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Permian palynostratigraphy: a global overview

MICHAEL H. STEPHENSON British Geological Survey, Keyworth, Nottingham NG12 5GG, UK (e-mail: [email protected])

Abstract: Permian palynostratigraphic schemes are used primarily to correlate coal- and hydro- carbon-bearing rocks within basins and between basins, sometimes at high levels of biostrati- graphic resolution. Up to now, their main shortcoming has been the lack of correlation with schemes outside the basins, coalfields and hydrocarbon fields that they serve, and chiefly a lack of correlation with the international Permian scale. This is partly because of phytogeographical provinciality from the Guadalupian onwards, making correlation between regional palynostrati- graphic schemes difficult. However, local high-resolution palynostratigraphic schemes for regions are now being linked either by assemblage-level quantitative taxonomic comparison or by the use of single well-characterized palynological taxa that occur across Permian phytogeographical provinces. Such taxa include: Scutasporites spp., Vittatina spp., Weylandites spp., Lueckisporites virkkiae, Otynisporites eotriassicus and Converrucosisporites confluens. These palynological cor- relations are being facilitated and supplemented with radiometric, magnetostratigraphic, indepen- dent faunal and strontium isotopic dating.

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Palynostratigraphy is the use of palynomorphs important biostratigraphic markers for the Permian (defined as organic-walled microfossils 5–500 mm of Gondwana and include several hundred species. in diameter) in correlating and assigning relative It is estimated that by the Lopingian about 60% of ages to rock strata. As such, it is a branch of biostra- the world’s flora consisted of seed plants (Gradstein tigraphy and follows the rules of biostratigraphic & Kerp 2012). practice: for example, those set out by Rawson These large-scale evolutionary changes in plants, et al. (2002). filtered by local and regional effects, are responsible The Permian, falling between 252.2 and 298.9 Ma, for the palynological succession that provides was a period of intense change in which the giant opportunities for subdivision on which palynostrati- continent of Pangea as a whole moved north, and graphic schemes are built. However, the pro- in which, through the early part of the Period, a tran- nounced phytogeographical differentiation of the sition from icehouse to greenhouse conditions occur- Permian has a powerful effect on palynostratigra- red (e.g. Fielding et al. 2008), alongside the decline phy, such that schemes differ considerably across in coal swamps and the establishment of widespread Pangea and correlation between schemes is even evaporite deposits (Henderson et al. 2012). The end now tentative or incomplete. In the Gondwana phy- of the Period saw a major extinction of fauna such as togeographical province, for example, it is difficult fusulinacean foraminifers, trilobites, rugose and tab- to correlate to the standard Permian stages; and ulate corals, blastoids, acanthodians, placoderms, the Carboniferous–Permian and Permian–Triassic and pelycosaurs; a dramatic reduction in bryozoans, boundaries are not precisely correlateable into brachiopods, ammonoids, sharks, bony fish, cri- Gondwana basins using palynology (Stephenson noids, eurypterids, ostracodes and echinoderms 2008a). (Henderson et al. 2012); and, although many coni- Until recently, progress in correlation was ham- fers (e.g. glossopterids, cordaites) became extinct pered by the lack of fundamental stratigraphic stan- at the end of the Permian, there is no evidence of dards such as stage Global Stratigraphic Sections major extinction in the plants (Gradstein & Kerp and Points (GSSPs); however, since 1997 (Jin 2012). Amongst the most important changes in et al. 1997; Henderson et al. 2012) a number of land plants is the replacement, near the end of the GSSPs have been established within the Pennsylva- Carboniferous, of arborescent lycophytes by arbo- nian–Permian succession, the most important of rescent tree ferns; arborescent lycophytes only which is the basal Permian GSSP at Aidaralash persisted into the Guadalupian in China. The arbo- Creek in the southern Urals (Jin et al. 1997; Hender- rescent horsetails also declined by the end of the Car- son et al. 2012), and the basal Triassic GSSP at boniferous. In the Permian, a great variety of new Meishan section D, Changxing County, Zhejiang seed plant groups appeared such as cycads, ginkgos, Province, South China (Yin et al. 2001). Since voltzialean conifers and glossopterids. The latter are these developments, there have also been other

From:Lucas,S.G.&Shen, S. Z. (eds) The Permian Timescale. Geological Society, London, Special Publications, 450, https://doi.org/10.1144/SP450.2 # 2016 The Author(s). Published by The Geological Society of London. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON advances contributing to the precision and utility of The approach taken in this paper is to sur- palynological biostratigraphy in this interval, vey the palynostratigraphic schemes in the main including radiometric and faunal dating of palyno- phytogeographic provinces and then to attempt logical biozones, and limited high-resolution corre- synthesis; and so the focus is on palynostratigraphy lation between continents using a well-defined not taxonomy. Given the plethora of palynological palynological species. literature on this interval, the review is necessarily Palynological research in the Permian is exten- selective. Most recent published palynostratigraphic sive, being partly driven by exploration for coal (e.g. schemes (e.g. since 2000) have emanated from in India and Australia), and oil and gas (e.g. in the South American and Middle Eastern basins. In Middle East, South America, Australia and the the following account, age assignments related to Barents Sea), but has tended to be regional or local these and other schemes reflect those of the original in focus (see Truswell 1980). A number of authors authors but may not necessarily use modern chrono- (Bharadwaj 1969; Kemp 1975; Bharadwaj & Sri- stratigraphic nomenclature, thus a variety of strati- vastava 1977; Balme 1980; Truswell 1980; Utting & graphic stage and other nomenclature is used in Piasecki 1995; Warrington 1996; Price 1997; Play- this paper. For the convenience of the reader, a ford & Dino 2005; Azcuy et al. 2007; Stephenson chart showing correlations of the main chronostrati- 2008a) have attempted to summarize the research or graphic subdivisions used internationally is shown to correlate the main biozones across regions, but in Figure 1. correlation has been tentative. Among the difficul- Permian palynostratigraphic schemes use pollen ties acknowledged by these previous reviewers are and spores almost exclusively. While it is recog- disparate stratigraphic and taxonomic methods nized that marine palynomorphs (acritarchs) may practised in different parts of the world, and differ- be present in Permian rocks, no study has sought ent standards of documentation of palynological data. to produce a palynostratigraphy based purely on

StandardRussia Tethys Western Europe China North America

Changhsingian ? Changhsingian 254.2 Dorashamian Thuringian ? Wuchiapingian Wuchiapingian ? Dzhulfian Lopingian Late Permian 259.8 Ochoan

Capitanian Capitanian Tatarian Midian 265.1 Saxonian Maokouian Wordian Wordian

Guadalupian Middle Permian 268.8 Murghabian ? Roadian Kazanian Roadian 272.3 Ufimian

Kubergandian Rotliegend Kungurian Kungurian Leonardian Bolorian 279.3 Luodianian

Arnskian Arnskian Yakhtashinian

Autunian 290.1 Wolfcampian Early Permian Cisuralian Sakmarian Sakmarian Sakmarian

295.5 Chuanshanian

Asselian Asselian Asselian 298.9

Fig. 1. Chronostratigraphy of the Permian, modified after Henderson et al. (2012). Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Permian acritarchs, although they may show future plant macrofossils differently (Utting & Piasecki potential (e.g. Lei et al. 2013). 1995). Balme (1970), Sullivan (1965), Turnau The range of morphology seen in palynomorphs (1978) and Van der Zwan (1981) surveyed the haz- in the Permian is illustrated simply in Figure 2. ards of the reconstruction of palaeophytogeograph- To improve readability, names of authors of taxa ical provinces by palynology. The value of pollen are excluded from the main text of the paper, but and spore taxa as indices of low-rank plant taxa is the main taxa and their authorship are listed in limited because the plant affinities of most Palaeo- Appendix A. zoic spore and pollen genera and species are unknown, and because botanical and palynological taxonomy are independent of one another. Despite Phytogeography of the Permian this, the broad palynological characteristics of a region at a certain time are thought to be representa- Phytogeographical provinciality makes correlation tive of the high-rank palaeobotanical characteristics difficult because it tends to reduce the number of of that region (Utting & Piasecki 1995). taxa in common between assemblages in different In broad terms, there was a gradual diversifica- provinces. In general, it seems reasonable to expect tion of phytogeographical provinces from relatively geographical parity between palaeobotanical prov- uniform palaeophytogeography in the Devonian to inces and palynological provinces since plants maximum provinciality in the Lopingian (Cleal & and palynomorphs are biologically linked (Hart Thomas 1991) when four main provinces existed: 1970). However, palynomorphs are subject to much Gondwana, Euramerica, Angara and Cathaysia wider distribution than plant remains; similar or (Fig. 3) (Utting & Piasecki 1995). Palaeobotani- identical palynomorphs may be produced by unre- cally, the Gondwana phytogeographical province lated plants (Meyen 1969); and taphonomic factors is distinct from other provinces in the Permian may affect the preservation of palynomorphs and because of the abundance of the glossopterids

