostic species have been identified within the late Frasnian-late F a m e n n i a n interval. These are: Teichertospora torquata (an eponym of the torquata-gracilis Zone in the Old Red Sandstone Continent zonation but not yet recorded in ArdenneRhenish regions), followed by Rugospora radiata, and at a slightly higher level, Vallatisporites hystricosus. This latter species first appears before Retispora lepidophyta and characterizes the recently defined VH Zone (Maziane et al. 1999). The youngest Devonian characteristic miospore event is the first occurrence ofRetispora lepidophyta at the base of the latest Famennian or 'Strunian'. R. lepidophyta is a well-known cosmopolitan species which permits accurate correlations all over the world. Its total range within the 'Strunian' is subdivided into three biozones, according to the recently revised miospore zonal scheme of the Ardenne region (Maziane et al,.1999). Each of these subdivisions is characterized by the first occurrence of a diagnostic species, viz.: Knoxisporites literatus (LL), Indotriradites explanatus (LE) and Verrucosisporites nitidus (LN). The 'Strunian' interval has been palynologically investigated in various areas of North Africa and the Middle East giving rise to different regional biozonations based on miospore assemblages. Several samples from well A1-37 in Cyrenaica (northeast Libya) have been assigned to the R. lepidophyta - K. literatus (LL) Interval Zone by Streel et al. (1988, p. 113, fig. 2). The biozonations defined in North Africa by Lanzoni & Magloire (1969), Attar et al. (1980) and Massa et al. (1980) were compared to biozones as currently envisaged by the present authors in northern Brazilian basins (Loboziak et al. in press, p. XXX, fig. 3). In the A1 Kufrah Basin (Grignani et al. 1991, p. 1175, fig. 4), the youngest miospore subdivision of the Binem Formation (palynozone 11), which overlies the supposed early to late Famennian palynozones 9 and 10, contains Verrucosisporites nitidus together with R. lepidophyta, K. literatus, I. explanatus and some other typical latest Famennian associates. A LN Biozono age is therefore ensured for this assemblage. This same assignment, or at least a LE-LN zonal range, also applies to the lowest assemblage of Khosh-Yeilagh in the GhosnaviTilabad cut of Eastern Elburz, northeast Iran (Coquel et al. 1977, p. 62, 63). In northern Brazilian basins, most of the latest Famennian miospore assemblages contain I. explanatus and a few other characteristic species, such as Vallatisporites verrucosus, VaUatisporites vallatus and Tumulispora malevkensis, which first occur only in the upper subdivisions of the R. lepidophy-
ta range. Therefore, the LL Zone, could not be identified in any of the investigated samples and may be entirely absent in northern Brazil. This indicates a biostratigraphic, and probably lithological, gap between at least the VH Zone (or still, older parts of the VCo Zone) and the base of the section containing R. lepidophyta. Due to the scarcity, or even local absence, of V. nitidus in most of the investigated sections, the R. lepidophyta range cannot be accurately subdivided. Therefore, a comprehensive LE-LN zonal attribution is preferably proposed for this interval. In the Paran~ Basin, LE-LN miospore assemblages are sometimes found reworked into Late Carboniferous glaciogenic strata bearing saccate pollen grains. Loboziak et al. (1995) reported a R. lepidophyta microflora from diamictites apparently lacking any pollen grains (well 2-O-1-PR). This may represent the first in situ occurrence of 'Strunian' miospores in the Paran~ Basin (Fig. 2).
CONCLUSION The existence, in various areas of Western Gondwana, of miospore biozones first defined in the Devonian of Western Europe, has been demonstrated in several recent publications. It is concluded that endemic Western Gondwanan species, in the light of criteria defined in the Ardenne-Rhenish regions, also represent good biostratigraphic markers for dating and correlating late Early - Late Devonian strata in those areas, in addition to the zonal species of the Southern Euramerican miospore zonations. A c k n o w l e d g e m e n t s - J.H.G. Melo publishes with the permission of PETROBRAS - PetrSleo Brasileiro S.A.
REFERENCES ATTAR A., CANDILIER A.M., COQUEL R. & FOURNIER J. 1980 - Etude palynologique du D~vonien terminal et du Carbonif'ere inf6rieur du bassin d'Illizi (Fort-Polignac), Alg6rie. Bulletin des
Centres de Recherches et d'Exploration - Production d'Elf Aquitaine, 17: 79-147. BOUMENDJEL K., LOBOZIAK S., PARIS F., STEEMANS P. & STREEL M. 1988 - Biostratigraphie des miospores et des chitinozoaires du Silurien sup6rieur et du D~vonien dans le Bassin d'Illizi (S.E. du Sahara alg~rien). Geobios, 32: 329-357. BUR JACK M.I.A., LOBOZIAK S. & STREEL M. 1987 - Quelques donn~es nouvelles sur les miospores d~voniennes du Bassin du Paran~. Sciences G~ologiques Bulletin, 40: 381-391. BYVSHEVAT.V. 1985 - Spores from Tournaisian and Vis~an stages of the Russian Plate. In MENNERV.V. & BYVSHEVAT.V. (eds), Atlas of spores and pollen of Phanerozoic oil - and gasbearing
FIGURE 5 (suite) - 2456(1): X28/2, well MG-1, cuttings sample at 2456 m, Hammadah Basin, Tunisia. 17. Cymbosporites catillus ALLEN, 1965. Slide 1995(1): $39/3, well MG-1, cuttings sample at 1995 m, Hammadah Basin, Tunisia. 18. Samarisporites triangulatus ALLEN, 1965. Slide 2241(1): L32, well MG-1, cuttings sample at 2241 m, Hammadah Basin, Tunisia. 19. Samarisporites sp. E STREEL& LOBOZIAK, 1987. Slide 950672: N46/3, shallow well Caima PH-2, sample 346 at 129.82/129.90 m, Amazon Basin, Brazil. Slides from wells A1-69, MG1, 2-NA-I-PA, 2-PM-I-MA and RSP-1 are housed in the palynological collection of the Laboratoire de Pal6obotanique, Universit~ de Lille, France. Slides t~om wells l-AM-I-AM, I-AM-7-AM, I-MS-3-AM and Caima PH-2 are housed in the palynological slide collection of the Biostratigraphy and Paleoecology Section of Petrobras Research Centre (Cenpes/Divex/Sebipe), Rio de Janeiro, Brazil. Miospore locations on slides are based on England Finder graticules. Magnification of illustrated specimens: x 500.
