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Late Carbonifeous to Permian core samples from two borehole sections, STRAT 1 and CKP 6, that penetrate the glacialto postglacial and coal-bearing Lower Karoo sequence, Kalahari Karoo Basin, Botswana, contain well preserved pollenand spore palynomorphs. The palynomorph assemblages comprise 165 species of spores and pollen, and few acritarchsand chlorophycean algae. This paper presents a local biozonation of three succeeding assemblage zones, recognisable ineach borehole section. The assemblage zones are named: the Hamiapollenites bullaeformis Biozone, theCyclogranisporites gondwanensis Biozone and the Platysaccus papilionis–Striatopodocarpites fusus Biozone, in as-cending order of stratigraphy. Assemblages are compared and correlated with assemblages described from other Gond-wana areas of Africa, Australia, Arabia, South America and Antarctica. A general analysis of taxa from the KalahariKaroo Basin indicates a distinct similarity with assemblages from the Paraná Basin of South America. TheHamiapollenites bullaeformis Biozone is comparable with the Vittatina costabilis Interval Zone of the Paraná Basin. As-semblages from the Cyclogranisporites gondwanensis and the Platysaccus papilionis–Striatopodocarpites fusus zonesare comparable with the Lueckisporites virkkiae Interval Zone. Age determination for the assemblages in the KalahariKaroo Basin is inferred from comparison with similar assemblages from previous Gondwana studies, and ranges fromthe Late Carboniferous (Kasimovian–Gzhelian) to latest Early or possibly earliest Middle Permian (Late Cisularian toEarly Guadalupian). • Key words: Kalahari Karoo Basin, Late Carboniferous–Permian, pollen, spores, assemblagezones.

MODIE, B.N. & LE HÉRISSÉ, A. 2009. Late Palaeozoic palynomorph assemblages from the Karoo Supergroup and theirpotential for biostratigraphic correlation, Kalahari Karoo Basin, Botswana. Bulletin of Geosciences 84(2), 337–358(11 figures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received January 16, 2009; accepted in re-vised form May 30, 2009; published online June 19, 2009; issued June 30, 2009.

Benson N. Modie (corresponding author), The Centre for Knowledge, P.O. Box 1382, Lobatse, Botswana;tesbnm@gmail.com • Alain Le Hérissé, Université de Brest, CNRS, UMR 6538 Domaines Océaniques, InstitutEuropéen de la Mer, CS 93837, 6 Avenue V. Le Gorgeu, 29238 Brest Cedex 3, France

Although the coal-bearing Karoo Supergroup sequence ofthe Kalahari Karoo Basin (Fig. 1) is known to be exten-sive in Botswana (e.g. Smith 1984), geological insightinto stratigraphic correlation remains minimal. This isgreatly compounded by the lack of surface exposure foroutcrop analysis and correlation due to extensive cover bya thin veneer of Cainozoic superficial deposits of the Ka-lahari Desert environment (Thomas & Shaw 1991, Car-ney et al. 1994). The heavy reliance on the traditionalmethod of stratigraphic subdivision, based only on litho-logical similarities, to correlate widely spaced boreholesections in the Kalahari Karoo Basin is not unequivocal.For example, studies of sedimentary environments and fa-cies distributions have shown that similar units can be de-posited at different times, in different areas and with no

physical connection to each other (Reading 1986, Mial1990). The lack of extensive surface exposure in the Kala-hari Karoo Basin and few macrofossils, leaves palyno-logy as the most effective method of dating and correla-ting the subsurface sequences. Evidence from previouspalynological studies (e.g., MacRae 1978, Dolby 1990,ECL 1990, Cadman 1995, Key et al. 1995, Stephenson& McClean 1999) indicates that the Lower Karoo sequ-ence contains pollen and spore microfossils suitable forstratigraphic applications. The results of a recent study(i.e. Modie 2007), on which this paper is largely based,have also revealed an overwhelming presence of pollenand spore palynomorph assemblages suitable for biostra-tigraphic analysis. The purpose of this paper, therefore, isto give an account of the nature of palynomorph assem-

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blages obtained from the Lower Karoo sequence as wellas discuss and highlight their significance to stratigraphiczonation.

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The Kalahari Karoo Basin of Botswana is one among se-veral contemporaneous Karoo basins (Smith et al. 1993,Johnson et al. 1996, ECL 1998) of southwestern Gon-dwana (e.g., Fig. 2) that became active in thePermo-Carboniferous times, with the earliest depositslaid down during the Late Palaeozoic glacial event. Theexact geodynamic setting for the Kalahari Karoo Basin,as well as for the other related Karoo basins, remains littleunderstood. However, several previous workers, amongstthem Daly et al. (1991), Cox (1992), ECL (1998) and Tur-ner (1999), consider the origin of these basins to be linkedto collision induced extensional tectonics related to theLate Palaeozoic to Early Mesozoic Gondwanide Capeorogeny (e.g., Thomas et al. 1992). The latter resultedfrom the subduction of a palaeo-Pacific plate under south-ern Gondwana, which subsequently led to the develop-ment of a retro-arc foreland basin, represented by theMain Karoo Basin in South Africa. The Kalahari KarooBasin and other subsidiary basins in the north, developedas intracratonic rift basins, in response to the orogenicpush from the south (e.g., Daly et al. 1991, ECL 1998).

The sub-surface structural framework of the KalahariKaroo Basin is little understood due to widespread coverby sediments of the Kalahari Desert environment. Conse-quently, much of the covered centrally located KalahariKaroo Basin has hitherto been considered to represent anintracratonic sag basin. Only along the eastern fringe of thebasin, where there is limited exposure and near-surfacebedrock, has it been possible to demonstrate the occurrenceof fault-bounded graben structures (Green et al. 1980). Inrecent times though, the use of geophysical data (e.g., seis-mic, aeromagnetics etc.) to delineate major structural ele-ments, such as faults and lineaments, indicates a rift basinsetting for parts of the covered interior of the KalahariKaroo Basin (Davison & Steenkamp 1995). Generally,both pre- and syn-Karoo fault systems are considered tohave had a local influence on the distribution and thicknessof the Karoo sequence (ECL 1998).

Stratigraphic sub-division (Fig. 3) of the sequence inthe Kalahari Karoo Basin is largely based on lithologicalcomparisons. On a regional scale, the lithostratigraphy issub-divided into lower and upper sequences based on thelocation of a major regional mid-Karoo unconformity(ECL 1998). Sequences that belong to the lower Karoo in-clude the Dwyka, Ecca and Beaufort groups, whilst thosethat belong to the upper Karoo include the Lebung andStormberg Lava groups. Overall, the entire sequence in theKalahari Karoo Basin, as in most other Karoo basins in theregion, records a transition from a glacial period throughfluvio-deltaic and swampy settings and ultimately turningarid before sedimentation came to a halt due to the extru-sion of continental flood basalts (Smith 1984, Smith et al.

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������� ! Locality map indicating the position of study boreholes: insetshows the location of Botswana in the region.

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�������"! The regional location of the Karoo basins of southern Africa(after Johnson et al. 1996).

1993, Johnson et al. 1996). The Karoo basaltic volcanism,considered to have been initiated by extensional tectonicsthat marked the beginning of the Gondwana break-up, rep-resents a major regional magmatic event in Karoo geologythat occurred between 185–177 Ma ago (Jourdan et al.2005).

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This paper is based on samples sourced from drill-coresprovided by the Botswana Department of Geological Sur-vey, as part of a broader project to investigate Karoo basinstratigraphy (Modie 2007). Two well-sampled cored-bore-holes, STRAT 1 and CKP 6, provided 120 and 95 samples,respectively, for palynological investigations. STRAT 1and CKP 6 boreholes are located in separate sub-basins ofthe Kalahari Karoo Basin i.e. Southwest Botswana andWestern Central Kalahari sub-basins, respectively (Fig. 1).The stratigraphic range of sampling from STRAT 1 bore-hole covered the Dwyka and Ecca groups. Sampling fromCKP 6 borehole covered the entire Karoo Supergroup clas-tic sequence i.e. from the Dwyka Group to the LebungGroup. All samples were prepared at the Laboratoire de Pa-laeontologie, Université de Brest, France, where all sampleresidues, as well as all specimens illustrated in this paper,are presently kept. Samples were processed using standardpalynological methods (e.g. Wood et al. 1996). Palynolo-gical slides prepared for each sample were examined usinga transmitted-light microscope (i.e. Leica DIAPLAN) toperform routine slides analyses for the systematic descrip-tions of palynomorphs.

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A total of one hundred and ninety-one productive samples,from both STRAT 1 and CKP 6 boreholes, yielded 165spore and pollen taxa, and 19 acritarchs and chlorophyceanalgae, which are listed in Appendix 1. The range of the taxaand proposed biozonation are indicated in Figs 4 and 5. Allproductive samples are from the Lower Karoo sequencei.e. Dwyka, Ecca and Beaufort groups, with the latter, ho-wever, only yielding very poorly preserved palynomorphsin the basal sections. Samples from the upper Karoo se-quence i.e. Lebung Group, which is largely dominated byred beds, are barren. The palynomorphs recovered from thelower Karoo are dominated by pollen and spores (e.g.,Figs 4, 5), however a few samples also indicate the rare oc-currences of microphytoplanktons (acritarchs and prasnip-hycean phycomata). There is a visible disparity in the pre-servation status of palynomorphs from STRAT 1 and CKP6, which perhaps reflects the palaeo-depositional settingsof separate sub-basins where the two boreholes are located.