Fig. 2. Range of morphology in Permian palynomorphs. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Fig. 3. Phytogeographical provinces of the Permian on a Wordian reconstruction (from the Scotese Palaeomap Project); palaeophytogeography after Utting & Piasecki (1995). whose leaf impressions (Glossopteris and Ganga- palynology of Angara is characterized by the abun- mopteris) and more rare associated fruiting bodies dance of trilete spores rather than by bisaccate and (e.g. Plumsteadia and Scutum) are characteristic. taeniate bisaccate pollen, which are common in the Some conifers, sphenophytes and ferns were also other Permian phytogeographical provinces. Mono- present (Cleal & Thomas 1991). In the northern saccate pollen of Cordaitina is common in the lower parts of Gondwana, marattialean ferns and lyco- parts of the Permian in Angara, whereas the mono- phytes were present (Cleal & Thomas 1991). Paly- colpate pollen Cycadopites is common in the upper nologically, the Gondwana province is, perhaps, parts. the most distinct because of its diversity of taeniate Palaeobotanically, the Cathaysia province is and monosaccate pollen, and the occurrence of distinguished by its gigantopterids, noeggerathia- restricted genera not yet recorded outside the prov- lean-like progymnosperms and plants with cycad- ince: for example, Guttulapollenites, Microbaculis- like foliage; however, lycophytes, sphenophytes pora, Dulhuntyispora and Corisaccites (Truswell and pteridosperms were also present (Cleal & Tho- 1980). However, mixing of floras between Gond- mas 1991). Both palaeobotanists and palynologists wana, Euramerica and Cathaysia are suggested by report a similarity between Cisuralian floras of palaeobotanical studies (Archangelsky & Wagner Cathaysia and Pennsylvanian floras of Euramerica 1983), and such mixing has been observed by paly- (Kaiser 1976; Cleal & Thomas 1991; Utting & Pia- nologists (e.g. Kar & Jain 1975; Kaiser 1976; Eshet secki 1995). Kaiser (1976) interpreted the relict flora 1990; Nader et al. 1993a, b). as being due to the continuing palaeotropical condi- Palaeobotanically, the Euramerican province is tions of Cathaysia, which provided a refuge for the distinguished by its abundant conifers, marattialean tropical vegetation of the Pennsylvanian swamps ferns and pteridosperms (Cleal & Thomas 1991). In of Euramerica. the Lopingian, aridification caused the vegetation to Apart from the Cisuralian abundance of Pennsyl- become sparser and of lower diversity (Schweitzer vanian Euramerican palynological taxa in Cathay- 1986). Palynologically, a similar impoverishment sia, there are also endemic palynological taxa that of species occurs in the Lopingian (Pattison et al. characterize Cathaysia: for example, Nixispora 1973; Smith et al. 1974). British and west European and Patellisporites (Utting & Piasecki 1995). The assemblages are characterized by an abundance of Carboniferous ‘relict flora’ of Cathaysia appears conifer pollen of Lueckisporites virkkiae, and by to persist into the Lopingian in eastern Yunnan, the less abundant genera Protohaploxypinus, Stria- China (Ouyang 1982), where diverse assemblages toabieites, Striatopodocarpites, Taeniaesporites, of genera such as Torispora, Crassispora, Triqui- Vittatina, Falcisporites and Klausipollenites (Smith trites and Laevigatosporites occur. et al. 1974). According to Utting & Piasecki (1995), Angara was dominated by a cool-temperate flora containing Palynostratigraphy diverse herbaceous sphenophytes and Cordaitales, Euramerica with the most northerly part (Siberia) having a cold- temperate climate that spread to most of Angara The phytogeographical province of Euramerica is by the Lopingian. According to Hart (1970), the now represented by the areas west of the Ural Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Mountains, Europe, parts of North Africa and North Russian Permian palynostratigraphy, compared America, and contains the historical type area for assemblages from the Ufimian and Kazanian type the Permian and the present type area for the Early areas in the southern Urals around Perm and west Permian (Cisuralian) in the southern Urals, includ- towards Kazan, with the sub-Angaran palynological ing the base Permian GSSP at Aidaralash Creek, biozones of the Sverdrup Basin of the Canadian southern Urals. Arctic. The Ufimian, an original Russian stage Knowledge of the palynostratigraphy of the recently abandoned by the All Russian Stratigraphic Permian of the southern Urals is hampered by the Commission (see Henderson et al. 2012), may form lack of work since the 1980s. Most work before part of the upper Kungurian, and the Kazanian is that date was published in Russian, but was conve- considered equivalent to the Roadian (Fig. 1). Well- niently summarized by Hart (1970) and Warrington preserved palynological assemblages occur in the (1996). However, the work was regionally based Ufimian and Kazanian, but Utting et al. (1997) and not gathered into a Urals-wide palynostrati- recovered no palynomorphs from the Tatarian of graphic scheme, perhaps because other palaeonto- the area. logical groups (e.g. ammonoids and fusulinacean Utting et al. (1997) found that many taxa range foraminifers) already provided ample resolution through the Ufimian and Kazanian: for example, for stratigraphic subdivision. Florinites luberae, Cordaitina uralensis, Alisporites Only a short, preliminary palynostratigraphic plicatus, Limitisporites monstruosus, Hamiapollen- study has been carried out at Aidaralash Creek ites tractiferinus and Weylandites striatus. Several (Dunn 2001). Dunn (2001) described a section of taxa appear first in the lower Kazanian (e.g. Lueck- approximately 50 m of strata from 24.2 m below, isporites virkkiae), while Crucisaccites ornatus, to 26 m above, the Carboniferous–Permian boun- Weylandites cincinnatus and Hamiapollenites bul- dary. The assemblages contain abundant Vittatina laeformis disappear in the lower Kazanian. Taxa (particularly Vittatina costabilis) and taeniate bisac- of note that are common in the Ufimian and Kaza- cate pollen. The non-taeniate bisaccate pollen Limit- nian sequences include Protohaploxypinus per- isporites monstruosus is common throughout and fectus, Weylandites striatus, Alisporites plicatus, spores are rare. No marked palynological change Limitisporites monstruosus and Hamiapollenites occurs at the Carboniferous–Permian boundary at tractiferinus (see Utting et al. 1997, fig. 7). At the the base of Bed 19.5 where the conodont Streptog- suprageneric level, it appears that Ufimian assem- nathodus isolatus first appears (Davydov et al. blages are more spore-rich than those of the Kaza- 1998). nian; the latter is characterized by abundant Faddeeva (1980) in a larger survey of southern taeniate bisaccate pollen, and common non-taeniate Urals palynology considered that Gzhelian (latest and polyplicate pollen (see Utting et al. 1997, fig. 8). Carboniferous) and Asselian assemblages are dis- Although Utting et al. (1997) did not recover tinguished by changes in proportions of spores and palynomorphs from the Tatarian in the southern pollen, in that the Gzhelian assemblages have com- Urals, they referred to other studies (see Utting mon spores and few Vittatina specimens and pollen, et al. 1997, p. 5) that characterize the Tatarian as while Asselian assemblages have common saccate being dominated by species of Protohaploxypinus, pollen (e.g. Cordaitina and Potonieisporites) and Vittatina and Florinites luberae. Gomankov (1992) Vittatina. A similar distinction is apparent to the refers to the presence in the lower Tatarian of Scu- SW in the Donetz Basin according to Inosova tasporites unicus, which is similar to Scutasporites et al. (1975). nanuki from the Wordian of the Sverdrup Basin, Later assemblages from the Sakmarian of the Canada (see later discussion); and Utting et al. Urals are dominated by saccate pollen, while Artin- (1997) noted some similarities between the Russian skian assemblages contain common spores includ- and Canadian assemblages, although they also ing Tuberculatosporites and Granulatisporites,as noted quantitative differences (see Utting et al. well as pollen (Cordaitina and Cycadopites) (Hart 1997, fig. 8), which they attributed to palaeoclimatic 1970). According to Hart (1970), the Kungurian of differences. the Urals is characterized by taeniate bisaccate pol- Gomankov et al. (1998) described few palyno- len, and the Kazanian by more diverse assemblages logical differences between the Tatarian and Kaza- of monosaccate and taeniate bisaccate pollen and nian, but identified four broad assemblage zones an increase in non-taeniate bisaccate pollen. Fad- for the Tatarian. In ascending order, Zone I is deeva (1980) also noted a change at the base of characterized by Weylandites and Lueckisporites the Kazanian, including an increase in taeniate and virkkiae, and species of Protohaploxypinus, Tae- non-taeniate taxa, as well as the appearance of niaesporites and Vittatina, as well as forms related Lueckisporites, Taeniaesporites and Vesicaspora. to Vittatina (e.g. Ventralvittatina and Duplivitta- Utting et al. (1997), in an important paper that tina). Zone II lacks Ventralvittatina and Duplivitta- constitutes one of the few modern surveys of tina, and contains the first appearance of spore Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON specimens similar to the distinctive Limatulaspor- which range from late Stephanian A to Stephanian ites fossulatus, as well as Cladaitina and Cordai- D; and two higher biozones (a lower Vittatina tina. Zone III contains fewer specimens of costabilis (VCI) Zone and an upper Vittatina costa- Cladaitina and Cordaitina, but also contains rare bilis (VCII) Zone) that Hochuli (1985) considered Scutasporites unicus. Zone IV sees the first appear- together to be equivalent to the Vittatina costabilis ance of Cedripites priscus, as well as sculptured (VC) Zone of Clayton et al. (1977) (Fig. 4). monolete spores of the genus Punctatosporites. Visscher (1980) and Edwards et al. (1997) Much of the European sediments of the Cisura- indicated that a long hiatus (the ‘post-Variscan lian to the west of the southern Urals were deposited interval’) exists between the Autunian and the in restricted basins and are therefore difficult to cor- Thuringian (Lopingian) of Europe (Fig. 1), although relate with beds of similar age in Russia (Utting & Schaarschmidt (1980) reported a Kungurian–Kaza- Piasecki 1995). Doubinger (1974) and Clayton nian assemblage from Germany. Apart from this et al. (1977) defined biozones based in the Carbon- assemblage, palynological assemblages from the iferous–Permian rocks of France and Germany. Guadalupian of Europe appear to be rare: however, Balme (1980) related the base of the Disaccites palynological assemblages from the Lopingian rocks Striatiti (DS) Zone of Clayton et al. (1977) to the of Western Europe have been intensively studied base of Unit III of Western Australia (see later dis- (Visscher 1973), but have been shown to be, palyno- cussion) due to the common expansion of the taeni- logically, rather homogeneous throughout the ate bisaccate pollen at these horizons. Doubinger sequence (Grebe & Schweitzer 1962; Schaarsch- et al. (1987) and Jerzykiewicz (1987) described a midt 1963; Clarke 1965; Smith et al. 1974); the only similar abundance of taeniate bisaccate pollen in notable change being a gradual overall impoverish- the Autunian (latest Pennsylvanian–Cisuralian: ment of the palynoflora occurring in the upper parts Fig. 1) of the Lode`ve Basin, France, and the Intrasu- of the sequence (Pattison et al. 1973). Although sev- detic Basin of SW Poland, respectively. eral important taxonomic studies emanate from the Hochuli (1985) documented four biozones based European Lopingian (e.g. Klaus 1963; Schaarsch- in the Carboniferous–Permian rocks of NE Switzer- midt 1963), possibilities for palynostratigraphic land. Hochuli (1985) recognized the Angulispori- subdivision lie with the variations within the tes splendidus–Latensina trileta (ST) and the Lueckisporites virkkiae palynodemes (lineages) of Potonieisporites novicus/bharadwaji–Cheiledonites Visscher (1971) and with changes in the relative major (NBM) biozones of Clayton et al. (1977), abundances of suprageneric groups in, for example,