406 strata of the Russian and Turanian plates. Vsesoyuznyy Nauchno¢-Issledovatel'skuy Geologicheskiy Institut (VNIGNI), Trudy, 253:80-158 [in Russian]. R., LOBOZIAKS., STAMPFLIG. & STAMPFLI-VuILLEB. 1977 Palynologie du D~vonien sup~rieur et du Carbonif'ere inf6rieur dans l'Elburz oriental (Iran Nord-Est). Revue de Micropaldontologie, 20: 59-71. DAEMON R.F. 1974 - Palinomorfos-guias do Devoniano Superior e Carbonifero Inferior das bacias do Amazonas e Parnaiba. Anais da Academia Brasileira de Ci~ncias, 46: 549-587. ..... 1976 - Correlaq~o bioestratigr~flca entre os sedimentos do Siluriano, Devoniano e Carbonffero Inferior das bacias do Amazonas, P a r n a ~ a e Paran~. Anais do 29 ° Congresso Brasileiro de Geologia, Ouro Preto, 2: 189-194. . . . . . & CONTREIRAS C.J.A. 1971 - Zoneamento palinolSgico da Bacia do Amazonas. Anais do 25 ° Congresso Brasileiro de Geologia, S~o Pau]o, 3: 79-88. . . . . . , QUADROS L . P . & SILVA L . C . 1967 - Devonian palynology and biostratigraphy of the Paran~i Basin. In BIGARELLAZ.J. (ed.), Problems in Brazilian Devonian Geology. Boletim Paranaense de Geoci~ncias, 21/22: 99-132. GRIGNANI D., LANZONI E . & ELATRASH H . 1 9 9 1 - P a l a e o z o i c a n d Mesozoic Subsurface Palynostratigraphy in the A1 Kufrah Basin, Libya. In SALEM M . J . , HAMMUDA O . S . & ELIAGOUBI B . A . (eds), The Geology of Libya (Third Symposium on the Geology of Libya, Tripoli, 27-30 sept., 1987), 4: 1159-1227. Elsevier. Amsterdam. JARDINI~ S. & YAPAUDJIANL. 1968 - Lithostratigraphie et palynologie du D~vonien-Gothlandien gr~seux du Bassin de Polignac (Sahara). Revue de l'Institut fran~ais du pgtrole et annales des combustibles liquides, 23: 439-467. LANZONIE. & MAGLOIREL. 1969 - Associations palynologiques et leurs applications stratigraphiques dans le D~vonien supSrieur et Carbonif'ere inf6rieur du Grand Erg Occidental (Sahara alg~rien). Revue de l'Institut fran~ais du pgtrole et annales des combustibles liquides, 24: 441-469. LOBOZIAKS. in press - Middle to early Late Devonian miospore biostratigraphy of Saudi Arabia. GeoArabia. . . . . . , MELO J.H.G., MATSUDA N.S. & QUADROS L.P. 1997a Miospore biostratigraphy of the type Barreirinha Formation (Curu~ Group, Upper Devonian) in the Tapaj6s River area, Amazon Basin, North Brazil. Bulletin des Centres de Recherches et d'Exploration - Production d'Elf Aquitaine, 21: 187-205. . . . . . , MELO J.H.G., QUADROS L . P . & STREEL M . 1997b - Palynological evaluation of the Famennian Protosalvinia (Foerstia) Zone in the Amazon Basin, northern Brazil: a preliminary study. Review of Palaeobotany and Palynology, 96: 3145. . . . . . , MELO J.H.G., RODRIGUES R., STREEL M . , QUADROS L.P. & BARRILARI I.M.R. 1996 - Age and correlation of the Barreirinha Formation (Curu~i Group, Amazon Basin): new evidence from the miospore biostratigraphy. Anais da Academia Brasileira de Ci~ncias, 68: 207-212. . . . . . , MELO J.H.G., STEEMANSP. & BARRILARII.M.R. 1995 - Miospore evidence for pre-Emsian and latest Famennian sedimentation in the Devonian of the Paran~ Basin, South Brazil. Anais da Academia Brasileira de Ci~ncias, 67:391392. . . . . . , MELO J.H.G. & STREEL M. in press - Latest Devonian and Early Carboniferous palynostratigraphy of northern Brazil and North Africa: a proposed integration of western European and Gondwanan miospore biozonations. Bulletin des COQUEL
Centres de Recherches d'Exploration - Production d'Elf Aquitaine, 22. D. 1992a - Biostratigraphie par miospores du D~vonien inf~rieur ~ sup~rieur du sondage MG-1 (Bassin d'Hammadah, Tunisie) - Comparaison avec les donn~es des faunes. Review of Palaeobotany and Palynology, 74: 193-205. . . . . . ~,~STREELM. 1989 - Middle-Upper Devonian miospores from the Ghadamis Basin (Tunisia-Libya): systematics and stratigraphy. Review of Palaeobotany and Palynology, 58: 173196. . . . . . & STREEL M. 1995a - West Gondwanian aspects of the Middle and Upper Devonian miospore zonation in North Africa and Brazil. Review of Palaeoboany and Palynology, 86: 147-155.
& STREELM. 1995b - Late Lower and Middle Devonian miospores from Saudi Arabia. Review of Palaeobotany and Palynology, 89: 105-113. . . . . . , STREELM. & BURJACKM.I.A. 1988 - Miospores du D~vonien moyen et sup~rieur du bassin du Paran~, Br~sil: syst~matique et stratigraphie. Sciences ggologiques, 41: 351-377. ..... , STREELM. • BURJACKM.I.A. 1989 - D~ductions pal~oclimatiques d'une comparaison entre les assemblages de miospores du D~vonien moyen et sup~rieur de Libye et du Br~sil. Geobios, 22: 247-351. ..... , STREELM., CAPUTOM. V. & MELO J.H.G. 1991 - Evidence of West European defined miospore zones in the uppermost Devonian and Lower Carboniferous of the Amazonas Basin (Brazil). Geobios, 24: 5-11. ..... , STREEL M., CAPUTO M.V. & MELO J.H.G. 1992b - Middle Devonian to Lower Carboniferous miospore stratigraphy in the Central P a r n a ~ a Basin (Brazil). Annales de la Socidtg Ggologique de Belgique, 115: 215-226. ..... , STREEL M., CAPUTO M.V. & MELO J.H.G. 1993 - Middle Devonian to Lower Carboniferous miospores from selected boreholes in Amazonas and P a r n a ~ a basins (Brazil): additional data, synthesis, and correlation. Documents des Laboratoires de Ggologie de la Facultg de Lyon, 125: 277-289. MASSAD. & MOREAU-BENOITA. 1976 - Essai de synth~se stratigraphique et palynologique du Syst~me D6vonien en Libye occidentale. Revue de l'Institut fran~ais du pgtrole, 31: 287333. ..... , COQUELR., LOBOZIAKS. & TAUGOURDEAU-LANTZJ. 1980 - Essai de synth~se stratigraphique et palynologique du Carbonif'ere en Libye occidentale. Annales de la Socigtd ggologique du Nord, 99: 429-442. MAZIANE N., HIGGS K. & STREEL M. 1999 - Revision of the late Famennian miospore zonation scheme in eastern Belgium. Journal of Micropaleontology, 18: 17-25. RICHARDSONJ.B. & McGREGORD.C. 1986 - Silurian and Devonian spore zones of the Old Red Sandstone Continent and adjacent regions. Geological Survey of Canada, 364: 1-79. RODRIGUES R . , LOBOZIAK S., MELO J.H.G. & ALVES D.B. 1995 Geochemical characterization and miospore biochronostratigraphy of the Frasnian anoxic event in the P a r n a ~ a Basin, Northeast Brazil. Bulletin des Centres de Recherches d'Exploration - Production d'Elf Aquitaine, 19: 319-327. STREEL M., FAIRON-DEMARETM. & LOBOZIAKS. 1990 - GivetianF r a s n i a n phytogeography of Euramerica and western Gondwana based on miospore distribution. In McKERROW W.S. ~,5 SCOTESE C.R. (eds), Palaeozoic Palaeogeography and Biogeography. The Geological Society Memoir, 12: 291-296. ..... , HIGGS K., LOEOZIhKS., RIEGELW. & STEEMANSP. 1987 - Spore stratigraphy and correlation with faunas and floras in the type marine Devonian of the Ardenne-Rhenish region. Review of Palaeobotany and Palynology, 50: 211-229. ..... , PARISF., RIEGELW. & VANGESTAINEM. 1988 - Acritarch, chitinozoan and spore stratigraphy from the Middle and Upper Devonian subsurface of Northeast Libya. In EL-ARNAUTIA . , OWENSS. & THUSUB. (eds), Palynostratigraphy of Northeast Libya, Benghazi-Libya. Garyounis University Publications, Benghazi: 111-128. YOLKIN E.A., KIM A.I., WEDDIGE K., TALENT J.A. & HOUSE M.R. 1997 - Definition of the Pragian/Emsian Stage boundary. Episodes, 20: 235-240. .....