The palynomorph content indicates a strong lithofaciesinfluence, with the sandstone facies producing the poorestyields in comparison with the siltstone and mudstone fa-cies. This observation is considered to reflect preferentialsorting with respect to grain-size.

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A detailed assessment of the palynomorph content i.e. qua-litative analysis, revealed the occurrence of taxa suitablefor subdividing the Lower Karoo sequence into biostrati-graphic units or biozones. Although the range charts couldnot produce a perfect correlation between STRAT 1 andCKP 6, consideration of the common taxa in terms of theassemblage zones, allowed the strata in both sections to besubdivided into three biozones which are assemblage zo-nes (Figs 4, 5).

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�������$! Karoo Supergroup lithostratigraphic subdivision and palaeo-environments.

Comparison of the data between STRAT 1 and CKP 6indicates that the base of the sections is characterised by afew common taxa of typically long range nature. There issome disparity in occurrence data between the two sec-tions, which makes the level of comparison in the basalsections rather poorly constrained. The most significanttaxon that reflects some degree of correlation between thebasal sections is the rare and relatively short to mediumrange Punctatisporites ubischii species. A perhaps signifi-cant point to consider with regard to the basal sections isthe fact that these marks the junction between older taxa ofpre-Karoo strata and those originating in early Karootimes. However, in the case of this study, the youngestpre-Karoo sequence consists of barren earliest Cambrianred beds, which unfortunately are not useful for a strati-graphic review of taxa ranges.

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The lower limit of the zone is placed at the lowermost oc-currences of taxa Apiculatisporis parmatus and Potonie-isporites brasiliensis. The upper limit of the zone is pla-ced at the last occurrences of taxa Punctatisporitesgracilis, Densosporites rotundus, and Cristatisporitesmicrovacuolatus. The zone has been named after the spe-cies Hamiapollenites bullaeformis, which forms one ofthe diagnostic taxa, and is restricted to the zone in bothSTRAT 1 and CKP 6, and occurs in other regionally asso-ciated areas.

The diagnostic taxa characteristic of this zone arerepresented by spores, with the most common being:Converrucosisporites, Cristatisporites, Horriditriletesand Punctatisporites spp. The most significant taxa are:Apiculatisporis unicus, Cirrabaculisporites spp., Converru-cosisporites pseudoreticulatus, Converrucosisporites irreg-ularis, Cristatisporites inconstans, Cristatisporites menen-dezii, Divaricrassus minor, Horriditriletes curvibaculosus,Lophotriletes rarus, Lundbladispora gracila, Punc-tatisporites gretensis, Punctatisporites parvus, Raistrickiacrenata and Retusotriletes diversiformis. The VittatinaBisaccate Group appears in this biozone. It also contains im-portant pollen taxa such as Caheniasaccites flavatus,Cycadopites cymbatus, Pakhapites fusus, Protohaplo-xypinus rugatus, Potonieisporites congoensis, Weylanditeslucifer. The most notable similarity between STRAT 1 andCKP 6, in terms of species range, is a major decrease in di-versity indicated by the Cristatisporites spp. at the top ofthe zone. Coincidently, the top of the zone in both sectionsalso mark a major lithofacies change from a predominantlyargillaceous facies, and hence reflecting the possibility offacies-control of the Cristatisporites spp. In general, therest of the zone indicates a close affinity between the

ranges of the taxa and the lithofacies distribution, an obser-vation that may indicate the restriction of ranges due toecological variations and the geographical isolation of taxa(Miall 1990). Nevertheless, some of the less common butsignificant and restricted species, such as Hamiapollenitesbullaeformis and Raistrickia crenata appear to have theirterminations not closely associated with the top boundaryof the zone and hence are probably not related to changescontributing to litho-facies variations. Therefore, theranges of Hamiapollenites bullaeformis and Raistrickiacrenata are considered to broadely represent time-equiva-lent events occurring in STRAT 1 and CKP 6. However, inorder to improve the confidence level in the reality of thiszone, these events require to be verified from other sectionsas well as searching for other diagnostic taxa in future stud-ies. It should be noted that Granulatisporites angularis,present in the biozone, with a long range in the two sectionsis considered here as a junior synonym of Granu-latipsorites austroamericanus Archangelsky & Gamerro,1979. The latter is also considered a possible junior syn-onym of Microbaculispora tentula Tiwari, 1975, byPlayford & Dino (2002). Some specimens referred to inthis study as Converrucosisporites irregularis (Anderson)comb. nov., are similar to some Pseudoreticulatisporapseudoreticulata of Backhouse (1991) which exibit a largemorphological variation. We also put in synonymy withConverrucosisporites pseudoreticulatus (Balme & Henel-ly) comb. nov., Converrucosisporites confluens (Archan-gelsky & Gamerro) Playford & Dino, 2002, which is wellrepresented in the biozone. In terms of thelithostratigraphic divisions this zone corresponds to thelowermost Ecca Group (i.e. Bori/Kobe formations) and theDwyka Group. However, the lower boundary of the zoneindicates the possibility of diachronosity, with the upperDwyka section in CKP 6 reflecting time equivalence withthe lower Dwyka in STRAT 1.

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This is a rather poorly defined zone (Figs 4, 5), which ischaracterised largely on long-range and some medium-range taxa. Compared to the Hamiapollenites bullaeformisAssemblage Zone, this zone has only a slight dominance ofspores over pollen in its list of diagnostic taxa. The moststriking characteristic of this zone in both STRAT 1 andCKP 6 is a drop in the consistency of frequency of occur-rence of taxa, a feature related to the change in the lithofa-cies observed at the upper limit of the Hamiapollenites bu-ellaeformis Assemblage Zone. As a result, there is a lack ofcommon restricted species to form diagnostic taxa and pro-vide time equivalent events for correlation. The lower limitof this zone is determined by the upper limit of the prece-

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ding zone i.e. Hamiapollenites bullaeformis AssemblageZone. The upper limit of the zone is placed at the last oc-currence of the taxon Cyclogranisporites gondwanensis.The latter has provided a name for the zone because it isthe only species that defines the upper limit in bothSTRAT 1 and CKP 6 borehole sections. The Biozone

KK 2 is also characterized by, amongst others: Alisporitesaustralis, Alisporites ovatus, Alisporites potoniei, Conca-visporites mortonii, Horriditriletes brevis, Laevigatispori-tes vulgaris, Lophotriletes rarus, Lueckisporites virkkiae,Protohaploxypinus perexiguus and Marsupipollenitestriradiatus.

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�������%! Lithology and range chart of spore and pollen species identified in core samples from STRAT 1 borehole, with inferred palynozonal attributions.

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This zone (Figs 4, 5) represents a narrow uppermost produc-tive section of the sampled strata, dominated by bisaccatepollen. The lower limit of the zone is placed at the first oc-currence of the taxa Platysaccus papilionis and Striatopodo-carpites fusus. The upper limit is not defined due to thenon-availablity of core material for sampling beyond thetopmost sampled level. Consequently, this zone possibly re-presents a lowermost subzone of a broader zone. The zone isnamed after these two species because they are the most sig-nificant taxa restricted within the zone, as noted in boreholeSTRAT 1 and CKP 6, as well as in equivalent zones from ot-her previous local studies. The species Luekisporites vir-kkiae ranges into Biozone KK 3 in STRAT 1, and Laeviga-tisporites vulgaris on the other hand appears in the samerange in CKP 6 whereas it disappears in STRAT 1. Otherdiagnostic short range taxa in the biozone are: Vestigispori-tes ventrisaccatus, common to the two cores and Floriniteseremus, Hamiapollenites saccatus, Polypodiisporites muta-bilis and Striatopodocarpites pantii.

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Most of the limited palynology studies undertaken fromBotswana either present only palynostratigraphic inter-pretations in terms of biostratigraphic units and their agesor assemblage descriptions without occurrence data (e.g.,Dolby 1990, Key et al. 1995). This approach presents prob-lems for comparison and correlation due to the unavailabi-lity of ranges of taxa, and as a result, only broad compari-sons can be considered. A few of the previous studieshowever (e.g., MacRae 1978, ECL 1990, Stephenson &McClean 1999) do present results that give insight into thevertical characteristics of taxa, which allows for a limitedbut significant comparison and correlation (e.g., Fig. 6).