Doubinger 1974 Clayton et al. 1977 Hochuli 1985

Upper Autunian A3 DS

VCII Lower Autunian A2 VC VCI

Stephanian C/D A1 NBM

Stephanian B ST

Stephanian A OT Westphalian D

Fig. 4. Correlations of western European Carboniferous–Permian palynostratigraphic schemes, after Hochuli (1985). Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY the Zechstein Sea Basin (Pattison et al. 1973). thorsteinssonii Scutasporites nanuki and the Road- Visscher (1971) erected six biozones within the ian Alisporites plicatus–Jugasporites compactus Thuringian that are based on a palynodeme of concurrent range zones (Fig. 5). The ages of the bio- Lueckisporites virkkiae. zones are based mainly on ammonoids, conodonts, Recent studies of European Thuringian sedi- brachiopods and foraminifers from the associated mentary rocks have concentrated on the palaeo- fossiliferous beds of shallow and deep basinal environment (e.g. Bercovici et al. 2009) or on the facies. palynological character of the Permian–Triassic Utting (1994) considered that the two Sverdrup boundary in the Alps (Looy et al. 2001; Twitchett Basin biozones were Sub-Angaran in character and et al. 2001; Spina et al. 2015). In North America, correlateable with assemblages in western Canada, only a small number publications, which concen- Alaska, Greenland, Svalbard, the Barents Sea, the trate on taxonomy, emanate from the Guadalupian Pechora Basin and the northern Russian Platform. and Lopingian, including those of Wilson (1962), However, the Wordian Ahrensisporites thorste- Jizba (1962), Tschudy & Kosanke (1966) and Clap- inssonii–Scutasporites nanuki Concurrent Range ham (1970). Zone differs from assemblages further south in the In North Africa, a palynostratigraphic scheme Urals in the type area of the Kazanian Stage (see for a Carboniferous–Permian transition sequence the previous discussion). The Sverdrup Basin assem- was defined by Brugman et al. (1988) from cuttings blages are dominated by trilete spores, non-taeniate samples from borehole A1-19 in NE Libya. On the and taeniate bisaccate pollen, and polyplicate taxa. basis of the ranges of taxa, Brugman et al. (1988) Utting’s detailed studies do not show large differ- proposed two assemblage biozones: a lower Assem- ences in the overall suprageneric character of assem- blage Zone A and an upper Assemblage Zone B. blages across the Roadian and Wordian (see Utting The top of the lowest biozone was defined by Brug- 1994, fig. 7), but a number of taxa are confined to man et al. (1988) as the stratigraphic level of the particular biozones including Crinalites sabinensis, disappearance of Lycospora pusilla, a spore type Cladaitina kolodae and Sverdrupollenites agluatus particularly characteristic of the Euramerican coal to the Alisporites plicatus–Jugasporites compactus belts of the Pennsylvanian. Brugman & Visscher Concurrent Range Zone, and Striatoabieites bore- (1988) and Brugman et al. (1988) believed this alis and the eponymous species to the Ahrensispor- level to approximate to the Permian–Carboniferous ites thorsteinssonii–Scutasporites nanuki Current boundary. The main difference across the boundary Range Zone. The latter biozone differs markedly between Assemblage Zone A and an upper Assem- from the succeeding Griesbachian Tympanicysta blage Zone B is the loss in the latter of taxa of Eur- stoschiana–Striatoabieites richteri Assemblage american affinity, including Endosporites ornatus, Zone, which was accounted for by Utting (1994) Crassispora kosankei and Spelaeotriletes spp. as being due to a sedimentary hiatus and climatic Brugman & Visscher (1988) also identified differences. Sakmarian–?Ufimian assemblages in A1-19.Assem- Mangerud (1994) established two biozones blages containing Lueckisporites virkkiae was re- for the Cisuralian–Guadalupian in the offshore cordedfromTunisia byKilani-Mazraouietal.(1990). Norway Finnmark Platform: the Dyupetalum sp.– Hamiapollenites bullaeformis Assemblage Zone Angara of ?Kungurian–Ufimian age, and the Kazanian– ?Tatarian Scutasporites cf. unicus–Lunatisporites Angara was divided into four by Utting & Piasecki spp. Assemblage Zone, as well as recognizing an (1995), including the Sub-Angara, Far Eastern, older biozone previously established in Spitsbergen Pechoran and Siberia subprovinces (Fig. 3). Sub- by Mangerud & Konieczny (1993) – the Cisuralian Angara occupied a more southerly position in Perm- Hamiapollenites tractiferinus Assemblage. The ian palaeogeographical reconstructions, while Sibe- Hamiapollenites tractiferinus Assemblage is domi- ria was to the NE, at high Permian latitudes. nated by species of Vittatina including V. costabilis The palynostratigraphy of Sub-Angara is the best and V. saccata, but also contains taeniate bisaccate studied of these subprovinces as a result of work pollen including Protohaploxypinus and Striatopo- carried out in the 1980s and 1990s in the Canadian docarpites ; and distally taeniate bisaccate pollen Arctic Sverdrup Basin by the Canadian Geological such as Hamiapollenites tractiferinus and H. bul- Survey, and work related to oil exploration in the laeformis. The Dyupetalum sp.–Hamiapollenites Barents Sea and Svalbard. bullaeformis and Scutasporites cf. unicus–Lunatis- Utting (1989) established five preliminary porites spp. assemblage zones are similar except for palynological biozones in the Sverdrup Basin from the occurrence in the upper biozone of Scutasporites shallow basin margin facies; later, Utting (1994) cf. unicus and the occurrence of Hamiapollenites refined the ages and detailed characteristics of bullaeformis in the lower biozone. According to two of the biozones: the Wordian Ahrensisporites Mangerud (1994) and Nilsson et al. (1996), the Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Standard Russia Arcc Canada, Arcc Canada, East Greenland, Spitsbergen, Barents Shelf, Ung 1889 Ung 1994 Piasecki 1984 Mangerud & Konieczny Mangerud 1994 1993