S. L O B O Z I A K Universit~ des Sciences et Technologies de Lille UPRESA 8014 du CNRS, Sciences de la Terre F-59655 Villeneuve d'Ascq cedex
. . . . . , STEEMANS P., STREEL M . & VACHARD
JoH.G. M E L O Petrobras/CenpesfDivex/Sebipe Cid. Univ., Q. 7, I. Fund~o 21949-900 Rio de Janeiro, RJ, Brazil
APPENDIX
A. SPECIES
LISTED
Camarozonotriletes ? concavus LOBOZIAK& S T R E E L , 1989 Camarozonotriletes sextantii MCGREGOR8: CAMFIELD, 1976 Chelinospora ligurata ALLEN, 1965
407 Craspedispora ghadamisensis LOBOZIAK& STREEL,1989 Craspedispora paranaensis LOBOZIAK,STREEL & BURJACK, 1988 Crassispora catenata HIGGS,1975 Cymbosporites catillus ALLEN, 1965 Emphanisporites annulatus McGREGOR,1961 Geminospora lemurata BALMEemend. PLAYFORO,1983 Geminospora punctata OWENS,1971 Geminospora piliformis LOBOZIAK,STREEL& BURJAEK,1988 Grandispora douglastownense McGREGOR, 1973 Grandispora gabesensis LOBOZL~K& STREEL,1989 Grandispora incognita (KEDo) McGREGOR & CAMFIELD,1976 Grandispora megaformis (RICHARDSON)McGREGOR,1973 Grandispora permulta LOBOZIAK,STREEL & MELO,1999 Grandispora protea (NAUMOVA)MOREAU-BENoIT,1980 Indotriradites explanatus (LuBER)PLAYFORD,1991
Knoxisporites dedaleus (NAUMOVA)STREEL,1977 Knoxisporites literatus (WALTZ)I%AYFORD,1963 Retispora lepidophyta (KEDo) PLAYFORD,1976 Rugospora bricei LOBOZlAK& STREEL,1989 Rugospora flexuosa (JUSHKO)STREEL,1974 Rugospora radiata (JUSHKO)BYVSHEVA,1985 Samarisporites triangulatus ALLEN,1965 Samarisporites sp. E STREEL & LOBOZIAK,1987 Teichertospora torquata (Higgs) McGREGOR • PLAYFORD,1990 Tumulispora malevkensis (KEDo) TURNAU,1978 Vallatisporites hystricosus (WINSLOW)BYVSHEVA,1985 Vallatisporites vallatus HACQUEBARD,1957 VaUatisporites v e r r u c o s u s HACQUEBARD,1957 Verrucosisporites bulliferus RICHARDSON& McGREGOR,1986 Verrucosisporites nitidus PLAYFORD,1964
STANISLASL O B O Z I A K & J o s e HENRIQUEG. MELO LOBOZIAK S. & MELO J.H.G. 2000. Miospore events from late Early to Late Devonian strata of Western Gondwana. [Stratigraphie ~v~nementielle des miospores du DSvonien ancien terminal au D~vonien r~cent du Gondwana occidental]. GEOBIOS, 33, 4: 399-407. Villeurbanne, le 31.08.2000. Manuscrit d~pos5 le 23.06.1999; accept~ d5finitivement le 22.11.1999. ABSTRACT - A miospore zonal scheme for late Early - Late Devonian strata of Western Gondwana is presented. It is based on the first occurrences of both distinctive zonal taxa or groups of taxa of the most significative biozonations of Southern Euramerica and endemic Western Gondwanan species. KEYWORDS: MIOSPORES, BIOSTRATIGRAPHY, DEVONIAN, WESTERN GONDWANA. RESUMt~ - Une zonation des miospores du D~vonien ancien terminal au D~vonien r~cent du Gondwana occidental est propos~e. Elle est basSe sur les apparitions ~ la fois de taxons ou groupes de taxons distincts des biozonations les plus significatives d'Euram~rique du Sud et d'esp~ces end~miques du Gondwana occidental. MOTS-CLI~S: MIOSPORES, BIOSTRATIGRAPHIE, DEVONIEN, GONDWANA OCCIDENTAL.
INTRODUCTION D u r i n g the l a s t decade, i n t e n s i v e r e s e a r c h on the Devonian miospore biozonation of W e s t e r n Gondw a n a h a s led to a considerable r e f i n e m e n t of its s t r a t i g r a p h i c subdivision a n d correlation. The a i m of this p a p e r is to p r e s e n t a zonal s c h e m e for late E a r l y to L a t e D e v o n i a n sections of t h a t province u s i n g t h e c r i t e r i a defined in S o u t h e r n E u r a m e r i c a n b i o z o n a t i o n s a n d the occurrence of W e s t e r n G o n d w a n a n species. T h e p r o p o s e d s c h e m e derives f r o m r e c e n t l y concluded a n d ongoing palynological i n v e s t i g a t i o n s c a r r i e d out by t h e s a m e group of w o r k e r s in v a r i o u s a r e a s of N o r t h Africa (Tunisia, Libya), Middle E a s t (Saudi Arabia) a n d S o u t h A m e r i c a (Brazil) (Fig. 1). FIGURE 1 - Occurrences of Devonian palynofloras discussed in
MIOSPORE INVESTIGATIONS IN WESTERN GONDWANA: A BRIEF REVIEW T h e earliest a n d m o s t significant m i o s p o r e synt h e s e s for the D e v o n i a n of W e s t e r n G o n d w a n a w e r e e s t a b l i s h e d d u r i n g the 70's. T h e y m a i n l y concern t h e Illizi (Fort-Polignac) B a s i n of A l g e r i a n Sahara (Jardin~ & Yapaudjian 1968), t h e H a m m a d a h ( R h a d a m ~ s ) B a s i n of w e s t e r n L i b y a (Massa & M o r e a u - B e n o i t 1976) a n d the B r a z i l i a n A m a z o n , P a r n a ~ a a n d P a r a n ~ B a s i n s ( D a e m o n et al. 1967; D a e m o n & C o n t r e i r a s 1971; D a e m o n 1974, 1976).