Correlation with the zonation scheme of MacRae (1978). –Although displaying some considerable variations in taxo-nomic terminology, the unpublished work of MacRae(1978) presents the first complete preliminary palynozona-tion scheme available for correlation. The scheme comprisesthree concurrent range zones based on the distribution ofsixty eight miospore taxa, namely zones I, II, and III, in as-cending order of the stratigraphy, and recognised in bore-hole N1/3 situated in the northeast part of the basin. A broadcomparison (Fig. 6) indicates that the section of STRAT 1that corresponds to Biozone KK 1 can be correlated withMacRae’s Biozone 1. Species most significant and commonto both sections include: Vittatina minima, Caheniasaccitesflavatus, Caheniasaccites ovatus, Plicatipollenites densus,Plicatipollenites gondwanensis, Punctatisporites gretensis

and Calamospora aplata. The stratigraphic range of thecommon taxa indicates a correlation of the middle to upperDwyka Group (e.g., Khuis and Middlepits formations) andlower Ecca Group (e.g., Kobe Formation) in STRAT 1 with amuch condensed similar section of the Karoo Supergroupdescribed in MacRae (1978). The upper section of STRAT 1,which includes biozones KK 2, KK 3, and the topmost sectionof Biozone KK 1 can be broadly correlated with the sectioncovered by MacRae’s concurrent range zones II and III inN1/3. Species common to both sections include: Alisporitessplendens, Striatopodocarpites fusus, Protohaploxypinuslimpidus, Alisporites ovatus and Laevigatosporites vulgaris.A comparison of the suprageneric quantitative trends revealsa rise in the relative abundance of bisaccate striate pollen inboth sections. Overall, STRAT 1 allows for only a broad andgeneral comparison. As a result there are no visible detailsthat make it possible to comparatively analyse the lithofaciesfor important factors such as diachronous characteristics.

Correlation with the zonation scheme of ECL (1990). –ECL (1990) erected a regional zonation scheme of five pa-lynozones based on analysis of borehole and field samplesfrom Namibia, Botswana, Zimbabwe, and Zambia. The pa-lynozonation scheme of ECL (1990) is largely based on va-riations determined by the relative abundance of taxa,which makes it difficult to correlate with the present studyin which the zonation is largely based on species’ ranges.Nevertheless, few diagnostic taxa allow for a broad compa-rison and correlation (Fig. 6). The first half of BiozoneKK 1 of STRAT 1 can be correlated with Zone I (MZ I) onthe basis of common taxa that are dominated by monosac-cates and consist of Plicatipollenites gondwanensis and re-latively diverse Potoniesporites spp. Other species of note,albeit in different taxonomic terminology, include Vesti-gisporites rudis and Cristatisporites morungavensis. Thesecond half of Biozone KK 1 is correlated with Zone MZ IIand the base of Zone MZ III on the basis of the common oc-currence of Cirrabaculisporites plumsteadi, an increase innon-striate pollen in the upper part of the zones and finallythe upper limit of the regular monosaccates i.e. Plicatipol-lenites and Cannanoropollis species. The KK 2 Biozone iscorrelated with the upper part of Zone MZ III and ZoneMZ IV, based on the occurrence of common significanttaxa including Lueckisporites virkkiae, Marsupipollenitestriradiatus and Laevigatosporites spp. Other characteristictaxa interesting in terms of correlation include Protohaplo-xypinus spp., Weylandites lucifer and Cycadopites cymba-tus. The uppermost KK 3 Biozone is correlated with ZoneMZ V based on a first appearance of Striatopodocarpitespantii as well as the following common taxa: Lueckispori-tes virkkiae, Weylandites lucifer, Polypodiisporites muta-bilis and Laevigatosporites vulgaris. The base of the zoneis generally reflected by a marked increase in the relativeabundance of the striate bisaccates.

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�������(! Range chart ofspore and pollen speciesidentified in the core sam-ples from CKP 6 borehole,with inferred palynozonalattributes.

Correlation with the work of Stephenson & McLean(1999). – Stephenson & McLean (1999) described an EarlyPermian palynological assemblage from a Lower Karoo se-quence, represented by the Morupule Formation, in the eas-tern fringe of the Kalahari Karoo Basin. There is remarkablesimilarity between STRAT 1 and the Morupule section interms of taxa content, albeit with some variations in taxono-mic terminology. However, the majority of the commonspecies depict a long range characteristic in the relatively ex-tensively sampled STRAT 1 section. The Morupule Forma-tion represents a narrow zone in terms of Karoo strata andless than 15 m was sampled by Stephenson & McLean(1999). Consequently, the initiations and terminations ofmost taxa most likely fall outside the sampled section andhence species ranges remain poorly constrained. Neverthe-less, comparison of the common taxa allows only for a broadand general tentative correlation (Fig. 6). The most compa-ratively significant taxa that correlate STRAT 1 to the Moru-pule section include Laevigatosporites vulgaris, Alisporitesovatus, Alisporites potoniei, Florinites eremus, Protohaplo-xypinus limpidus, Striatoabieites multistriatus and Platysac-cus spp. The ranges of the significant taxa broadly corres-pond to biozones KK 2 and KK 3 of STRAT 1.

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Palynological studies pertinent to Karoo Supergroup stra-tigraphy have been undertaken by various workers in seve-ral of the contemporary Karoo basins of southwestern Gon-

dwana (e.g. Manum & Duc Tien 1973; Jardiné 1974;Falcon 1975–1978; Utting 1976, Anderson 1977; Wright& Askin 1987; MacRae 1988; Hankel 1992–1993; Aitken1994; d’Engelbronner 1996; Nyambe & Utting 1997; Kal-kreuth et al. 1999; Millsteed 1994–1999; Semkiwa et al.2003 etc.). The results of such studies reveal some impor-tant common features that allow for a regional comparisonand correlation of the Karoo sequence in the various ba-sins. A selected number of the regional studies are discus-sed below and compared with the results from STRAT 1section (e.g., Fig. 7).

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Correlation with the zonation scheme of MacRae (1988). –MacRae (1988) described palynomorph assemblages fromthe northern Karoo basins of South Africa and subsequ-ently erected six palynozones i.e. A–F, with which a broadcorrelation with STRAT 1 borehole is possible (Fig. 7).MacRae’s (1988) Biozone A is tentatively correlated withthe basal section of STRAT 1 where although no signifi-cant taxa exist for direct correlation, common taxa of ratherlong ranges occur. The top of Biozone B is equated with theupper part of Biozone KK 1 based on the terminations ofPlicatipollenites trigonalis and Vestigisporites rudis, bothof which occurs in the two zones. On the other hand, thebase of Biozone B favourably equates with the base of Bio-zone KK 1 as defined on the basis of the initiation of thefollowing species: Weylandites lucifer, Verrucosisporitespseudoreticulatus, Horriditriletes tereteangulatus and

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�������)! Correlation of the zonation scheme from STRAT 1 borehole (core section for reference) with zonation schemes from previous local studies.

Striatoabieites multistriatus. Biozone C (MacRae 1988),described as the least palynologically productive, favou-rably equates with the top of Biozone KK 1, characterisedby a remarkable decrease in the frequency of occurrence aswell as the termination of most species, and the base ofBiozone KK 2 of STRAT 1. The top boundary of BiozoneKK 2 equates with the top boundary of Biozone D on thebasis of the range termination of Laevigatosporites vulga-ris in STRAT 1, even though the species is known higherup in CKP 6. Biozone KK 3 is correlated with Biozone E,albeit with difficulty, due to the lack of any common signi-ficant trends for direct correlation. A perhaps more signifi-cant observation is the occurrence of Platysaccus papilio-nis and Striatopodocarpites fusus, both of which arerestricted to this zone in STRAT 1. The assemblage ofcommon taxa among these zones includes: Deltoidosporadirecta, Striatoabieites multistriatus, Weylandites lucifer,Alisporites potoniei, Indotriradites australensis, Apicula-tisporis levis, Protohaploxypinus limpidus, Protohaploxy-pinus hartii and Striatopodocarpites cancellatus.

Lithologicallly, the overall trend deduced from compar-ative analysis of the palynological assemblages reflects adiachronous Karoo sequence between the north Karoo bas-ins of South Africa (e.g., borehole ET 61 in MacRae 1988)and the sequence at STRAT 1 borehole, especially with re-gard to the basal successions. MacRae’s (1988) biozones Aand B which largely fall within the lower Ecca Group suc-cession (i.e. Wellington Formation) but also include a muchcondensed section of the Dwyka Group, correlate to a largeextent with the Dwyka Group (i.e. Malogong, Khuis, andMiddlepits formations), as well as a basal section of the EccaGroup (i.e. Kobe Formation) in STRAT 1 borehole. Simi-larly, Biozone C (MacRae 1988), which represent the lowerparts of the middle Ecca succession (i.e. Enkelbult Forma-tion), correlates with the uppermost part of the lower Ecca(i.e. Kobe Formation), but also encroaches into the basalsection of the middle Ecca, in STRAT 1 borehole. The up-permost successions, although relatively poorly constraineddue to low palynomorph yields, indicate a much closerlithological correlation between the respective successions.The latter reflects the possibility of a shift from adiachronous setting to a near time equivalent deposition dur-ing the middle to upper Ecca.