Changhsingian

Wuchiapingian ? Lopingian

Capitanian Tatarian ?

Wordian Ahrensisporites Protohaploxypinus Taeniaesporites sp. thorsteinssonii – Scutasporites Kraeuselisporites nanuki Guadalupian Guadalupian

Alisporites plicatus – Scutasporites cf. unicus – Roadian Kazanian Jugasporites ? Lunasporites spp. Alisporites insignis– compactus Triadispora sp.

Viana Dyupetalum sp. - Kungurian Kungurian Hamiapollenites bullaeformis

Limisporites monstruosus – V. costabilis Arnskian Arnskian

Hamiapollenites Hamiapollenites tracferinus tracferinus

Potonieisporites W. striatus – P.

Cisuralian Cisuralian perfectus Sakmarian Sakmarian

Asselian Asselian Potonieisporites spp. - Viana sp.

Fig. 5. Sub-Angara palynostratigraphic schemes, adapted from Utting (1989, 1994) and Mangerud (1994).

Finnmark Platform and other Barents Sea sequences Province at least prior to the Cisuralian. According can be correlated across the northern Atlantic, to to Zhu et al. (2005), the Junggar Basin Permian con- Spitsbergen, Greenland and the Canadian Arctic tains mainly terrestrial sediments and the Lower (1), (Fig. 5) (Mangerud & Konieczny 1993; Utting 1994). Middle (2) and Upper (3) Permian palynofloras are There is very little modern palynological work characterized, respectively, by: (1) the overwhelm- on the basins to the north of Sub-Angara. Hart ing dominance of taeniate bisaccate pollen; (2) the (1970) and Utting & Piasecki (1995) summarized high content of Cordaitina and taeniate bisaccate palynological work from the Taimyr, Kuznets and pollen; and (3) by the appearance of many newly Tungus areas in Siberia, concluding that the main evolved forms with a ‘Mesophytic aspect’. These difference between the lower and upper parts of latter include pollen and spores of ‘advanced coni- the Permian is that monosaccate pollen are replaced fers and ferns’ including distinctive taxa such as by monosulcate pollen such as Cycadopites. To the Lueckisporites virkkiae, Scutasporites xinjiangen- east of Angara, in the Far Eastern subprovince, sis, Klausipollenites schaubergeri, Falcisporites Utting & Piasecki (1995) summarized the general- zapfei, Eucommiidites, Dictyophyllidites and abun- ized palynology in the Xinjiang, Tianshan, Kunlun dant small bisaccate pollen (e.g. Vitreisporites). and Junggar areas. More recent work by Zhu et al. Zhu et al. (2005, fig. 2) showed stratigraphic ranges (2005) compared the Junggar Basin with the of important taxa in the Permian sequences of the Tarim Basin, which belonged to the Euramerican Junggar and Tarim basins. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Metcalfe et al. (2009) focused on the Permian– and the Platysaccus minus–Gulisporites cochlear- Triassic non-marine sequence at Dalongkou and ius (MC) assemblage zones. Liu et al. (2008) Lucaogou in the Junggar Basin, and defined three considered that the SL Assemblage Zone correlates assemblages, two of Lopingian age and an upper approximately with the western European SL Zone assemblage of probable Early Triassic age. The of Clayton et al. (1977: Westphalian C–D). The oldest is the Tuberculatosporites homotubercula- VK Assemblage Zone is believed to span the Car- ris–Potonieisporites sp. Q assemblage. Apart from boniferous–Permian boundary. According to Liu the eponymous species, this assemblage contains et al. (2008), the upper TH and MC assemblage monosaccate pollen, including Cordaitina uralen- zones are of Cisuralian age. Liu et al. (2011) sis, and rare specimens of Scutasporites cf. unicus. described Pennsylvanian–Lopingian megaspores The youngest Permian assemblage, the Klausipol- from the same sections, establishing four megaspore lenites schaubergeri–Reduviasporonites chalas- assemblage zones extending from the Pennsylva- tus–Syndesmorion stellatum assemblage, is again nian to the Lopingian. This palynostratigraphic dominated by the eponymous taxa, but also contains scheme is correlated with the assemblage zones of common non-taeniate bisaccate pollen, abundant Liu et al. (2008). The lowest biozone is Carbonifer- algal remains (see Foster & Afonin 2006) and com- ous, but the succeeding Bentzisporites margarita- mon Scutasporites cf. unicus. Other taxa include tus–Spencerisporites radiatus (MS) assemblage Lueckisporites virkkiae, Lunatisporites transver- zone is Kasimovian–Roadian in age based on fusu- sundatus, Lunatisporites pellucidus, Platysaccus linids and conodonts. The Biharisporites queenslandi, Alisporites splendens and Striatoa- grosstriletus (G) assemblage zone is of Wordian– bieites richteri. Among spores making their appear- Capitanian age, while the highest of the assemblage ance in this assemblage are Leptolepidites jonkeri, zones of Liu et al. (2011), the Biharisporites cf. fos- Limatulasporites fossulatus and Naumovaspora kettensis (F) assemblage zone, is believed to be of striata. The probable alga Reduviasporonites cha- Wuchiapingian age. Liu et al. (2011) commented lastus is also common. that several Carboniferous megaspores characteris- The upper assemblage is the Lundbladispora tic of Euramerica persist into the Guadalupian in foveota–Pechorosporites disertus–Otynisporites Shanxi, North China, indicating that a warm and eotriassicus assemblage. Of note here is the pre- humid climate prevailed in this area during the sence of the distinctive megaspore Otynisporites Pennsylvanian–Roadian, whereas the climate of eotriassicus. This taxon is considered a useful mar- Euramerica had already become arid by the end of ker for the base of the Triassic because it occurs in the Carboniferous. This warm and humid climate both marine and non-marine sections in Greenland, in northern China made it a refuge for some typi- Italy, Russia and Poland (Foster & Afonin 2005). cally Euramerican Carboniferous plants. The only study of the Permian–Triassic boun- Cathaysia dary sequence in the Cathaysian province and of the GSSP of the basal Triassic is that of Ouyang & Tha Cathaysia province is associated with the Utting (1990). The Changhsing Formation yields a South China and Indochina blocks of the eastern low-diversity assemblage dominated by acritarchs, Palaeotethys Ocean, including the present-day but contains rare Klausipollenites sp., scolecodonts regions of Yunnan, Shanxi, Meishan and Hunan. and Reduviasporonites chalastus. The lower part In eastern Yunnan, Ouyang (1982) described of the succeeding Griesbachian Chinglung For- a Lopingian Torispora gigantea–Patellisporites mation contains Lueckisporites virkkiae, Klausipol- meishanensis assemblage dominated by spores lenites schaubergeri, Alisporites cf. nuthallenis, similar to many that characterize the Pennsylvanian Protohaploxypinus spp., Weylandites sp., Cedri- of Euramerica (a so-called Carboniferous ‘relict pites spp. and Reduviasporonites chalastus, as well flora’), with a rather small representation of gym- as ‘taxa of Triassic aspect’ (e.g. Aratrisporites cf. nosperm pollen. The succeeding Yunnanospora yunnanensis). Species of Aratrisporites have previ- radiata–Gardenasporites assemblage from the ously been used to correlate the base of the Triassic Lopingian Changhsing Formation contains more (e.g. see Metcalfe et al. 2009), although strong evi- common taeniate bisaccate pollen and other gymno- dence for their reliability is lacking because species sperm pollen. of Aratrisporites are known from the Carboniferous Liu et al. (2008) described palynological of the Middle East and North Africa (e.g. Aratris- assemblages from the Cathaysian province of porites saharaensis; Loboziak et al. 1986). Never- Shanxi, North China, establishing four assemblage theless, Aratrisporites cf. yunnanensis first appears zones, in ascending order, the Torispora securis– 2.7 m above the base of the Triassic at Meishan Torispora laevigata (SL), the Torispora verru- according to Ouyang & Utting (1990), and thus cosa–Pachetisporites kaipingensis (VK), the Thy- this taxon might be taken as a local marker for the mospora thiessenii–Striatosporites heyleri (TH) base of the Triassic. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Gondwana (e.g. Palynozone I of Azcuy 1980) (Fig. 6) to be dominated by monosaccate pollen (e.g. Potonieis- Of the four phytogeographical provinces, Gond- porites) and zonate spores (e.g. Ancistrospora and wana underwent the greatest changes through the Lundbladispora), with taeniate bisaccate pollen Permian chiefly because the landmass of Gondwana being rare. Permian assemblages (e.g. Palynozone was comprehensively ice-bound at the beginning of III of Azcuy 1980) contain more taeniate bisaccate the period and then swiftly underwent deglaciation pollen, Vittatina and Cristatisporites. Superimposed in the Cisuralian. on these general trends are a large number of other The Late Palaeozoic glaciation of Gondwana more subtle changes that indicate variation between probably comprised three distinct episodes (Isbell South American basins and which have allowed a et al. 2003); the third and last glaciation was geo- number of separate basin-specific palynostratigra- graphically widespread and had continental ice phic schemes to evolve (e.g. see Vergel 1993; Arch- sheets spanning the Carboniferous–Permian boun- angelsky & Vergel 1996; Ce´sari & Gutie´rrez 2000; dary through the Moscovian to the Artinskian (Fiel- Playford & Dino 2000, 2002; di Pasquo 2003; Di ding et al. 2008). The extent of palaeobotanical Pasquo et al. 2003; Souza & Marques-Toigo 2003; isolation and the poverty of flora and fauna during Souza et al. 2003; Pe´rez Loinaze & Ce´sari 2004; this glacial period mean that palynological assem- Souza 2006; Balarino 2014) (Fig. 6). blages are particularly difficult to relate to the inter- In general, the biostratigraphy of the basins is national stages and to the Carboniferous–Permian difficult to relate to the international stages of the boundary, with the result that even now Carboni- Carboniferous and Permian (Archangelsky et al. ferous and Permian palynological assemblages are 1980) because of the scarcity of marine faunas: difficult to distinguish in Gondwana (Stephenson however, indirect comparisons with sequences con- 2008a). taining palynomorphs and cosmopolitan faunas and The sedimentary rocks deposited at this time floras (e.g. Marques-Toigo 1974; Cisterna et al. occur in cratonic basins that are now spread widely 2011) allow some tentative dates to be assigned. apart across Australia, India, Antarctica, southern Since 2007, the most marked progress has been and central Africa, and South America. The Permian made in integrating radiometric dates with paly- rocks of these cratonic basins contain commercial nological biozones, allowing limited – not always deposits of coal (e.g. in India, Australia, southern reconcilable – calibration of the latter with the inter- Africa and South America), and oil and gas (in national scale. Amongst the most important of these Australia, South America and the Middle East), studies are those of Ce´sari (2007), Guerra-Sommer and thus are amongst the most intensely studied et al. (2008), Ce´sari et al. (2011), Mori et al. (2012) Permian sequences in the world. The need in explo- and di Pasquo et al. (2015). ration to access stratigraphic sequences through In the first of the studies, Ce´sari (2007) noted borehole material (cuttings, core and sidewall core) radiometric dates in the San Rafael Basin in central has meant that palynology is also, by far, the most western Argentina and in the Parana´ Basin in south- important biostratigraphic tool used in locally corre- ern Brazil that suggested numerical ages for bio- lating sequences in the Gondwana Permian, mainly zones established by Ce´sari & Gutie´rrez (2000) for resource exploitation rather than for academic and Souza & Marques-Toigo (2003) in those basins, study. Thus, palynostratigraphic schemes tend to be respectively. Thus, the Lueckisporites–Weylandites locally focused, although attempts have been made Assemblage Biozone of Ce´sari & Gutie´rrez (2000) recently to correlate more widely across Gondwana, in the San Rafael Basin contains a horizon dated and from Gondwana to the international Permian at 266.3 + 0.8 Ma (Wordian), while the Lueckis- scale. Because of this local focus, this part of the porites virkkiae Interval Biozone of Souza & review concentrates on parts of Gondwana, and Marques-Toigo (2003) in the Parana´ Basin contains then attempts a synthesis at the end of the section. a dated horizon of 278.4 + 2.2 Ma (Kungurian). Guerra-Sommer et al. (2008) reported an age South America. Palynostratigraphy has progressed of 285.4 + 8.6 Ma (Artinskian) within the Parana´ most recently in four regions: the Tarija and Chaco- Basin Faxinal coal seam, which is assigned to the parana´ basins in northern Argentina, the Paganzo Hamiapollenites karooensis Sub-biozone of the Vit- and other basins in central western Argentina, the tatina costabilis Interval Biozone of Souza & Claromeco´ Basin in eastern Argentina, and the Marques-Toigo (2003). Mori et al. (2012) noted a Parana´ and Amazonas basins in Brazil. For reviews date of 281 + 3.4 Ma (Artinskian) for another hori- of earlier literature pertaining to South America zon within the Lueckisporites virkkiae Interval Bio- see Azcuy (1980) and Archangelsky et al. (1980). zone of the Parana´ Basin in the Candiota coal mine. Referring generally to South American basins, Ce´sari et al. (2011, fig. 6) produced a helpful Azcuy (1980) and Archangelsky et al. (1980) con- synthesis of the palynostratigraphy and radiometric sidered Pennsylvanian palynological assemblages dating of the Carboniferous and Cisuralian sequence Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

South N Argenna, E Argenna, Brazil, Paraná Brazil, Chaco-Paraná, Argenna NW Argenna System or America Bolivia Claromecó Basin Amazonas Playford &Dino (2002) Césari & Guérrez (2000) part Azcuy (1980) Tarija Basin, Di Basin, Balarino Souza & Marques- Basin Vergel (1993) Pasquo (2003) (2014) Toigo (2003) Playford & Dino (2000)

Upper ? T. toreutos –R. T. toreutos –R. Permian chalastus ? ? chalastus Assemblage Assemblage Zone Zone

Middle Striates Zone Permian L. virkkiae LW Palynozone IV Upper L. virkkiae C.confluens – Lower V. costabilis Zone V. vifera Permian Middle ? Assemblage V. costabilis Zone

Palynozone III FS V. costabilis Lower Cristasporites ? R. cephalata TB Upper Upper C. monoletus S. heyleri Carb.