the text and respective data sources. 1. Amazon Basin, northern Brazil (Daemon & Contreiras 1971; Daemon 1974, 1976; Loboziak et al. 1991, 1993, 1996, 1997a, b; S. Loboziak & J.H.G. Melo, unpublished data). 2. Parna~a Basin, northern Brazil (Daemon 1974, 1976; Loboziak et al. 1992b, 1993; Rodrigues et al. 1995; S. Loboziak & J.H.G. Melo, unpublished data). 3. Paran~ Basin, southern Brazil (Daemon et al. 1967; Daemon 1976; Burjack et al. 1987; Loboziak et a1.,1988, 1995). 4. Grand Erg occidental, Algerian Sahara (Lanzoni & Magloire 1969). 5. Illizi (Fort-Polignac), Algerian Sahara (Jardin~ & Yapaudjian 1968; Attar et al. 1980; Boumendjel et al. 1988). 6. Hammadah (Rhadam~s-Ghadamis) Basin, Tunisia-Libya (Massa & MoreauBenoit 1976; Massa et al. 1980; Loboziak & Streel 1989; Loboziak et al. 1992a). 7. Cyrenaica, Libya (Streel et al. 1988). 8. Al Kufrah Basin, Libya (Grignani et al. 1991). 9. Northern Arabia, Saudi Arabia (Loboziak & Streel 1995; Loboziak in press). 10. Eastern Elburz, Iran (Coquel et al. 1977).
400 Since then several additional studies have been carried out in Libya (Streel et al. 1988; Loboziak & Streel 1989), then in Tunisia (Loboziak et al. 1992a) and Brazil (Burjack et al. 1987; Loboziak et al. 1988, 1991, 1992b, 1993, 1995). These studies have revealed some similarity in the succession of first occurrences of many of the characteristic species used to erect two miospore zonations, viz., those from the Old Red Sandstone Continent and adjacent regions (Richardson & McGregor 1986) and particularly from the Ardenne-Rhenish regions (Streel et al. 1987). They have also demonstrated the potential of recognizing Western European standard miospore zones in the regional Devonian of Western Gondwana. During the past few years, palynological research has been intensively carried out in the Amazon and Parnafba Basins of northern Brazil in the form of a PETROBRAS/U.S.T.L. joint project, the main goal of which was to contribute to a greater refinement of the Devonian and Lower Carboniferous palynozonation in these basins. Some important results obtained from these investigations were the subject of several publications (Rodrigues et al. 1995; Loboziak et al. 1996, 1997a, b).
A
B
During the same period a palynostratigraphic analysis of late Early to Middle Devonian sequences in Northern Saudi Arabia was carried out as a joint Saudi Aramco/CIMP project (Loboziak & Streel 1995b; Loboziak in press). This work has demonstrated that the miospore zonal scheme defined in Western Europe, which has been successfully extended to several parts of Western Gondwana, appears equally applicable to this Middle East region. The biozonal schemes obtained from these miospore investigations in the various Western Gondwanan regions are summarized in figure 2.
SOUTHERN EURAMERICA - WESTERN GONDWANA: A SINGLE PHYTOGEOGRAPHIC PROVINCE IN M I D D L E A N D LATE D E V O N I A N T I M E S
A quantitative approach based on the relative abundance of miospore taxa, first within Western Gondwana Loboziak et al. 1989) then between
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Miosporebiozonesafter: (1) Streel et al. 1987 (modified) (2) Richardson& McGregor1986 (modified)
Missingor condensed sedimentarysections
FIGURE 2 - Late Early-Late Devonian miospore biozonal schemes in the main investigated sections of Western Gondwanan regions. Amazon Basin after Loboziak et al. (1991, 1993, 1996, 1997a, b); S. Loboziak & J.H.G. Melo, unpublished data. Parnafba Basin after Loboziak et al. (1992b, 1993); Rodrigues et al. (1995); S. Loboziak & J.H.G. Melo, unpublished data. Paran~ Basin after Burjack et al. (1987); Loboziak et al. (1988, 1995). Northern Hammadah Basin after Loboziak et al. (1992a). Southern Hammadah Basin after Loboziak & Streel (1989). Northern Saudi Arabia after Loboziak & Streel (1995); Loboziak (in press).
401 Western Gendwana and Southern Euramerica (Streel et al. 1990), indicates a remarkable uniformity of land plant vegetation, and therefore of climate, from paleotropical to paleopolar realms during at least the Givetian and Frasnian. Although certain species are restricted to Western Gondwana (Loboziak & Streel 1995a), the forementioned studies demonstrated that Southern Euramerica and Western Gondwana constituted a single phytogeographic province during the Middle and Late Devonian. Several of the endemic species, in the light of biostratigraphic criteria defined in "Western Europe and mainly in the Ardenne-Rhenish regions (Streel et al. 1987), represent good zonal markers in Western Gondwanan areas.
MIOSPORE BIOHORIZONS The main characteristic miospore features of late Early to Late Devonian sections of Western Gondwana are summarized in figure 3 which shows successive miospore-defined events characterized by the first occurrence of one or more well known
taxa. These events are dated by correlation with faunally calibrated interval zones of the ArdenneRhenish regions (Streel et al. 1987) and the interregional assemblage zones of the Old Red Sandstone Continent and adjacent regions (Richardson & McGregor 1986). During most of the Early Devonian, i.e. from Lochkovian to the early late Emsian, miospores were diverse and rather small-sized. The base of Emsian, as recently defined (Yolkin et al. 1997) by the appearance of the conodont P. k i t a b i c u s , has not yet been characterized in terms of miospore data. However it is considered to be somewhere below the first occurrence of E m p h a n i s p o r i t e s a n n u l a t u s and C a m a r o z o n o t r i l e t e s s e x t a n t i i . Their inceptions define the base of the a n n u l a t u s - sext a n t i i Assemblage Zone of Richardson & McGregor's zonation which, according to Streel et al. (1987, fig. 13), corresponds to the E. a n n u l a t u s - B. b e l l a t u l u s (AB) and E. f o v e o l a t u s - V. d u b i a (FD) Oppel Zones of the Streel et al.'s zonation. C. sext a n t i i has not yet been recorded in Brazil, but it is present in North Africa. It has been reported from the Illizi Basin as ' A p i c u l a t i s p o r i t e s sp. I; forme zone interradiale' by Jardin~ & Yapaudjian (1968, pl. 1, fig. 15, 18), from the Hammadah Basin (Lobo-
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Miospore biozones after: (1) Streel et al. 1987 (modified) (2) Richardson & McGregor 1986 (modified)
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FIGURE 3 - Late Early to Late Devonian main miospore events of Western Gondwana.