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Correlation with the scheme of Falcon (1975). – Palynostra-tigraphic studies from the Mid-Zambezi Karoo Basin inZimbabwe by Falcon (1975) allow for a broad but limitedcorrelation (Fig. 7), based on rare similarities reflected byqualitative and quantitative analyses. Falcon (1975) erecteda zonation scheme that subdivided the Lower Karoo sequ-ence in the Mid-Zambezi Basin into four major assemblage

zones, including eight sub-zones. Biozone KK 1 inSTRAT 1 borehole can be equated with Falcon’s (1975) As-semblage I. One notable and comparable characteristic arethe ranges of Converrucosisporites pseudoreticulatus andCycadopites cymbatus species, both considered significantfor Assemblage I. However, only Converrucosisporitespseudoreticulatus form a significant taxon in the basal sec-tion of Biozone KK 1 of STRAT 1. The quantitative analy-ses also indicate some rare but significant similarities, whichinclude a relatively marked upward increase in the zonola-minatitriletes, as well as an increase in the bisaccates pollendefining the top boundary. A rare direct comparison is ref-lected by the initiations of Retusotriletes diversiformis andAlisporites ovatus which both correlate the base of Falcon’s(1975) Assemblage Sub-Zone B with the middle part of Bio-zone KK 1 of STRAT 1. The biozones KK 2 and KK 3 canbe equated with Falcon’s (1975) Assemblage II, albeit withdifficulties. The lower boundary for the correlated zones isplaced at the level where there is a marked upward increasein the relative abundances of bisaccates pollen. Taxa of no-table and comparable ranges include Marsupipollenitesspp., Striatoabieites spp., Striatopodocarpites spp., Laevi-gatosporites spp., Alisporites splendens and Alisporites po-toniei. The introduction of Laevigatosporites spp. appears tobe the most significant in defining the lower boundary forcorrelation.

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Correlation with the zonation scheme of Utting (1976). –Two palynological assemblages described by Utting (1976)from the Lower Karoo Luwumbu Coal Formation of the Lu-angwa Valley in Zambia broadly compare (Fig. 7) with theassemblages identified in STRAT 1. Based on comparisonof limited taxon ranges, the section of borehole STRAT 1corresponding to Biozone KK 1 can be equated with the ol-der assemblages obtained from the Mukumba SiltstoneMember. The few comparable and characteristic taxa, occa-sionally with variable taxonomic nomenclature, include Pli-catipollenites densus, Plicatipollenites trigonalis, Conver-rucosisporites pseudoreticulatus and Cristatisporites lestai.The latter assemblage broadly correlates the Dwyka Groupand the Kobe Formation (i.e. lower Ecca Group) of the Kala-hari Karoo Basin with the Mukumba Siltstone Member ofthe Luwumbu Coal Formation in Zambia. The upper sectionof STRAT 1 corresponding to biozones KK 2 and KK 3,compares favourably with the younger assemblage determi-ned from the Mpwashi Carbonaceous Member. The mostsignificant taxa in terms of comparative analysis of the ran-ges include Laevigatosporites spp., Alisporites potoniei,Protohaploxypinus perexiguus, Marsupipollenites striatus,Weylandites lucifer and Striatoabieites spp. This assem-blage allows for a broad correlation of a section of the Otshe

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Formation which corresponds to the middle Ecca, with theMpwashi Carbonaceous Member of the Luwumbu CoalFormation in Zambia.

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Correlation with the zonation scheme of Semkiwa et al.(2003). – Semkiwa et al. (2003) gave a brief account of pa-lynological analysis from the Lower Karoo sequence of theSongwe-Kiwira Coalfield in Tanzania. A comparison ofcommon taxa reflects the possibility of a broad correlationbetween STRAT 1 section and the sequence atSongwe-Kiwira Coalfield (Fig. 7). The basal section ofBiozone KK 1 can be equated with the Cannanoropollis–Plicatipollenties Zone on the basis of the common occur-rence of monosaccates, dominated by Cannanoropollisspp. and Plicatipollenites spp. There is also a notable simi-larity with regard to the common occurrence of Cristatis-porites spp. This comparison generally correlates theDwyka Group with the Idusi Formation at the Kiwira CoalMine. The upper sections of STRAT 1 reveal a much moreevident similarity as reflected by the occurrence of severalspecies of comparable ranges. The most significant taxa in-clude Alisporites ovatus, Alisporites potoniei, Laevigatos-porites spp., Marsupipollenites striatus, Protohaploxypi-

nus limpidus, Protohaploxypinus rugatus, Vittatinascutata and Weylandites lucifer. This assemblage allowsfor a correlation of the section corresponding to the upperhalf of Biozone KK 1 and Biozone KK 2 with the Scheurin-gipollenites–Protohaploxypinus Zone (e.g., Semkiwa et al.2003). This consequently equates a section of the upper-most Dwyka, as well as the Kobe Formation and much ofthe Otshe Formation with the Mchuchuma Formation ofthe Songwe-Kiwira Coalfield in Tanzania.

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Correlation with the zonation scheme of Henkel (1992). –Henkel (1992) documented Late Permian to Early Triassicassemblages from the Karoo Sequence of the MombasaBasin in Kenya. Sampling for this work covered a very nar-row section of the Maji ya Chumvi Formation, which ma-kes correlation very poor and largely inconclusive. Of thetwo assemblages described, only the older assemblage,which unfortunately is limited to one sampling level, canbe loosely equated with biozones KK 2 and KK 3. Taxaconsidered for comparative analysis include: Lueckispori-tes virkkiae, Weylandites spp., Plicatipollenites spp., Pla-tysaccus spp., Striatopodocarpites spp. and Protohaplox-pinus spp.

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�������*! Correlation of STRAT 1 zonation scheme with a select few zonation schemes from previous regional studies.

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Correlation with the scheme of Jardiné (1974). – Palynolo-gical studies by Jardiné (1974) on Karoo equivalent sequen-ces revealed three palynological assemblages from the “Sé-rie de l’Agoula”, which overlies glacial deposits. Thelowermost assemblage P II can be equated (e.g., Fig. 7) withlower part of Biozone KK 1 in STRAT 1 on the basis ofcommon characteristic taxa such as Pakhapites fusus, Pak-hapites ovatus, Caheniasaccites ovatus and Striomonosac-cites spp. The succeeding assemblage P III is loosely corre-lated with upper part of Biozone KK 1 based on thesignificant common occurrence of Protohaploxypinus spp.and Striatoabieites spp. In addition, the latter zone is, in the“Série de l’Agoula”, immediately succeeded by a non-productive interval, which compares well with the intervalcorresponding to the topmost section of Biozone KK 1, andto Biozone KK 2. Lastly, the uppermost assemblage P IV ofthe “Série de l’Agoula” can be correlated with Biozone KK3 based on the common occurrence of Lueckisporites vir-kkiae and Platysaccus papilionis, which are characteristicspecies in both of the sections.

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Palynological assemblages described by Wright & Askin(1987) from Karoo sequences of the Morondava Basin in Ma-dagascar indicate some comparable characteristics with thesequence at STRAT 1 borehole. The assemblage that charac-terizes the Lower Sakamena section of the Madagascar stra-tigraphy compares favourably with the assemblages at the topof Biozone KK 1, and in biozones KK 2 and KK 3 (Fig. 7).Taxa with closely comparable ranges of occurrence includeLueckisporites virkkiae, Striatopodocarpites pantii, Alispori-tes ovatus, Plicatipollenites gondwanensis, Protohaploxypi-nus limpidus, Platysaccus spp. and Weylandites spp.

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Although long-distance (e.g., between continents) paly-nostratigraphic correlation is a common practice amongbiostratigraphers, there is always a great deal of uncerta-inty associated with the frequency of distribution and thepalaeogeography of the respective palynomorphs. Firstly,the frequency distribution of taxa, even on a local scale, isgreatly skewed such that there are major disparities bet-ween different sections under investigation (Agterberg &Gradstein 1999), which ultimately gives rise to imprecisecorrelations. The extrapolation of such inherent problemsto a large-scale or long-distance correlation is likely to re-sult in exaggeration and mis-correlations. A common prob-lem observed in the present study, but also reported by

other workers (e.g., Souza 2006, Stephenson 2008), invol-ves the realization that certain short-range taxa occur aslong-range taxa in other regions, making correlation imprac-tical. Secondly, studies have revealed that global florasprogressively became palaeogeographically restricted, par-ticularly during the latest Carboniferous to Permian, a phe-nomenon that has greatly reduced the efficacy of long-distance palynostratigraphic correlation (Playford & Dino2005). The above observations call for a concerted effort toupgrade and refine palynological data at the local level be-fore attempting to erect long-distance correlation schemes.Therefore, with the above limitations in mind, the resultsfrom the present study will be compared and correlated withonly a few selected Gondwana regions (e.g., Fig. 8).