Palynozone II Zone MR DM I. unicus

BC S. incrass. A. cristatus Lower Potonieisporites- Lundbladispora Palynozone I S. triangulus

Fig. 6. Correlation of South American palynostratigraphic schemes after Stephenson (2008a), Balarino (2014) and di Pasquo et al. (2015). Biozone codes: Ce´sari & Gutie´rrez (2000) DM, Raistrickia densa–Convolutispora muriornata; FS, Fusacolpites fusus–Vittatina subsaccata; LW, Lueckisporites–Weylandites; di Pasquo (2003) BC, Dictyotriletes bireticulatus–Cristatisporites chacoparanensis; MR, Granulatisporites micronodosus– Reticulatisporites reticulatus; TB, Marsupipollenites triradiatus–Lundbladispora braziliensis. across Argentina and Brazil correlating the San possible that Lueckisporites virkkiae has a diachro- Rafael and Parana´ basin biozones and using radio- nous first occurrence strongly reducing its value as metric dates to relate South American palynological a possible Euramerica–Gondwana ‘bridging taxon’. biozones to those of Namibia and Australia. Another possibility is that Lueckisporites virk- di Pasquo et al. (2015) gave radiometric dates kiae is being misidentified or that the conception from five volcanic ash beds within the Cisuralian of the taxon being used by taxonomists is too Copacabana Formation in central Bolivia (Tarija wide. The original concept of Lueckisporites virk- Basin). The five dates (cited as preliminary and pub- kiae Potonie´ & Klaus (1954) was of a diploxylonoid lished only in the non-peer reviewed Permian ICS bisaccate pollen grain with a wide separation of Newsletter Permophiles, 53, Supplement 1) are sacci (e.g. see Potonie´ & Klaus 1954, text-fig. 5, 298, 295.4–295.1 and 293 Ma (for two ash layers pl. 10, fig. 3; Klaus 1963, p. 300). Clarke (1965) approximately 25 m apart stratigraphically), and allowed more haploxylonoid specimens within 292.1–291.3 Ma. According to di Pasquo et al. Lueckisporites virkkiae in his emendation of the (2015), these dates suggest an Asselian age for the species, referring to these as ‘variant B’. di Pasquo ‘Vittatina costabilis assemblage’ and an Asselian– et al. (2015) did not illustrate the specimens that Sakmarian age for the ‘Lueckisporites virkkiae they attribute to Lueckisporites virkkiae, but the assemblage’ of di Pasquo et al. (2015, fig. 4). This specimen illustrated by Mori et al. (2012, fig. 3j) latter date is clearly inconsistent with other dates is strongly haploxylonoid and lacks evidence of a for the Lueckisporites virkkiae Interval Biozone of prominent cappa or exoexinal taeniae. It appears Souza & Marques-Toigo (2003) in the Parana´ closer to Corisaccites alutas. Haploxylonoid speci- Basin (e.g. see Ce´sari et al. 2011). mens of Lueckisporites virkkiae (using the concep- The accuracy of radiometric dates is important tion of Clarke 1965) are difficult to separate from for discussion of the stratigraphic occurrence of C. alutas, although Venkatachala & Kar (1966) Lueckisporites virkkiae, and inaccuracies inherent regarded C. alutas as ‘subsaccate’, and subsequent in radiometric dating may be the cause of apparent authors have described C. alutas as having poorly discrepancy in its first appearance, otherwise it is inflated ‘leathery’ sacci, the exoexine of which is Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON structurally indistinguishable from that of the cor- on extensive description, and particularly illustra- pus (see Stephenson 2008b). tion, of important taxa. Backhouse (1991) used To maintain the value of Lueckisporites virkkiae these data for a comparison and correlation with pal- as a biostratigraphical marker may mean rejecting ynological biozones established in the Western Aus- the emendation of Clarke (1965) and retaining the tralian Collie Basin (Fig. 7). This study remains one original concept of Lueckisporites virkkiae Potonie´ of the most rigorous intercontinental palynological & Klaus, 1954 as a diploxylonoid bisaccate pollen correlations yet completed for the Gondwana phyto- grain with a wide separation of sacci (see Stephen- geographical province, and, although correlation to son 2008b). It may also be valuable to start compar- the international stages is necessarily tentative, the ative studies between South American Gondwanan and Euramerican localities focusing on the genera Lueckisporites and Corisaccites. Karoo Basin Collie Basin Such uncertainties over correlation mean that, at System/Stage biozones, biozones, present, it is not possible to accurately correlate Anderson (1977) Backhouse (1991) South American biozones to the international scale Strat Biozone (Fig. 6). However, Vergel (1993) and Playford & Dino (2002) correlated the Carboniferous–Permian Protohaploxypinus boundary in the Chacoparana´ Basin to the boundary rugatus between the Potonieisporites–Lundbladispora Zone and the Cristatisporites Zone. Didecitriletes ?Ufimian Africa. Much palynological research in southern 4d ericianus African Permian sequences has been aimed at the correlation of coal seams (Falcon et al. 1984; Aitken Upper Ecca Dulhuntyispora 4c 1994; Millsteed 1999): however, more general granulata schemes have been proposed (Hart 1969; Falcon 1975; Anderson 1977; MacRae 1988). Microbaculispora villosa In the most detailed and extensive study covering 3d more than 35 boreholes, four surface localities and ?Kungurian more than 1000 samples, Anderson (1977) erected 3c Praecolpates seven biozones for the Permian. The lowest, Zone sinuosus 3b

1 (Anderson 1977, fig. 8), contains common Micro- Middle Ecca baculispora, monosaccate pollen (grouped mainly 3a Microbaculispora under Vestigisporites) and non-taeniate bisaccate trisina

pollen (grouped mainly under Pityosporites). Ander- Permian son (1977) considered Zone 1 to be Sakmarian 2d Striatopodocarpites in age. The succeeding Zone 2 (Sakmarian– 2c fusus Artinskian), divided into four subzones, is chiefly distinguished by its higher numbers of zonate 2b Pseudoreculaspora spores, although the highest subzone of Zone 2, Lower Ecca pseudoreculata and Zone 3 (Artinskian), are characterized by very abundant non-taeniate bisaccate pollen. Taeniate 2a bisaccate pollen become common only in the highest subzone of Anderson’s Zone 3.

Anderson’s Zone 4 (Artinskian–Wordian) con- Sakmarian Arnskian Converrucosisporites tains common taeniate bisaccate pollen with small confluens numbers of proximal taeniae which were referred to Lueckisporites by Anderson (1977), but would 1 be referred by modern taxonomists to Corisaccites, Guttulapollenites and, possibly, Taeniaesporites, as well as to Lueckisporites. Zone 5 (Capitanian) Dwyka is marked by common Vittatina. Amongst other Asselian Stage 2 ? changes, a marked drop in Vittatina marks the base of Zone 6. Zones 6 and 7 (Lopingian) are sim- Carboniferous ilar, but are distinguished by the differing propor- tions of their constituent species (Anderson 1977). The work of Anderson (1977) was not only Fig. 7. Correlation of Karoo and Collie Basin based on a large palynological database, but also biozones, after Backhouse (1991). Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY correlation of lithological units between Western Middle East (e.g. see Nader et al. 1993a; Stolle Australia and southern Africa is probably robust. 2007; Angiolini & Stephenson 2008; Jan et al. Other studies have also applied Australian paly- 2009; Stephenson et al. 2013; Stephenson & Powell nostratigraphic schemes in southern Africa (Mills- 2013, 2014). The chief unifying features of the teed 1993, 1999; Stephenson & McLean 1999). In standard palynozonation that occur throughout the east and central Africa, palynostratigraphic schemes Arabian Peninsula are, towards the base, the pres- were developed by Hart (1965a) and Utting (1978). ence of common monosaccate pollen, as well as Extensive taxonomic studies in that region were car- taxa such as Anapiculatisporites concinnus (charac- ried out by Bose & Kar (1966, 1976) and Bose & terizing Oman and Saudi Arabia Palynological Zone Maheshwari (1968). 1 (OSPZ1): ?Pennsylvanian) (Fig. 8), and above this Unlike South America, only a few ash layers the appearance of Microbaculispora and Horriditri- provided opportunities for direct dating of palyno- letes (characterizing OSPZ2: ?Pennsylvanian– logical biozones in Africa. However, Stephenson Asselian). (2009) recovered palynological assemblages of the The base of OSPZ3 is defined chiefly by Converrucosisporites confluens Oppel Zone from the abrupt increase in the small non-taeniate bisac- the Ganigobis Shale Member of Namibia, close to cate pollen Pteruchipollenites indarraensis from Ash layer IIb, which was radiometrically dated as approximately 10 to 50 or 60% of assemblages (Ste- 302.0 + 3.0 Ma (Pennsylvanian: Gzhelian or Kasi- phenson 2015). Other taxa that occur first consis- movian) (see Bangert 2000). Thus, the Converruco- tently in OSPZ3 are the taeniate bisaccate pollen sisporites confluens Oppel Zone may range earlier Striatopodocarpites cancellatus and S. fusus, and than previously thought and may influence age the taeniate ‘circumstriate’ pollen taxa, such as Cir- considerations of non-calibrated palynological bio- cumstriatites talchirensis and Striasulcites tectus. zones, such as those of Backhouse (1991) discussed Kingiacolpites subcircularis is common throughout earlier. Recent work on the palynology of South African coal seams (Go¨tz & Ruckwied 2014, Ruckwied et al. 2014) concentrated on using perceived cli- Chronostragraphy Palynostragraphy, Palynostragraphy, matic changes expressed in palynological sequences Stephenson Penney et al. 2003 et al. 2008 to correlate across South African basins. Barbolini & Bamford (2014) established two biozones in the Capitanian Mmamantswe coalfield (Lower Ecca) in Botswana. OSPZ6