402
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~
) I
FIGURE 4 - 1. Grandispora protea (NAuMOVA)MOREAU~BENOIT,1980. Slide 2639(1): $28, well MG-1, cuttings sample at 2639 m, Hammadah Basin, Tunisia. 2, Grandispora megaformis (RICHARDSON)McGREGOR,1973. Slide 1870(1): N38/3, well A1-69, cuttings sample at 1870 ft, Hammadah Basin, Libya. 3. Grandispora douglastownense McGREGOR,1973. Slide 1550(2): J35/3, well A1-69, cuttings sample at 1550 ft, Hammadah Basin, Libya. 4. Grandispora incognita (KEDo) McGREGOR& CAMFIELD, 1976. Slide 2285(1): 030/4, well MG-1, cuttings sample at 2285 m, Hammadah Basin, Tunisia. 5. Grandispora gabesensis LOBOZL~K& STREEL, 1989. Slide C6(2): V26/3, well MG-1, core 6 at 2161.8 m, Hammadah Basin, Tunisia. 6. Grandispora permulta (DAEMON)LOBOZIAK,STREEL & MELO, 1999. Slide 315,30(1): J40/1, well RSP-1, core at 315.30 m, Paran~ Basin, Brazil. 7. Chelinospora ligurata ALLEN,1965. Slide 2476(1): Q25/3, well MG-1, cuttings sample at 2476 m, Hammadah Basin, Tunisia. 8,9. Chelinospora sp. 8. Slide 1995(1): 025/4, well MG-1, cuttings sample at 1995 m, Hammadah Basin, Tunisia. 9. Slide C5(1): L43/2, well MG-1, core 5 at 2160.6 m, Hammadah Basin, Tunisia. 10. Camarozonotriletes ? concavus LOEOZIAK& STREEL, 1989. Slide 2285(1): U24, well MG-1, cuttings sample at 2285 m, Hammadah Basin, Tunisia. 11. Tumulispora malevkensis (NAuMOVA)TURNAU, 1978. Slide P.B.99.38-40(1): L24, well 2-PM-1-MA, core 38-40 at 1195.6/1198.6 - 1253.4/1256.4 m, Parna~a Basin, Brazil. 12. Verrucosisporites nitidus (NAuMOVA)PLAYFORD,1964. Slide P.B.99.34(1): W42/3, well 2-PM-1-MA, core 34 at 1034.4/1037.4 m, Parnaiba Basin, Brazil. 13. Verrucosisporites bulliferus RICHARDSON• McGREGOR,1986. Slide C6(1): R41/2, well MG-1, core 6 at 2161.8 m, Hammadah Basin, Tunisia. Slides from wells A1-69, MG-1, 2-NA-1-PA, 2-PM-1-MA and RSP-1 are housed in the palynological collection of the Laboratoire de Pal~obotanique, Universit~ de Lille, France. Slides from wells 1-AM-I-AM, 1-AM-7-AM, 1-MS-3-AM and Caima PH2 are housed in the palynological slide collection of the Biostratigraphy and Paleoecology Section of Petrobras Research Centre (Cenpes/Divex/Sebipe), Rio de Janeiro, Brazil. Miospore locations on slides are based on England Finder graticules. Magnification of illustrated specimens: × 500.
403 ziak & Streel 1989; Loboziak et al. 1992a) and also as Craspedispora craspeda by Massa & MoreauBenoit (1976, pl. 2, fig. 15). It is also represented in the Middle East (Loboziak & Streel 1995b). A renewal of the microflora occurred in the late Emsian, mainly due to a dramatic increase in size of the specimens. This event is well documented by the appearance of large, prominently-spined zonates and pseudosaccates belonging to the Samarisporites/Grandispora complex and the disappearance of several smaller forms, particularly reticulate elements well known in the Early Devonian. The large spinose zonates-pseudosaccates first occur and proliferate in the transition between the annulatus - sextantii and the overlying douglastownense - eurypterota Zones of Richardson & Mc Gregor (1986, p. 13, fig. 2). This same interval of latest Emsian age correlates with the transition between the E. foveolatus - V. dubia (FD) and the succeeding A. apiculatus - G. protea (AP) Zones of the Ardenne-Rhenish regions (Streel et al. 1987, fig. 13). This group of miospores is abundant throughout the Eifelian, attains its epibole in the Givetian and becomes gradually rarer in t h e Frasnian. Some specimens, mainly belonging to Grandispora incognita, still persist into palynophase 'IV', which is the highest miospore subdivision of the Frasnian according to the Streel et al. (1987) zonal scheme. Therefore, rich assemblages of large zonates-pseudosaccates are useful in identifying the above mentioned stratigraphic interval. However, they still occur, though quite sparsely, as part of reworked Middle Devonian palynofloras within the latest Famennian marine or glacio-marine sections of Gondwanan regions bearing Retispora lepidophyta. Several successive miospore-defined events are recorded within the in situ range of the large zonates-pseudosaccates. In ascending order, the first event corresponds to the appearance of several endemic zonates and pseudosaccates in Western Gondwana areas, such as Grandispora permulta, G. gabesensis and Craspedispora ghadamisensis. This event occurs within a stratigraphic interval equivalent to the Ardenne-Rhenish AP/A. acanthomammillatus - D. devonicus (AD) transition of late early Eifelian age. Of all these species, G. permulta is the most representative pseudosaccate form in the Middle Devonian microfloras of Western Gondwana. The second event, at the base of the G. lemurata (lem) Interval Zone within the earliest Givetian, corresponds to the first occurrence of Geminospora lemurata, a species widely dispersed all over the world. It is usually well represented in Western Gondwanan regions, and its entry is coeval with the proliferation of highly verrucate and baculate patihate forms of the genus Chelinospora. The next horizon, at the base of the S. triangulatus -A. ancyrea (TA) Zone of late early Givetian age, corresponds to the first occurrence of Samarisporites triangulatus and allied congeneric forms which become widely distributed in the later Middle Devonian and slightly younger strata.
The two next events, of earlier and late early Frasnian ages, correspond to the appearance of miospore taxa bearing tabulate sculpture: Verrucosisporites bulliferus at the base of the V. bulliferus - C.jekhovskyi (BJ) Zone, and Geminospora piliformis, an endemic Western Gondwanan species, within a stratigraphic interval equivalent to the BJ/K bulliferus - L. media (BM) zonal transition of the Ardenne-Rhenish zonation. The succeeding event, of late Frasnian age, occurs at the base of the palynophase 'IV' and corresponds to the first occurrence ofRugospora bricei, a smallsized pseudosaccate species with small/fine rugulate sculpture. A return to palynofloras dominated by smallersized miospores occurs close to, but below, the Frasnian/Famennian boundary and was probably related to new strategies in plant reproduction. From this level to the upper Famennian, miospore assemblages in the Ardenne-Rhenish regions include various small-sized, spinose pseudosaccate species of the genus Grandispora, whose successive first occurrences characterize the bases of most of the miospore subdivisions between the 'IV'c patyaophase and the A. verrucosa - V. hystricosus (VH) Oppel Zone. Unfortunately, the lower to upper Famennian has not been sufficiently documented by us in North Africa and the Middle East. A late Famennian miospore assemblage (Fa2c) was recorded in the Gazelle sup~rieure Formation of the Illizi Basin (Boumendjel et al. 1988). It is characterized by the presence ofRugospora flexuosa, a name incorrectly used, according to Byvsheva (1985), by Western European palynologists for R. radiata. Results from the Aouinet Ouenine IV Formation of the Hammadah Basin of western Libya (Massa & Moreau-Benoit 1976, p. 307-309, fig. 5) and from the upper part of the Binem Formation (palynozones 9 and 10) of the A1 Kufrah Basin in southeastern Libya (Grignani et al. 1991, p. 1174, fig. 4), are not considered reliable because of the absence of diagnostic Famennian species within the described assemblages and the lack of any faunal control. In the northern Brazilian basins, index species of latest Frasnian to middle Famennian palynozones from Ardenne-Rhenish regions are rare or even unknown. Possible causes of such problem may include palynologically unsuitable lithologies (distal marine black shales), biostratigraphic gaps related to sedimentary condensation, or the scarcity of contemporary vegetation cover in those areas due to climatic constraints. Therefore, no Western European-defined miospore zone between palynophase 'V' and Oppel Zone GF have been individualized on an indisputable basis, in spite of very erratic occurrences, above the inception of Rugospora bricei, of other significant miospore species with late Frasnian or younger age, such as Teichertospota torquata and Crassispora catenata, as well as Knoxisporites sp. aff. dedaleus (in the Parna~a Basin). Sedimentary sections of same age are seemingly unrepresented in the Paran~ Basin.