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Correlation with the zonation scheme of Souza & Marques-Toigo (2003, 2005), and Souza (2006). – Palynological as-semblages described by Souza and Marques-Toigo (2003,2005) and Souza (2006), from the Brazilian Paraná Basin,indicate a remarkable similarity with the assemblages obtai-ned from the Kalahari Karoo Basin in Botswana. Consequ-ently, the palynozones erected for the sequence at STRAT 1borehole can, to a certain degree, be compared and correla-ted (Fig. 8). The lowermost basal section of STRAT 1 (e.g.,below Biozone KK 1) is poorly constrained but can be loose-ly correlated with the Crucisaccites monoletus Interval Zoneon the basis of the common occurrence of monosaccatessuch as Cannanoropollis spp., Plicatipollenites spp., Poto-niesporites spp., and Caheniasaccites spp. In addition, bothazonotriletes spores (i.e. laevigate and apiculate), compri-sing Punctatisporites gretensis, Horriditriletes spp. etc., andzonolaminatitriletes (e.g., Cristatisporites spp. and Vallatis-porites spp.) form significant constituents. The section ofSTRAT 1 borehole confined between the base of BiozoneKK 1 and the lower section of Biozone KK 2 can be equatedwith the Vittatina costabilis Interval Zone on the basis of asignificant incoming of several Vittatina species. In addi-tion, the species after which the zone was named (e.g., Souza& Marques-Toigo 2005, fig. 2f) also form a significant spe-cies in STRAT 1 (described as Vittatina sp. in this study).The top of Biozone KK 1 in STRAT 1 can be directly corre-lated with the top of the Protohaploxypinus goraiensis Sub-zone on the basis of the terminations of Plicatipollenites tri-gonalis and Potoniesporites congoensis. The upper sectionsof STRAT 1 borehole, stretching from the upper part of Bio-zone KK 2 up to and including Biozone KK 3, compares fa-vourably with the Lueckisporites virkkiae Interval Zone.Taxa considered pertinent in recognizing the basal sectionsof the zones of correlation comprise Lueckisporites virkiaeand Laevigatosporites spp. However, there are several othertaxa characteristic of these zones, the most notable being:

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Marsupipollenites striatus, Striatopodocarpites fusus, Stria-topodocarpites pantii and Weylandites lucifer.

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Correlation with the zonation scheme of Backhouse(1991). – Backhouse (1991) established the most widelycompared palynostratigraphic scheme from the GondwanaPermian sequences of the Collie Basin of Australia. Thereare however some difficulties in correlating the zonationscheme established here with that of Backhouse (1991) forthe Collie Basin. The most important limitation is due tothe fact that between the two areas certain key species re-veal variable ranges and abundances, an observation alsorecognized by Backhouse (1991) when comparing the as-semblages from the Collie Basin with other Australian ba-sins. Due to the difficulties outlined above, only a broadcorrelation (Fig. 10), based on general similarities in theranges of common taxa is considered here. The most clo-sely comparable assemblages indicate a probable correla-tion of biozones KK 2 and KK 3 of STRAT 1 borehole withthe Striatopodocarpites fusus Zone of the Collie Basin.The most significant taxa pertinent to this correlation com-prise Florinites eremus, Horriditriletes tereteangulatus,Laevigatosporites vulgaris forma colliensis, Cycadopitescymbatus which disappears in KK 2, and Striatopodocar-pites fusus, all of which are initiated at the same level in theCollie Basin. Other significant species that indicate a com-parable range with Striatopodocarpites fusus in the CollieBasin, and also form characteristic taxa in the correlatablesection of STRAT 1 borehole includes: Cymatiosphaeragondwanensis, Weylandites lucifer, Marsupipollenites tri-radiatus, Marsupipollenites striatus, Alisporites ovatus,Alisporites potoniei, Striatoabieites multistriatus and Pro-tohaploxypinus limpidus. Based on the range of a few se-lected taxa, the lower part of the two cores, STRAT 1 andCKP 6, corresponding to KK 1 Biozone and the interval be-low, compares favourably with the section at Collie Basinthat precedes the Striatopodocarpites fusus Zone, and in-cludes the Pseudoreticulatispora pseudoreticulata Zone,the Pseudoreticulatispora confluens Zone and the top ofStage 2. The assemblage pertinent to this correlation inclu-des: Plicatipollenites spp., Converrucosisporites naumo-viae, Puncatisporites gretensis, Cycadopites cymbatus,Verrucosisporites andersonii and Cristatisporites spp.

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Palynological studies undertaken by various workers in An-tarctica reveal the occurrence of assemblages with commonand comparable taxa to those recognized in this study. As aresult, the sequence at STRAT 1 can be generally compared

and broadly correlated with stratigraphic sections from An-tarctica (Fig. 8). Permian assemblages described by Farabeeet al. (1991), from the Buckley Formation of the centralTransantarctic Mountains, closely compares with assembla-ges from the upper sections of STRAT 1 borehole, whichcomprises the upper part of Biozone KK 1, and biozones KK2 and KK 3. The most characteristic taxa pertinent to thiscorrelation include: Marsupipollenites triradiatus, Marsupi-pollenites striatus, Protohaploxypinus limpidus and Laevi-gatosporits spp. Lindström (1995a, b) also described somepalynomorph assemblages from Antarctica which comparefavourably with those recognized in the Karoo supergroup.The assemblages recognized from the Vestfjella mountain-range (e.g., Lindström 1995a) compare favourably with thoseobtained from the interval comprised of the upper part of Bio-zone KK 1 as well as biozones KK 2 and KK 3, as was thecase with the Buckley Formation above (e.g., Farabee et al.1991). The most significant taxa for comparison with the se-quence at the Vestfjella mountain-range include: Alisporitesovatus, Alisporites potoniei, Cymatiosphaera gondwanensis,Florinites eremus, Laevigatosporites vulgaris, Lueckisporitesspp., Marsupipollenites striatus, Marsupipollenites triradia-tus, Striatopodocarpites fusus, Striatoabieites multistriatus,Weylandites lucifer and Weylandites magmus.

A comparison of the assemblages from STRAT 1 withthose described by Lindström (1995b) from the Heime-frontfjella mountain-range reveals the occurence of twogroups of correlatable taxa. Firstly, an older assemblagecharacterized by Cahenniasaccites ovatus, Cycadopites ne-vesi, Limitisporites rectus and Verrucosisporites andersonii,which broadly correlates the lower section of STRAT 1, i.e.the basal section of Biozone KK 1, with the lower half ofLocality A. Secondly, a younger assemblage that correlatesthe upper part of Biozone KK 1 and biozones KK 2and KK 3, with the upper half of Locality A (Lindström1995b) is comprised of typical taxa as observed in the earliersections above. Such taxa include: Alisporites ovatus, Ali-sporites potoniei, Laevigatosporites vulgaris, Protohaplo-xypinus limpidus, Striatopodocarpites fusus, Striatoabieitesmultistriatus and Weylandites magmus.

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Palynological studies from Oman and Saudi Arabia (e.g.,Besems & Schuurman 1987; Love 1994; Penney et al.2008; Stephenson & Osterloff 2002; Stephenson et al.2003; Stephenson 2004, 2008) present an opportunity tocompare and correlate the sequence in the Kalahari KarooBasin with the Late Palaeozoic Gondwana sequence of theArabian Peninsula. However, there is a lack of any signifi-cant similarities between taxa from either area, and henceonly a limited comparative analysis that allows for only abroad and general correlation is possible.

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Penney et al. (2008) proposed a complete correlationchart of palynostratigraphical schemes for Oman. Thelower sections of STRAT 1 and CKP 6 boreholes, equiva-lent to KK 1 Biozone, can be broadly correlated with sev-eral assemblage zones defined in Oman and Saudi Arabia(Fig. 8): Besems & Schuurman’s (1987) AssemblageGroup A; an equivalent of Love’s (1994) Microbaculi-spora Assemblage; the succession of Penney et al.’s(2008) biozones 2165A and 2165B, and the base ofStephenson et al.’s (2003) OSPZ2 Biozone in Saudi Ara-bia. Those assemblages, as in the Karoo, are dominated bycingulocamerate trilete spores, such as Cristatisporitesspp., and monosaccate pollen that includes Plicati-pollenites spp. and Cannanoropollis spp. In Saudi Arabia,the base of OSPZ2 is defined by first occurrence ofHorriditriletes spp., which are also well represented in thebase of sequences of STRAT 1 and CKP 6.

Besems & Schuurman’s (1987) Assemblage Group B,which is characterized by striate and non-striate bisaccates,and Penney et al.’s (2008) biozones 2141A, 2141B and2141C, can be broadly correlated with the upper part ofBiozone KK 1 and part of Biozone KK 2.

Palynological assemblages from the Rahab and LowerGharif members of the Aklata and Gharif formations, re-spectively, described by Stephenson & Osterloff (2002)and Stephenson et al. (2003), can also be compared in ageneral and broad perspective. Taxa characteristic of the

Microbaculispora tentula and the Converrucosisporitessp. A – Microbaculispora grandegranulata biozones andtop of Biozone OSPZ2, reveal close similarities to thatcharacterising the upper part of Biozone KK 1 and BiozoneKK 2. The most significant taxa pertinent to this compari-son include: Apiculatisporis cornutus, Apiculatisporis lep-toacaina, Apiculatisporis parmatus, Cycadopites cym-batus, Horriditriletes uruguaiensis, Pakhapites fusus andVerrucosisporites andersonii. The upper Biozone KK 3 ofSTRAT 1 and CKP 6 boreholes can be only poorly equatedwith the Alisporites indarraensis Biozone and the base ofStephenson et al.’s (2003) Biozone OSPZ3, based on acommon increase in diversity of bisaccate pollen. Rare sig-nificant taxa pertinent to this correlation include Striato-podocarpites fusus and Protohaploxypinus limpidus.