Arabia, the Middle East, Pakistan and India. The broad character of Cisuralian assemblages in the Wordian southern parts of the Middle East, and Pakistan Guadalupian and India, are similar to those of Africa and South Roadian OSPZ5 America in that monosaccate pollen and simple tri- lete spores dominate the lowest glacigene horizons, Kungurian and these are succeeded by horizons with more diverse assemblages containing taeniate and non- Arnskian OSPZ4 taeniate bisaccate pollen, cycad pollen, and fern c b spores. Above this level in the late Cisuralian and OSPZ3 Guadalupian, the assemblages of Arabia, the Middle Sakmarian a 2141C East and Pakistan begin to diverge from those of the Cisuralian more southerly part of Gondwana probably because 2141B of more rapid climate warming due to the fact that Asselian/ Sakmarian OSPZ2 these areas were already of lower latitude and ? 2141A were rapidly moving north (e.g. Stephenson et al. 2165B 2008b). In Arabia, there are palynostratigraphic schemes 2165A for Oman (Besems & Schuurman 1987; Love 1994; Stephenson & Osterloff 2002; Penney et al. 2008; OSPZ1 2159B Stephenson et al. 2008a) and Saudi Arabia (Stump & van der Eem 1995); and these, as well as in-house Carb.? 2159A oil company schemes, have been synthesized by Stephenson et al. (2003) and Stephenson (2006) to a standard palynozonation for the entire Arabian Fig. 8. Palynostratigraphic schemes of the Peninsula, which also has application in the wider Arabian peninsula. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON this biozone, occasionally reaching 50% of assem- Talchir, Karharbari, Barakar, Barren Measures blages. OSPZ3 is believed to be Sakmarian in age and Raniganj) and did not span larger parts of the (Stephenson 2015). sequence (Truswell 1980). However, attempts have The chief distinguishing characteristic of OSPZ4 been made to synthesize data into palynostrati- (?Artinskian–Kungurian: Stephenson et al. 2003) graphic schemes for the entire Indian Permian is the common occurrence of Barakarites rotatus, (e.g. Tiwari & Tripathi 1992) and some limited cor- K. subcircularis and P. limpidus, although C. alutas, relations of Indian palynostratigraphy with inter- P. amplus, Plicatipollenites spp., Striatopodocar- national stages have been made (Tiwari 1996; pites spp., S. tectus, Strotersporites spp., Vesicas- Vijaya 1996). pora spp., Vittatina costabilis and Vittatina spp. The biozones of Tiwari & Tripathi (1992) are are also present. OSPZ5 (?Roadian–Wordian) is well defined, but are not dated using the interna- characterized by, amongst others, Distriatites insoli- tional scale. The earliest of these biozones, the Poto- tus, Playfordiaspora cancellosa and Thymospora nieisporites neglectus Biozone, from the lowest opaqua; and OSPZ6 (?Wordian–Capitanian: Ste- glacigene sediments of the Talchir Formation is phenson 2008b) is characterized by ?Florinites dominated by monosaccate pollen, but lacks taeni- balmei and Protohaploxypinus uttingii. A high-res- ate bisaccate pollen. The base of the succeeding Pli- olution palynozonation mainly aimed at correlation catipollenites gondwanensis Biozone is marked by in hydrocarbon basins of Oman can be correlated diversification and the appearance of taeniate bisac- with the OSPZ scheme (Penney et al. 2008) (Fig. 8) cate pollen, and the biozone above (Parasaccites Palynological data on higher parts of the korbaensis Biozone) in the upper Talchir Formation sequence to the Permian–Triassic transition is not by fern spores such as Microbaculispora tentula. available in the western part of the Middle East The base of the succeeding Crucisaccites monoletus because lithologies are not suitable for palynomorph Biozone of the Karhabari Formation is marked by preservation. However, to the east of the region the first appearances of the eponymous taxon, as in the Salt Range of Pakistan, this part of the Perm- well as taxa such as Marsupipollenites. The ages ian sequence is represented by rocks that contain of these biozones are considered to range from the both palynomorphs and well-preserved fauna. Asselian to Sakmarian (Vijaya 1996). The succeed- Balme (1970), in a pioneering taxonomic work on ing biozones of Tiwari & Tripathi (1992) are, in the Salt Range, defined many of the taxa of value order of deceasing age, the Scheuringipollenites in palynostratigraphy of Permian–Triassic Tethyan barakarensis, Faunipollenites varius, Densipollen- and Gondwanan palynostratigraphy, although ites indicus, Gondisporites raniganjensis and Densi- Balme (1970) did not define biozones himself. More pollenites magnicorpus biozones. Tiwari (1996) recently, Hermann et al. (2012) defined two latest attempted to date ‘palynoevents’ through the Indian Permian assemblages in the Salt Range (‘Chhidru Permian: for example, the decline of monosaccates 1’ and ‘Chhidru 2’) present in the uppermost Chhi- (Palynoevent 3) was considered to be late Artin- dru Formation (Changhsingian). Chhidru 1 (the skian, and the first appearance of Densipollenites Protohaploxypinus spp.–Weylandites spp. Associa- (Palynoevent 4) as Kungurian–Ufimian – although tion) is dominated by gymnosperm pollen, including these were not related to the biozones of Tiwari & common Klausipollenites spp. (including K. schau- Tripathi (1992). bergeri) and Protohaploxypinus spp., (including Despite the fact that there has been some con- P. limpidus), as well as Weylandites lucifer and solidation of schemes in India, local palynostrati- Alisporites spp. Spores are very rare. Chhidru 2 graphic schemes continue to be developed in the (the Kraeuselisporites wargalensis–Protohaploxy- cratonic basins. In Andhra Pradesh, Aggarwal & pinus spp. Association) is characterized by abundant Jha (2013) defined eight Permian palynological bio- cavate trilete spores such as Kraeuselisporites war- zones for the Godavari Graben, linking these with galensis and Kraeuselisporites spp. According to climate changes, although not to the international Hermann et al. (2012), the range of K. wargalensis Permian scale. Jha et al. (2014), also working in is restricted to Chhidru 2. The top of Chhidru 2 the Andhra Pradesh Godavari Graben, identified two is characterized by the last occurrences of Klausi- assemblages (Palynoassemblage-I and -II) dated as pollenites schaubergeri and Lueckisporites spp. Lopingian. In northern India in Kashmir, Tewari (including Lueckisporites virkkiae). et al. (2015) identified Permian–Triassic assem- In India, much of the taxonomic work in the blages broadly correlated to the Densipollenites 1960s was directed at correlation between coal magnicorpus and Klausipollenites decipiens bio- seams, and not at correlation with other Gondwana zones of peninsular India (see Tiwari & Tripathi sequences or international stages of the Permian. 1992). Major taxonomic studies were confined mainly to one or other of the ‘lithostratigraphic stages’ of Australia. Australia has the best-documented the peninsular Indian Permian (in ascending order: Cisuralian sequences of Gondwana and, owing to Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY the presence of rare marine intervals, some cali- the Canning Basin, Western Australia (Foster & bration of palynological biozones with interna- Archbold 2001), dated as ‘post-Guadalupian’ by tional stages has been possible using marine Glenister et al. (1990) and ‘Capitanian–Dzhulfian’ macrofaunas (e.g. Leonova 1998; Archbold 1999). by Leonova (1998). Such tie points allow spot Later palynological biozones in the Guadalupian dates to be applied to the otherwise well-developed and Lopingian are more difficult to date due to palynostratigraphy. the lack of marine fauna (in coal and red beds), A feature of palynostratigraphy in the Australian although there are tie points (Foster & Archbold Permian is significant endemism, with the result that 2001): for example, the record of a single specimen separate palynozonations developed in western and of the ammonoid Cyclolobus persulcatus, from the eastern Australian basins in the earliest period of Cherrabum Member of the Hardman Formation, in palynological investigation (Fig. 9). In the west, eight assemblage biozones (units I–VIII), ranging in age from Pennsylvanian to Lopingian, were rec- ognized; while, in the east, five assemblage bio- Eastern Western System/ Australia Australia zones (stages 1–5) encompass the same interval Stage (Evans 1969; Kemp et al. 1977; Foster 1979; Trus- Ages aer Backhouse well 1980; Price 1983, 1997). The precise relation- (1991) Kemp et Price (1983, Balme Backhouse ship between these schemes, and the ages of the al. (1977) (1980) (1991) 1997) biozones, remains speculative (see Jones & Trus- well 1992) (Fig. 9), but there are broad similarities Protohaploxypinus reculatus across the continent (see Balme 1980, text-fig. 4). In the lower parts of the glacigene sequence (Stage 1), radially and bilaterally symmetrical APP5 Unit VII D. parvithola monosaccate pollen is dominant, and taeniate and