404
3
~
~,, •
,~•
9
•
FIGURE 5 - 1. Craspedispora ghadamisensis LOBOZIAK• STREEL, 1989. Slide 358,6(2): $37/4, well RSP-1, core at 358.6 m, Paran~i Basin, Brazil. 2. Craspedispora paranaensis LOBOZL~K,STREEL & BURJACK,1988. Slide 315,30(1): G40/2, well RSP-1, core at 315.30 m, Paran~i Basin, Brazil. 3. Indotriradites explanatus (LUBER)PLAYFORD,1991. Slide 940294: V39/2, well 1-AM-7-AM, core 21 at 1272.10 m, Amazon Basin, Brazil. 4. Retispora lepidophyta (KEDo) PLAYFORD,1976. Slide 9701042: P50, well 1-MS-3-AM, core 24 at 1184/1189 m, Amazon Basin, Brazil. 5. Geminospora punctata OWENS,1971. Slide 2270(1): G40/3, well MG-1, cuttings sample at 2270 m, H a m m a d a h Basin, Tunisia. 6. Geminospora lemurata BALMEemend. PLAYFORD,1983. Slide C6(1): W54/2, well MG-1, core at 2161.8 m, H a m m a d a h Basin, Tunisia. 7. Geminospora piliformis LOBOZIAK,STREEL & BURJACK,1988. Slide 2178(1): R26, well MG-1, cuttings sample at 2178 m, H a m m a d a h basin, Tunisia. 8, 9. Emphanisporites annulatus MCGREGOR, 1961. 8. Slide 2483(1): Q35, well MG-1, cuttings sample at 2483 m, H a m m a d a h Basin, Tunisia. 9. Slide C21(2): E38/1, well MG-1, core at 2450.6 m, H a m m a d a h Basin, Tunisia. 10. Teichertospora cf. torquata (Higgs) McGREGOR & PLAYFORD, 1990. Slide 950301: U44/3, shallow well Caima PH-2, sample 22(3) at 31.12/31.18 m, Amazon Basin, Brazil. 11. Rugospora radiata (JusHKO) BYVSHEVA, 1985. Slide A.B.99.25(2): 026/3, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 12. Rugospora bricei LOSOZlAK& STREEL,1989. Slide A.B.99.25(2): H34/3, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 13. VaUatisporites vallatus HACQUEBARD,1957. Slide 8502263: K49, well 1-AM-I-AM, core 9 at 1236.66/1239.56 m, Amazon Basin, Brazil. 14. Vallatisporites hystricosus (WINSLOW)BYVSHEVA,1985. Slide A.B.99.25(1): U31, well 2-NA-1-PA, core 25 at 1438.5/1440.5 m, Amazon Basin, Brazil. 15. Vallatisporites v e r r u c o s u s HACQUEBARD,1957. Slide P.B.99.38-40(1): H38, well 2-PM-1-MA, core 38-40 at 1195.6/1198.6 - 1253.4/1256.4 m, P a r n a ~ a Basin, Brazil. 16. Camarozonotriletes sextantii MCGREGOR & CAMFIELD,1976. Slide
405 The successive first appearances of only three diagnostic species have been identified within the late Frasnian-late F a m e n n i a n interval. These are: Teichertospora torquata (an eponym of the torquata-gracilis Zone in the Old Red Sandstone Continent zonation but not yet recorded in ArdenneRhenish regions), followed by Rugospora radiata, and at a slightly higher level, Vallatisporites hystricosus. This latter species first appears before Retispora lepidophyta and characterizes the recently defined VH Zone (Maziane et al. 1999). The youngest Devonian characteristic miospore event is the first occurrence ofRetispora lepidophyta at the base of the latest Famennian or 'Strunian'. R. lepidophyta is a well-known cosmopolitan species which permits accurate correlations all over the world. Its total range within the 'Strunian' is subdivided into three biozones, according to the recently revised miospore zonal scheme of the Ardenne region (Maziane et al,.1999). Each of these subdivisions is characterized by the first occurrence of a diagnostic species, viz.: Knoxisporites literatus (LL), Indotriradites explanatus (LE) and Verrucosisporites nitidus (LN). The 'Strunian' interval has been palynologically investigated in various areas of North Africa and the Middle East giving rise to different regional biozonations based on miospore assemblages. Several samples from well A1-37 in Cyrenaica (northeast Libya) have been assigned to the R. lepidophyta - K. literatus (LL) Interval Zone by Streel et al. (1988, p. 113, fig. 2). The biozonations defined in North Africa by Lanzoni & Magloire (1969), Attar et al. (1980) and Massa et al. (1980) were compared to biozones as currently envisaged by the present authors in northern Brazilian basins (Loboziak et al. in press, p. XXX, fig. 3). In the A1 Kufrah Basin (Grignani et al. 1991, p. 1175, fig. 4), the youngest miospore subdivision of the Binem Formation (palynozone 11), which overlies the supposed early to late Famennian palynozones 9 and 10, contains Verrucosisporites nitidus together with R. lepidophyta, K. literatus, I. explanatus and some other typical latest Famennian associates. A LN Biozono age is therefore ensured for this assemblage. This same assignment, or at least a LE-LN zonal range, also applies to the lowest assemblage of Khosh-Yeilagh in the GhosnaviTilabad cut of Eastern Elburz, northeast Iran (Coquel et al. 1977, p. 62, 63). In northern Brazilian basins, most of the latest Famennian miospore assemblages contain I. explanatus and a few other characteristic species, such as Vallatisporites verrucosus, VaUatisporites vallatus and Tumulispora malevkensis, which first occur only in the upper subdivisions of the R. lepidophy-
ta range. Therefore, the LL Zone, could not be identified in any of the investigated samples and may be entirely absent in northern Brazil. This indicates a biostratigraphic, and probably lithological, gap between at least the VH Zone (or still, older parts of the VCo Zone) and the base of the section containing R. lepidophyta. Due to the scarcity, or even local absence, of V. nitidus in most of the investigated sections, the R. lepidophyta range cannot be accurately subdivided. Therefore, a comprehensive LE-LN zonal attribution is preferably proposed for this interval. In the Paran~ Basin, LE-LN miospore assemblages are sometimes found reworked into Late Carboniferous glaciogenic strata bearing saccate pollen grains. Loboziak et al. (1995) reported a R. lepidophyta microflora from diamictites apparently lacking any pollen grains (well 2-O-1-PR). This may represent the first in situ occurrence of 'Strunian' miospores in the Paran~ Basin (Fig. 2).