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Correlation with the Gondwana Sequence of India isoverwhelmingly limited despite several decades of paly-nological studies that generated a plethora of data on mor-photaxonomy and palynostratigraphy (e.g., Bharadwaj1962, Bharadwaj & Salujha 1964, Bharadwaj & Srivas-tava 1969, Tiwari 1964, Kar 1968, Lele 1973, Tiwari &Singh 1981, Tiwari & Tripathi 1992, Singh et al. 1995,Meena 1999, Jana et al. 2002 etc.). Most palynostratig-

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�������+! Correlation of STRAT 1 zonation scheme with main palynozonation for the regions of Gondwana.

raphic studies involve quantitative analyses at the genuslevel and describe assemblages that are confined to andbased on the litho-stratigraphic subdivision, making itdifficult to compare and correlate in terms of species ran-ges. In addition, there is some noticeable degree of varia-tion in taxonomical nomenclature, which also contributesto the difficulties of correlation. Tiwari & Tripathi (1992)compiled a species-determined composite palynozona-tion scheme based on stratigraphic distribution data obtai-ned from previous studies undertaken in the DamodarGraben and several other pertinent basins of India. Thiswork (i.e. Tiwari & Tripathi 1992) allows for a limited butimportant comparative analysis of the assemblages fromIndia and those recognized from the Kalahari Karoo Ba-sin, as is described below.

A review of the ranges of common taxa reveals twogroups of assemblages that broadly correlate the sequenceat STRAT 1 borehole with the sequences in the Damo-dar-Rajmahal basins of India. The first group, comprisingan older assemblage, is characterized by: Caheniasaccitesovatus, Cannanoropollis spp. (as Parasaccites), Plicati-pollenites spp. (i.e. P. gondwanensis, P. densus, andP. trigonalis), and Potonieisporites spp., which closelycorrelates Biozone KK 1 with Tiwari & Tripathi’s (1992)assemblage zones I–III. This comparison broadly equatesthe Dwyka Group and the lowermost Ecca Group inSTRAT 1 with the Talchir Formation of India.

The second group, which forms a younger assemblageand is relatively more diverse, is characterized by the fol-lowing taxa: Alisporites ovatus (as Scheuringipollenitesovatus), Alisporites potoniei (as S. maximus), Laevi-gatosporites spp., Lophotriletes rectus, Marsupipollentiestriradiatus, Marsupipollenites striatus, Protohaploxypinuslimpidus (as Faunipollenites varius), Protohaploxypinusperexiguus (as F. perexiguus), and Weylandites lucifer.The latter assemblage favourably compares biozones KK 2and KK 3, with Tiwari & Tripathis’s (1992) assemblagezones III–VIII. Such a comparison broadly correlatesmuch of the Ecca Group in STRAT 1 section, with theKarharbari, Barakar and Kulti formations, as well as basalsections of the Raniganj Formation.

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The subject of age determination on the basis of biostrati-graphic character requires data of high density and qualitythe nature of which is beyond the scope of this study. Never-theless, some general inferences can be made to give insightinto chronostratigraphic relations. Age determination forbiostratigraphic units depend on reliable correlation withmarine sequences containing age-definitive faunas fromwhich internationally agreed reference sections or stratoty-pes have been developed (Dunn 2001, Playford & Dino2005, Stephenson 2008). Most of the early reference secti-ons used to erect international standard stages, with respectto the relative geological time-scale, developed in Europe,are largely based on the distribution of ammonoids, cono-donts, and foraminifers (Christopher & Goodman 1996).For most Gondwana sequences, where there is a general lackof marine fossils, zonation schemes are mainly based on ter-restrial stratigraphic intervals, and precise dating and corre-lation remains uncertain. This is compounded by the geo-graphic restriction of most Permian assemblages, whichresulted from the existence of distinct palaeoclimatic zoneswith associated floral provinces (ECL 1990, Playford &Dino 2005). In addition, radiometric age determination forUpper Palaeozoic strata of Gondwana, some of them for theSouth African Karoo Basin (e.g. Bangert et al. 1999), re-main limited, and hence age control for the diagnostic as-semblages remains equivocal. Consequently, age control forassemblages, and hence the zonation scheme, from the pre-sent study is inferred from comparison with other Gon-dwana palynofloras, and as a result remains tentative.

Comparison of the palynological assemblages from thepoorly constrained basal sections of the Karoo sequenceconsidered in this study, with other previous works (e.g.,MacRae 1988, Dolby 1990, Souza 2006), indicate LateCarboniferous ages (i.e. Kasimovian–Gzhelian). The latteris seemingly typified, in Gondwana sequences, by the com-mon dominance of monosaccates and trilete spores whencompared with the dominance of bisaccate pollen (e.g.,MacRae 1988, Souza 2006). Late Carboniferous ages havealso been suggested from equivalent lithofacies (e.g.,

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�������-! (Figs 9–11: Representative palynomorphs from the Lower Karoo sequence in STRAT 1 borehole. Slides are kept at the Laboratoire dePaléontologie, Université de Bretagne Occidentale, in Brest, France. Palynomorph locations on slides are recorded as England Finder (EF) coordinates.)Pollen and spores of the Platysaccus papilionis (A–F) and the Cyclogranisporites gondwanensis assemblage zones (G–O). • A – Lophotriletes rectusBharadwaj & Salujha, 1964 (slide: 1383, EF: J34). • B – Vestigisporites ventrisaccatus Anderson, 1977 (slide: 1556, EF: W42.2). • C – Alisporites ovatus(Balme & Hennelly) Jansonius, 1962 (slide: 1551, EF: O32.4). • D – Platysaccus papilionis Potonié & Klaus, 1954 (slide: 1554, EF: E47).• E – Lueckisporites virkkiae Potonié & Klaus, 1954 (slide: 1555, EF: E21). • F – Striatopodocarpites fusus (Balme & Hennelly) Potonié, 1958 (slide:1579, EF: V17.4). • G – Deltoidospora directa (Balme & Hennelly) Norris, 1965 (slide: 1562, EF: C19.4). – H Cyclogranisporites gondwanensisBharadwaj & Salujha, 1964 (slide: 1599, EF: P39.3). • I – Granulatisporites papillosus Hart, 1965 (slide: 1600, EF: U22.1). • J – Horriditriletes ramosus(Balme & Hennelly) Bharadwaj & Salujha, 1964 (slide: 1608, EF: T30.4). • K – Indotriradites australensis (Hart) Millsteed, 1999 (slide: 1578, EF:P24.2). • L – Lundbladispora braziliensa (Pant & Srivastava) emend. Marques-Toigo & Picarelli, 1984 (slide: 1599, EF: K42). • M – Alisporites potoniei(Lakhanpal, Sah & Dube) Somers, 1968 (slide: 1551, EF: O32.4). • N – Protohaploxypinus limpidus (Balme & Hennelly) Balme & Playford, 1967 (slide:1562, EF: P24.4). • O – Striatopodocarpites cancellatus (Balme & Hennelly) Hart, 1963 (slide: 1578, EF: Q22). Scale bar corresponds to 10 µm.

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Dwyka Group), by Key et al. (1995) based on studies fromsouthwest Botswana, as well as Visser (1990) who studiedglacigene deposits from southern Africa. Souza (2006) givesa brief account of limited radiometric ages from the DwykaGroup, which also indicates late Carboniferous stages.

Preliminary correlation of the biozones from STRAT 1and CKP 6 with other relevant palynological studies fromGondwana sequences (e.g., Wright & Askin 1987; ECL1990; Hankel 1992; Farabee et al. 1991; Lindström 1995a, b;Souza 2006) generally reveal an Early Permian age forBiozone KK 1, whereas the biozones KK 2 and KK 3 are oflate early Permian to early Middle Permian age. There are,however, difficulties in defining stage boundaries, whichrequires more refined data to allow direct correlation atshort stratigraphic intervals. Nevertheless, tentative age in-

ferences for the present zonation scheme from the KalahariKaroo Basin are made based on comparison with thezonation schemes for the Paraná Basin of Brazil in SouthAmerica (e.g., Di Pasquo et al. 2003a, b; Souza & Mar-ques-Toigo 2003, 2005; Souza 2006), and their calibrationby radiometric dates (Santos et al. 2006, Stephenson2008). It is also worth noting that the similarities indicatedby the assemblage taxa from the Kalahari Karoo Basin andthe Paraná Basin are also corroborated by the palaeo-geographic similarities of the two basins as reflected byseveral illustrations of their Late Palaeozoic reconstruction(e.g., Kalkreuth et al. 1999, Turner 1999, Wopfner 1999,Golonka & Ford 2000 etc.).