CONCLUSION The existence, in various areas of Western Gondwana, of miospore biozones first defined in the Devonian of Western Europe, has been demonstrated in several recent publications. It is concluded that endemic Western Gondwanan species, in the light of criteria defined in the Ardenne-Rhenish regions, also represent good biostratigraphic markers for dating and correlating late Early - Late Devonian strata in those areas, in addition to the zonal species of the Southern Euramerican miospore zonations. A c k n o w l e d g e m e n t s - J.H.G. Melo publishes with the permission of PETROBRAS - PetrSleo Brasileiro S.A.
REFERENCES ATTAR A., CANDILIER A.M., COQUEL R. & FOURNIER J. 1980 - Etude palynologique du D~vonien terminal et du Carbonif'ere inf6rieur du bassin d'Illizi (Fort-Polignac), Alg6rie. Bulletin des
Centres de Recherches et d'Exploration - Production d'Elf Aquitaine, 17: 79-147. BOUMENDJEL K., LOBOZIAK S., PARIS F., STEEMANS P. & STREEL M. 1988 - Biostratigraphie des miospores et des chitinozoaires du Silurien sup6rieur et du D~vonien dans le Bassin d'Illizi (S.E. du Sahara alg~rien). Geobios, 32: 329-357. BUR JACK M.I.A., LOBOZIAK S. & STREEL M. 1987 - Quelques donn~es nouvelles sur les miospores d~voniennes du Bassin du Paran~. Sciences G~ologiques Bulletin, 40: 381-391. BYVSHEVAT.V. 1985 - Spores from Tournaisian and Vis~an stages of the Russian Plate. In MENNERV.V. & BYVSHEVAT.V. (eds), Atlas of spores and pollen of Phanerozoic oil - and gasbearing
FIGURE 5 (suite) - 2456(1): X28/2, well MG-1, cuttings sample at 2456 m, Hammadah Basin, Tunisia. 17. Cymbosporites catillus ALLEN, 1965. Slide 1995(1): $39/3, well MG-1, cuttings sample at 1995 m, Hammadah Basin, Tunisia. 18. Samarisporites triangulatus ALLEN, 1965. Slide 2241(1): L32, well MG-1, cuttings sample at 2241 m, Hammadah Basin, Tunisia. 19. Samarisporites sp. E STREEL& LOBOZIAK, 1987. Slide 950672: N46/3, shallow well Caima PH-2, sample 346 at 129.82/129.90 m, Amazon Basin, Brazil. Slides from wells A1-69, MG1, 2-NA-I-PA, 2-PM-I-MA and RSP-1 are housed in the palynological collection of the Laboratoire de Pal6obotanique, Universit~ de Lille, France. Slides t~om wells l-AM-I-AM, I-AM-7-AM, I-MS-3-AM and Caima PH-2 are housed in the palynological slide collection of the Biostratigraphy and Paleoecology Section of Petrobras Research Centre (Cenpes/Divex/Sebipe), Rio de Janeiro, Brazil. Miospore locations on slides are based on England Finder graticules. Magnification of illustrated specimens: x 500.
406 strata of the Russian and Turanian plates. Vsesoyuznyy Nauchno¢-Issledovatel'skuy Geologicheskiy Institut (VNIGNI), Trudy, 253:80-158 [in Russian]. R., LOBOZIAKS., STAMPFLIG. & STAMPFLI-VuILLEB. 1977 Palynologie du D~vonien sup~rieur et du Carbonif'ere inf6rieur dans l'Elburz oriental (Iran Nord-Est). Revue de Micropaldontologie, 20: 59-71. DAEMON R.F. 1974 - Palinomorfos-guias do Devoniano Superior e Carbonifero Inferior das bacias do Amazonas e Parnaiba. Anais da Academia Brasileira de Ci~ncias, 46: 549-587. ..... 1976 - Correlaq~o bioestratigr~flca entre os sedimentos do Siluriano, Devoniano e Carbonffero Inferior das bacias do Amazonas, P a r n a ~ a e Paran~. Anais do 29 ° Congresso Brasileiro de Geologia, Ouro Preto, 2: 189-194. . . . . . & CONTREIRAS C.J.A. 1971 - Zoneamento palinolSgico da Bacia do Amazonas. Anais do 25 ° Congresso Brasileiro de Geologia, S~o Pau]o, 3: 79-88. . . . . . , QUADROS L . P . & SILVA L . C . 1967 - Devonian palynology and biostratigraphy of the Paran~i Basin. In BIGARELLAZ.J. (ed.), Problems in Brazilian Devonian Geology. Boletim Paranaense de Geoci~ncias, 21/22: 99-132. GRIGNANI D., LANZONI E . & ELATRASH H . 1 9 9 1 - P a l a e o z o i c a n d Mesozoic Subsurface Palynostratigraphy in the A1 Kufrah Basin, Libya. In SALEM M . J . , HAMMUDA O . S . & ELIAGOUBI B . A . (eds), The Geology of Libya (Third Symposium on the Geology of Libya, Tripoli, 27-30 sept., 1987), 4: 1159-1227. Elsevier. Amsterdam. JARDINI~ S. & YAPAUDJIANL. 1968 - Lithostratigraphie et palynologie du D~vonien-Gothlandien gr~seux du Bassin de Polignac (Sahara). Revue de l'Institut fran~ais du pgtrole et annales des combustibles liquides, 23: 439-467. LANZONIE. & MAGLOIREL. 1969 - Associations palynologiques et leurs applications stratigraphiques dans le D~vonien supSrieur et Carbonif'ere inf6rieur du Grand Erg Occidental (Sahara alg~rien). Revue de l'Institut fran~ais du pgtrole et annales des combustibles liquides, 24: 441-469. LOBOZIAKS. in press - Middle to early Late Devonian miospore biostratigraphy of Saudi Arabia. GeoArabia. . . . . . , MELO J.H.G., MATSUDA N.S. & QUADROS L.P. 1997a Miospore biostratigraphy of the type Barreirinha Formation (Curu~ Group, Upper Devonian) in the Tapaj6s River area, Amazon Basin, North Brazil. Bulletin des Centres de Recherches et d'Exploration - Production d'Elf Aquitaine, 21: 187-205. . . . . . , MELO J.H.G., QUADROS L . P . & STREEL M . 1997b - Palynological evaluation of the Famennian Protosalvinia (Foerstia) Zone in the Amazon Basin, northern Brazil: a preliminary study. Review of Palaeobotany and Palynology, 96: 3145. . . . . . , MELO J.H.G., RODRIGUES R., STREEL M . , QUADROS L.P. & BARRILARI I.M.R. 1996 - Age and correlation of the Barreirinha Formation (Curu~i Group, Amazon Basin): new evidence from the miospore biostratigraphy. Anais da Academia Brasileira de Ci~ncias, 68: 207-212. . . . . . , MELO J.H.G., STEEMANSP. & BARRILARII.M.R. 1995 - Miospore evidence for pre-Emsian and latest Famennian sedimentation in the Devonian of the Paran~ Basin, South Brazil. Anais da Academia Brasileira de Ci~ncias, 67:391392. . . . . . , MELO J.H.G. & STREEL M. in press - Latest Devonian and Early Carboniferous palynostratigraphy of northern Brazil and North Africa: a proposed integration of western European and Gondwanan miospore biozonations. Bulletin des COQUEL
Centres de Recherches d'Exploration - Production d'Elf Aquitaine, 22. D. 1992a - Biostratigraphie par miospores du D~vonien inf~rieur ~ sup~rieur du sondage MG-1 (Bassin d'Hammadah, Tunisie) - Comparaison avec les donn~es des faunes. Review of Palaeobotany and Palynology, 74: 193-205. . . . . . ~,~STREELM. 1989 - Middle-Upper Devonian miospores from the Ghadamis Basin (Tunisia-Libya): systematics and stratigraphy. Review of Palaeobotany and Palynology, 58: 173196. . . . . . & STREEL M. 1995a - West Gondwanian aspects of the Middle and Upper Devonian miospore zonation in North Africa and Brazil. Review of Palaeoboany and Palynology, 86: 147-155.