As described earlier, the KK 1 Biozone globally corre-lates with the Vittatina costabilis Interval Zone (Souza

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������� 5! Pollen and spores of the Hamiapollenites bullaeformis Assemblage Zone. • A – Punctatisporites gracilis Anderson, 1977 (slide: 1602, EF:H47.1). • B – Retusotriletes diversiformis (Balme & Hennelly) Balme & Playford, 1967 (slide: 1603, EF: S20.1). • C – Lophotriletes rarus Bharadwaj& Salujha, 1964 (slide: 1619, EF: F37). • D – Cristatisporites microvacuolatus (Dias-Fabrício) emend. Picarelli & Dias-Fabrício, 1990 (slide: 1598, EF:D39.3). • E – Raistrickia crenata Foster, 1979 (slide: 1627, EF: N37.4). • F – Cristatisporites morungavensis (Dias-Fabrício) emend. Picarelli& Dias-Fabrício, 1990 (slide: 1622, EF: J31.2). • G – Densosporites rotundus Falcon, 1978 (slide: 1581, EF: R21.4). • H – Weylandites magmus (Bose& Kar) Backhouse, 1991 (slide: 1563, EF: V30.3). • I – Limitisporites rectus Leschik, 1956 (slide: 1571, EF: T24). • J – Apiculatisporis parmatus (Balme& Hennelly) comb. nov. (slide: 1603, EF: V49.3). • K – Vittatina scutata (Balme & Hennelly) Bharadwaj, 1962 (slide: 1565, EF: Q23).• L – Hamiapollenites bullaeformis (Samoilovich) Jansonius, 1962 (slide: 1623, EF: T35.4). Scale bar corresponds to 10 µm.

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2006). From the latter correlation, Early Permian age rang-ing from the Asselian stage to the Artinskian stage are de-duced. The upper part of KK 1 Biozone plus KK 2 andKK 3 biozones generally correlate with the Lueckisporitesvirkkiae Interval Zone (Souza 2006), and in the Parana Ba-sin, Shrimp U-Pb zircon dating of the Irati Formation re-vealed an age of 278.4 Ma (Artinskian; Gradstein et al.2004) for the base of the Luekisporites virkkiae Zone(Santos et al. 2006). Consequently, a tentative late EarlyPermian to earliest Middle Permian age range is proposedfor the upper biozones KK 2 and KK 3. A rare absolute ageof 270 Ma reported from the Collingham Formation of

South Africa (e.g., Turner 1999), which overlies theWhitehill Formation, supports a late Early Permian age(i.e. Kungurian) for the uppermost part of Biozone KK 2.The latter represent a lithofacies section indicated in John-son et al. (1996) to immediately overlie the Whitehill For-mation in the Kalahari Karoo Basin and, hence probablyequates with the Collingham Formation. Comparisons ofthe assemblage characteristic of Biozone KK 3 with sev-eral regional and Gondwana-wide studies indicates MiddlePermian (Guadalupian) ages (e.g., Wright & Askin 1987,Farabee et al. 1991, Hankel 1992, Lindström 1995a). Asdiscussed above, the top of Biozone KK 2, is considered to

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������� ! A selected few significant pollen and spores used to determine the range zones for STRAT 1 and CKP6 boreholes. • A – Punctatisporitesubischii Foster, 1979 comb. nov. (slide: 1571, EF: T47.4). • B – Cirrabaculisporites lageniformis (Anderson) comb. nov. (slide: 1564, EF: K38.3).• C – Laevigatosporites vulgaris (Ibrahim) Ibrahim, 1933 (slide: 1580, EF: G33). • D – Converrucosisporites irregularis (Anderson) comb. nov. (slide:1643, EF: U43.3). • E – Laevigatosporites plicatus Kar, 1968 (slide: 1561, EF: T30.3). • F – Potonieisporites congoensis Bose & Maheshwari, 1968(slide: 1629, EF: K17.2). • G – Horriditriletes gondwanensis (Tiwari & Moiz) Foster, 1975 (1580, EF: E19). • H – Vittatina sp. (slide: 1621, EF: Y26.2).• I – Pakhapites fusus (Bose & Kar) Menéndez, 1971(slide: 1613, EF: W37.1). • J – Divaricrassus minor (Anderson) Millsteed, 1999 (slide: 1599, EF:R22.4). • K – Weylandites lucifer (Bharadwaj & Salujha) Foster, 1975 (slide: 1558, EF: O37.1). • L – Marsupipollenites triradiatus Balme & Hennelly,1956 (slide: 1559, EF: M44.1). Scale bar corresponds to 10 µm (except for Fig. 11F where the scale bar represents 20 µm).

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represent late Early Permian age, therefore, the succeedingBiozone KK 3 is considered to represent early MiddlePermian ages (i.e. Roadian). Consequently, the base ofBiozone KK 3 is tentatively equated with the boundary be-tween Early and Middle Permian for the sequence atSTRAT 1 borehole in the Kalahari Karoo Basin.

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The Lower Karoo sequence of the Karoo Supergroup in theKalahari Karoo Basin contains relatively well preserved po-llen and spore palynomorphs suitable for biostratigraphicclassifications. Taxa range analysis, determined on the basisof first and last occurrences datum, revealed the occurrenceof long, medium, and short range species, which allowed forthe subsequent erection of biostratigraphic units or biozo-nes. Three assemblage zones have been erected for each ofthe study sections i.e. STRAT 1 and CKP 6 boreholes. Al-though there appears to be considerable variation in taxaranges between the sections considered, the frequent occur-rence of common events nevertheless highlights great po-tential for palynological comparisons. A preliminary directcorrelation between STRAT 1 and CKP 6 borehole sectionsis attainable on the basis of the three assemblage zoneswhich are common to both sections. The assemblage zoneserected are comprised of, in ascending order of stratigraphy,the Hamiapollenites bullaeformis Assemblage Zone (Bio-zone KK 1), the Cyclogranisporites gondwanensis Assem-blage Zone (Biozone KK 2), and the Platysaccus papilio-nis–Striatopodocarpites fusus Assemblage Zone (BiozoneKK 3). Only limited comparisons and broad correlationswith other local previous palynology studies have been es-tablished, owing to the non-availability of taxa range data inmost such studies. Comparison and tentative correlation ofthe zones with similar zones erected in the region, as well asGondwana-wide, have been established, albeit cautiously,in recognition of several limiting factors that commonlyhamper long-distance correlations. Amongst the notable li-miting factors is the disparity in the frequency distribution oftaxa between sections, also relatively common in local cor-relations, as well as factors of global scale such as palaeoge-ographic restriction of flora or provincialism, which is con-sidered to have been well established during the latestCarboniferous to Permian times. Generally, there appears tobe a closer degree of similarity between assemblages fromthe Paraná Basin of South America and those from the Kala-hari Karoo Basin. Age control for the zonation schemeserected in this study is inferred from comparison with simi-lar assemblages from previous Gondwana studies, andhence remains tentative. The overall age range is from theLate Carboniferous (Kasimovian–Gzhelian) to the latestEarly or possibly earliest Middle Permian (Late Cisularianto Early Guadalupian).

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This paper is published with the permission of the Director of theDepartment of Geological Survey, Botswana. It is part of a Ph.D.study prepared under the guidance of A. Le Hérissé, Université deBrest, France. B. Modie wish to extend his gratitude to J.-J.Tiercelin and B. Le Gall, also of the Université de Brest, who pro-vided assistance and helpful discussions during the study, butmost importantly played a pivotal role in initiating collaborationbetween the Botswana Department of Geological Survey and theUniversité de Brest. Many thanks go to M.H. Stephenson of theBritish Geological Survey (Keyworth, U.K.), J. Broutin of theUniversity of Paris (Jussieu), and B.D. Millsteed, then of the Uni-versity of Durban-Westville (Durban, South Africa), all of whomprovided valuable comments, suggestions, and assistance duringthe course of the study. F. Paris and R. Brocke are also thanked fortheir helpful comments on the earlier version of the manuscript.

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1. Calamospora aplata Bharadwaj & Salujha, 19642. Concavisporites mortonii (de Jersey) de Jersey, 19623. Concavisporites sp.4. Deltoidospora directa (Balme & Hennelly) Norris, 19655. Pachytriletes splendens (Saksena) Millsteed, 19996. Punctatisporites gretensis Balme & Hennelly 19567. Punctatisporites gracilis Anderson, 19778. Punctatisporites parvus Anderson, 19779. Punctatisporites ubischii Foster 1979 comb. nov.