& STREELM. 1995b - Late Lower and Middle Devonian miospores from Saudi Arabia. Review of Palaeobotany and Palynology, 89: 105-113. . . . . . , STREELM. & BURJACKM.I.A. 1988 - Miospores du D~vonien moyen et sup~rieur du bassin du Paran~, Br~sil: syst~matique et stratigraphie. Sciences ggologiques, 41: 351-377. ..... , STREELM. • BURJACKM.I.A. 1989 - D~ductions pal~oclimatiques d'une comparaison entre les assemblages de miospores du D~vonien moyen et sup~rieur de Libye et du Br~sil. Geobios, 22: 247-351. ..... , STREELM., CAPUTOM. V. & MELO J.H.G. 1991 - Evidence of West European defined miospore zones in the uppermost Devonian and Lower Carboniferous of the Amazonas Basin (Brazil). Geobios, 24: 5-11. ..... , STREEL M., CAPUTO M.V. & MELO J.H.G. 1992b - Middle Devonian to Lower Carboniferous miospore stratigraphy in the Central P a r n a ~ a Basin (Brazil). Annales de la Socidtg Ggologique de Belgique, 115: 215-226. ..... , STREEL M., CAPUTO M.V. & MELO J.H.G. 1993 - Middle Devonian to Lower Carboniferous miospores from selected boreholes in Amazonas and P a r n a ~ a basins (Brazil): additional data, synthesis, and correlation. Documents des Laboratoires de Ggologie de la Facultg de Lyon, 125: 277-289. MASSAD. & MOREAU-BENOITA. 1976 - Essai de synth~se stratigraphique et palynologique du Syst~me D6vonien en Libye occidentale. Revue de l'Institut fran~ais du pgtrole, 31: 287333. ..... , COQUELR., LOBOZIAKS. & TAUGOURDEAU-LANTZJ. 1980 - Essai de synth~se stratigraphique et palynologique du Carbonif'ere en Libye occidentale. Annales de la Socigtd ggologique du Nord, 99: 429-442. MAZIANE N., HIGGS K. & STREEL M. 1999 - Revision of the late Famennian miospore zonation scheme in eastern Belgium. Journal of Micropaleontology, 18: 17-25. RICHARDSONJ.B. & McGREGORD.C. 1986 - Silurian and Devonian spore zones of the Old Red Sandstone Continent and adjacent regions. Geological Survey of Canada, 364: 1-79. RODRIGUES R . , LOBOZIAK S., MELO J.H.G. & ALVES D.B. 1995 Geochemical characterization and miospore biochronostratigraphy of the Frasnian anoxic event in the P a r n a ~ a Basin, Northeast Brazil. Bulletin des Centres de Recherches d'Exploration - Production d'Elf Aquitaine, 19: 319-327. STREEL M., FAIRON-DEMARETM. & LOBOZIAKS. 1990 - GivetianF r a s n i a n phytogeography of Euramerica and western Gondwana based on miospore distribution. In McKERROW W.S. ~,5 SCOTESE C.R. (eds), Palaeozoic Palaeogeography and Biogeography. The Geological Society Memoir, 12: 291-296. ..... , HIGGS K., LOEOZIhKS., RIEGELW. & STEEMANSP. 1987 - Spore stratigraphy and correlation with faunas and floras in the type marine Devonian of the Ardenne-Rhenish region. Review of Palaeobotany and Palynology, 50: 211-229. ..... , PARISF., RIEGELW. & VANGESTAINEM. 1988 - Acritarch, chitinozoan and spore stratigraphy from the Middle and Upper Devonian subsurface of Northeast Libya. In EL-ARNAUTIA . , OWENSS. & THUSUB. (eds), Palynostratigraphy of Northeast Libya, Benghazi-Libya. Garyounis University Publications, Benghazi: 111-128. YOLKIN E.A., KIM A.I., WEDDIGE K., TALENT J.A. & HOUSE M.R. 1997 - Definition of the Pragian/Emsian Stage boundary. Episodes, 20: 235-240. .....
S. L O B O Z I A K Universit~ des Sciences et Technologies de Lille UPRESA 8014 du CNRS, Sciences de la Terre F-59655 Villeneuve d'Ascq cedex
. . . . . , STEEMANS P., STREEL M . & VACHARD
JoH.G. M E L O Petrobras/CenpesfDivex/Sebipe Cid. Univ., Q. 7, I. Fund~o 21949-900 Rio de Janeiro, RJ, Brazil
APPENDIX
A. SPECIES
LISTED
Camarozonotriletes ? concavus LOBOZIAK& S T R E E L , 1989 Camarozonotriletes sextantii MCGREGOR8: CAMFIELD, 1976 Chelinospora ligurata ALLEN, 1965
407 Craspedispora ghadamisensis LOBOZIAK& STREEL,1989 Craspedispora paranaensis LOBOZIAK,STREEL & BURJACK, 1988 Crassispora catenata HIGGS,1975 Cymbosporites catillus ALLEN, 1965 Emphanisporites annulatus McGREGOR,1961 Geminospora lemurata BALMEemend. PLAYFORO,1983 Geminospora punctata OWENS,1971 Geminospora piliformis LOBOZIAK,STREEL& BURJAEK,1988 Grandispora douglastownense McGREGOR, 1973 Grandispora gabesensis LOBOZL~K& STREEL,1989 Grandispora incognita (KEDo) McGREGOR & CAMFIELD,1976 Grandispora megaformis (RICHARDSON)McGREGOR,1973 Grandispora permulta LOBOZIAK,STREEL & MELO,1999 Grandispora protea (NAUMOVA)MOREAU-BENoIT,1980 Indotriradites explanatus (LuBER)PLAYFORD,1991
Knoxisporites dedaleus (NAUMOVA)STREEL,1977 Knoxisporites literatus (WALTZ)I%AYFORD,1963 Retispora lepidophyta (KEDo) PLAYFORD,1976 Rugospora bricei LOBOZlAK& STREEL,1989 Rugospora flexuosa (JUSHKO)STREEL,1974 Rugospora radiata (JUSHKO)BYVSHEVA,1985 Samarisporites triangulatus ALLEN,1965 Samarisporites sp. E STREEL & LOBOZIAK,1987 Teichertospora torquata (Higgs) McGREGOR • PLAYFORD,1990 Tumulispora malevkensis (KEDo) TURNAU,1978 Vallatisporites hystricosus (WINSLOW)BYVSHEVA,1985 Vallatisporites vallatus HACQUEBARD,1957 VaUatisporites v e r r u c o s u s HACQUEBARD,1957 Verrucosisporites bulliferus RICHARDSON& McGREGOR,1986 Verrucosisporites nitidus PLAYFORD,1964