10. Punctatisporites sp.11. Retusotriletes diversiformis (Balme & Hennelly) Balme &

Playford, 196712. Retusotriletes golatensis Staplin, 196013. Cyclogranisporites gondwanensis Bharadwaj & Salujha,

196414. Cyclogranisporites sp.15. Granulatisporites papillosus Hart, 196516. Granulatisporites angularis (Staplin) Ybert, 197517. Granulatisporites trisinus Balme & Hennelly, 195618. Granulatisporites sp.19. Granulatisporites sp. A20. Anacanthotriletes sp.21. Anacanthotriletes sp. A22. Apiculatisporis cornutus (Balme & Hennelly) Høeg & Bose,

196023. Apiculatisporis unicus (Tiwari) Bharadwaj & Srivastava

comb. nov.24. Apiculatisporis levis (Balme & Hennelly) Segroves, 197025. Apiculatisporis leptoacaina (Jones & Truswell) comb. nov.26. Apiculatisporis parmatus (Balme & Hennelly) comb. nov.27. Didecitriletes eoericiana (Anderson) Millsteed, 199928. Didecitriletes ericianus (Venkatachala & Kar, 1965)29. Divaricrassus minor (Anderson) Millsteed, 199930. Lophotriletes rectus Bharadwaj & Salujha, 196431. Lophotriletes novicus Singh, 196432. Lophotriletes rarus Bharadwaj & Salujha, 196433. Lophotriletes sp.34. Lophotriletes sp. A35. Cirrabaculisporites lageniformis (Anderson) comb. nov.36. Cirrabaculisporites plumsteadi (Hart) Anderson comb. nov.37. Cirrabaculisporites ramosus Maheshwari & Bose, 196938. Cirrabaculisporites sp.39. Horriditriletes brevis Bharadwaj & Salujah, 196440. Horriditriletes ramosus (Balme & Hennelly) Bharadwaj &

Salujha, 196441. Horriditriletes tereteangulatus (Balme & Hennelly) Back-

house, 1991

42. Horriditriletes uruguaiensis (Marques-Toigo) Archangel-sky & Gamerro, 1979

43. Horriditriletes curvibaculosus Bharadwaj & Salujha, 196444. Horriditriletes gondwanensis (Tiwari & Moiz) Foster, 197545. Horriditriletes sp.46. Horriditriletes sp. A47. Raistrickia crenata Foster, 197948. Converrucosisporites naumoviae (Hart) Backhouse, 199149. Converrucosisporites pseudoreticulatus (Balme & Hen-

nelly) comb. nov.50. Converrucosisporites irregularis (Anderson) comb. nov.51. Converrucosisporites sp.52. Converrucosisporites sp. A53. Verrucosisporites morulatus (Knox) emend. Smith & But-

terworth, 196754. Verrucosisporites andersonii (Anderson) Backhouse, 198855. Verrucosisporites trisecatus Balme & Hennelly, 195656. Verrucosisporites sp.57. Verrucosisporites sp. A58. Convolutispora intrareticulata (Anderson) comb. nov.59. Convolutispora sp.60. Convolutispora sp. A61. Convolutispora sp. B62. Foveosporites karrooensis Anderson comb. nov.63. Foveosporites sp.64. Foveosporites sp. A65. Foveosporites sp. B66. Microreticulatisporites sp.67. Microreticulatisporites sp. A68. Clavatisporites sp.69. Indotriradites australensis (Hart) Millsteed, 199970. Cristatisporites crassilabratus Archangelsky & Gamerro,

197971. Cristatisporites inconstans Archangelsky & Gamerro, 197972. Cristatisporites lestai Archangelsky & Gamerro, 197973. Cristatisporites microvacuolatus (Dias-Fabrício) emend. Pi-

carelli & Dias-Fabrício, 199074. Cristatisporites morungavensis (Dias-Fabrício) emend. Pi-

carelli & Dias-Fabrício, 199075. Cristatisporites spinosus (Menendez & Azcuy) Playford,

197876. Cristatisporites menendezii Playford, 197877. Cristatisporites sp.78. Cristatisporites sp. A79. Densoisporites sp.80. Densosporites rotundus Falcon, 197881. Densosporites sp.

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82. Lundbladispora braziliensa (Pant & Srivastava) emend.Marques-Toigo & Picarelli, 1984

83. Lundbladispora gracila Stephenson & Osterloff, 200284. Vallatisporites sp.85. Vallatisporites sp. A86. Laevigatosporites vulgaris (Ibrahim) Ibrahim, 193387. Laevigatosporites plicatus Kar, 196888. Polypodiisporites mutabilis Balme, 197089. Polypodiisporites detritus (Leschik) Anderson, 197790. Florinites eremus Balme & Hennelly, 195591. Striomonosaccites brevis Bose & Kar, 196692. Striomonosaccites crucistriatus Ybert, 197593. Striomonosaccites sp.94. Barakarites rotatus (Balme & Hennelly) Bharadwaj & Ti-

wari, 196495. Bascanisporites undosus Balme & Hennelly, 195696. Cannanoropollis janakii Potonie & Sah, 196097. Cannanoropollis sp.98. Plicatipollenites gondwanensis (Balme & Hennelly) Lele,

196499. Plicatipollenites densus Srivastava, 1970

100. Plicatipollenites trigonalis Lele, 1964101. Plicatipollenites sp.102. Plicatipollenites sp. A103. Plicatipollenites sp. B104. Caheniasaccites ovatus Bose & Kar, 1966105. Caheniasaccites flavatus Bose & Kar, 1966106. Potonieisporites novicus Bhardwaj, 1954107. Potonieisporites brasiliensis (Nahuys, Alpern & Ybert) Ar-

changelsky & Gamerro, 1979108. Potonieisporites congoensis Bose & Maheshwari, 1968109. Illinites spectabilis (Leschik) comb. nov.110. Limitisporites rectus Leschik, 1956111. Limitisporites monstruosus (Luber in Luber & Waltz) Hart,

1965112. Limitisporites perspicuous (Leschik) comb. nov.113. Limitisporites sp.114. Vestigisporites rudis Balme & Hennelly, 1955115. Vestigisporites ventrisaccatus Anderson, 1977116. Alisporites ovatus (Balme & Hennelly) Jansonius, 1962117. Alisporites australis de Jersey, 1962118. Alisporites potoniei (Lakhanpal, Sah & Dube) Somers, 1968119. Alisporites splendens (Leschik) Foster, 1979120. Platysaccus papilionis Potonié & Klaus, 1954121. Platysaccus sp.122. Pteruchipollenites sp.123. Hamiapollenites bullaeformis (Samoilovich) Jansonius, 1962124. Hamiapollenites fusiformis (Marques-Toigo) emend. Ar-

changelsky & Gamerro, 1979125. Hamiapollenites saccatus Wilson, 1962126. Hamiapollenites tractiferinus (Samoilovich) Jansonius, 1962127. Hamiapollenites sp.128. Lueckisporites virkkiae Potonié & Klaus, 1954129. Lunatisporites nubilus (Leschik) comb. nov.130. Lunatisporites variesectus Archangelsky & Gamerro, 1979131. Lunatisporites sp.132. Protohaploxypinus haigii Foster, 1979

133. Protohaploxypinus hartii Foster, 1979134. Protohaploxypinus limpidus (Balme & Hennelly) Balme &

Playford, 1967135. Protohaploxypinus perexiguus (Bharadwaj & Salujha) Fos-

ter, 1979136. Protohaploxypinus rugatus Segroves, 1969137. Protohaploxypinus sp.138. Striatoabieites multistriatus (Balme & Hennelly) Hart, 1964139. Striatopodocarpites cancellatus (Balme & Hennelly) Hart,

1963140. Striatopodocarpites fusus (Balme & Hennelly) Potonié,

1958141. Striatopodocarpites pantii (Jansonius) Balme, 1970142. Striatopodocarpites rarus (Bharadwaj & Salujha) Balme,

1970143. Ephedripites sp.144. Ephedripites sp. A145. Vittatina densa Anderson, 1977146. Vittatina foveolata Tschudy & Kosanke, 1966147. Vittatina scutata (Balme & Hennelly) Bharadwaj, 1962148. Vittatina minima Jansonius, 1962149. Vittatina saccifer Jansonius, 1962150. Vittatina subsaccata Samoilovich, 1953151. Vittatina sp.152. Vittatina sp. A153. Weylandites lucifer (Bharadwaj & Salujha) Foster, 1975154. Weylandites magmus (Bose & Kar) Backhouse, 1991155. Pakhapites fusus (Bose & Kar) Menéndez, 1971156. Pakhapites ovatus (Bose & Kar) Menéndez, 1971157. Gnetaceaepollenites sp.158. Pretricolpipollenites sp.159. Marsupipollenites striatus (Balme & Hennelly) Foster, 1975160. Marsupipollenites triradiatus Balme & Hennelly, 1956161. Cycadopites cymbatus (Balme & Hennelly) Hart, 1965162. Cycadopites glaber (Luber & Waltz) Hart, 1965163. Cycadopites nevesi (Hart) Hart, 1965164. Cycadopites follicularis Wilson & Webster, 1946165. Cycadopites sp.166. Inapertisporites inapertus (Anderson) Falcon, 1978167. Micrhystridium sp.168. Tetraporina sp.169. Veryhachium sp. A170. Veryhachium sp. B171. Veryhachium sp. C172. Species A173. Species B174. Species C175. Species D176. Species E177. Botryoccocus sp. cf. B. braunii178. Arabisphaera bellula Hemer & Nygreen, 1967179. Arabisphaera sp.180. Cymatiosphaera gondwanensis (Tiwari) Backhouse, 1991181. Cymatiosphaera sp. A182. Cymatiosphaera sp. B183. Maculatasporites sp.184. Tasmanites sp.

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