Stage 5 P. rugatus D. ericianus Main trends ?Ufimian APP4 D. granulata Geochronology and Chronostragraphy type spores M. villosa pollen bisaccate Taeniate bisaccate Cycadopites Monosaccate Cheilocardioid Stage 4 Nontaeniate ?Kungurian APP3 Unit VI Kungurian P. sinuosus Unit V

M. trisina Arnskian

Unit IV Arnskian Stage 3 APP2 S. fusus Sakmarian Unit III P. pseudo. Lower Permian (Cisuralian) Permian Lower Asselian C. confl. Sakmarian Gzhelian Permian

Stage 2 APP1 Unit II ? Asselian Kazimovian Very rare Moscovian

?Gzhelian Rare Stage 1 Pennsylvanian pars Bashkirian Carboniferous Common ?Kazimovian Fig. 10. Main trends in palynomorph types across the Fig. 9. Correlation of eastern and western Australian Gondwana phytogeographical province from the palynostratigraphic schemes, after Kemp et al. (1977), Pennsylvanian to the Cisuralian, after Stephenson Foster (1979), Balme (1980), Backhouse (1991) and (2008a). Correlation of events to the international scale Price (1983, 1997). is approximate. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY non-taeniate bisaccate pollen is subordinate. In Lopingian palynological assemblages from Oman Stage 2 above, small triangular fern spores (cheilo- and Saudi Arabia are very different to those of cardioid spores) and Cycadopites become common; Australia. Similarly, the Guadalupian palynology while, around the bases of Stage 3, taeniate and non- of South America differs considerably to that of taeniate bisaccate pollen become common. In Stage Australia. These palynological differences, gener- 4, a diversity of pollen and spores develops, includ- ated in the late Cisuralian following deglaciation ing distinctive colpate pollen such as Praecolpatites of the Gondwana continent, are probably due to sinuosus; while Stage 5 is partly characterized by the northwards drift of Gondwana through the Perm- the very distinctive spore genus Dulhuntyispora, ian, and the different order in which the various which is distinguished by expanded blister-like parts of Gondwana moved through latitudinal belts. exoexine in each of the inter-radial areas. For the Cisuralian, the main palynological events that can be tracked with reasonable certainty Antarctica. Early work in Antarctica was carried out through all basins of Gondwana appear to be the by Balme & Playford (1967) and Kemp (1973). appearance and diversification patterns of: (1) mono- Kyle (1977) erected two informal palynological bio- saccate pollen; (2) cheilocardioid spores (such as zones from study of a wide range of Antarctica Microbaculispora); (3) Cycadopites pollen; and localities. Her Parasaccites Zone is dominated by (4) taeniate and non-taeniate bisaccate pollen. The monosaccate pollen, along with rare bisaccate taeni- precise ages of these events in relation to the inter- ate pollen, and, locally, high frequencies of Micro- national scale are not known, and recent data from baculispora tentula and Cycadopites cymbatus. radiometric dating are still being collected and According to Truswell (1980), this biozone com- assimilated (e.g. Stephenson 2009; Ce´sari et al. pares closely with the upper part of Stage 2 in Aus- 2011; di Pasquo et al. 2015) (Fig. 10). tralia. The succeeding Protohaploxypinus Zone of Having said this, a few high-resolution corre- Kyle (1977) is distinguished by abundant bisaccate lations within Gondwana that involve younger pollen, including many taeniate taxa. In gross com- Permian rocks have been possible based on close position and species content, this biozone resembles taxonomic comparison and accurate stratigraphic Stage 4 assemblages from eastern Australia. Work information: for example, the correlation of the following that of Kyle (1977) has tended to apply Collie Basin, Western Australia by Backhouse Australian palynostratigraphy to Antarctica, rather (1991) with the South African biozones of Anderson than develop more sophisticated local palynostrati- (1977) (see Fig. 7). Dino & Playford (2002) and graphic schemes (e.g. Farabee et al. 1990; Larrsson Stephenson (2008a) identified taxa that are com- et al. 1990; Lindstro¨m 1995a, b; Lindstro¨m& mon to South America and Australia in the Carbon- McLoughlin 2007). iferous–Permian. Converrucosisporites confluens occurs across most Gondwana sequences: Australia Synthesis of Gondwana palynostratigraphy. It is (Foster & Waterhouse 1988), India (Srivastava & clear that there are quantitative and qualitative sim- Bhattacharyya 1996), Argentina (Archangelsky & ilarities in the late Pennsylvanian and Cisuralian that Gamerro 1979), Uruguay (Beri & Goso 1996), Ant- allow correlation across Gondwana with reasonable arctica (Lindstro¨m 1995b), and Oman and Saudi accuracy; after this period, however, palynological Arabia (Stephenson et al. 2003). Ahrensisporites characteristics of the various parts of Gondwana cristatus is considered to have a near-synchronous begin to diverge. For example, Guadalupian and first appearance in the early Pennsylvanian, while

Fig. 11. Photomicrographs of taxa of possible biostratigraphical value between and within phytogeographical provinces of the Permian. (1) Lueckisporites virkkiae, F46, V-619, 2187; from the basal Khuff clastics, Saudi Arabia; width of specimen approximately 50 mm. (2) Lueckisporites virkkiae, E42, V-619, 2187; from the basal Khuff clastics, Saudi Arabia; width of specimen approximately 50 mm. (3) Converrucosisporites confluens, V42, MM-151, 3167, distal focus; from the Al Khlata Formation, Oman; width of specimen approximately 40 mm. (4) Converrucosisporites confluens, V42, MM-151, 3167, proximal focus; from the Al Khlata Formation, Oman; width of specimen approximately 40 mm. (5) Weylandites sp. slide: 1563, EF: V30.3; from the Lower Karoo sequence, Kalahari Karoo Basin, Botswana; courtesy Alain Le He´risse´; specimen featured in Modie & Le He´risse´ (2009); width of specimen approximately 35 mm. (6) Vittatina sp., M27, RA-2, 4256; from the Al Khlata Formation, Oman; width of specimen approximately 35 mm. (7) Vittatina sp., M27, TL-16, 982, from the Al Khlata Formation, Oman; width of specimen approximately 35 mm; (8) Scutasporites unicus , collection number GBA 2010/013/0027 (holotype); single grain preparation, slide No. 404, England-finder K34r; Gro¨den Formation ¼ Gro¨dner Sandstein (Arenaria di Val Gardena); courtesy of Prof. Hans Egger; specimen featured in Draxler (2010); width of specimen approximately 50 mm. (9) Otynisporites eotriassicus; Otyn IG 1 borehole, depth 817.5 m; Balic Formation (Lower Buntsandstein), Induan; courtesy of Dr Teresa Marcinkiewicz; specimen featured in Marcinkiewicz et al. (2014); width of specimen approximately 400 mm. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON the last appearances of Psomospora detecta, Spe- precision of taxonomic classification in different laeotriletes ybertii and Rattiganispora apiculata areas such that the utility of potentially useful were also considered to be possibly coeval in both widely correlateable palynological taxa are reduced continents (Dino & Playford 2002). (e.g. Lueckisporites virkkiae). The benefits of a better integrated general paly- nostratigraphy are very great scientifically because Discussion and conclusions there are numerous events of global scientific inter- est in the Permian: for example, the timing and order The purpose of palynostratigraphy, like biostratigra- of deglaciation events across the continent of Gond- phy generally, is to provide correlation. Correlation wana that can, arguably, only be established through helps palynologists, stratigraphers and geologists to a Gondwana-wide palynostratigraphic scheme (see relate sedimentary rocks deposited in one place to Stephenson et al. 2007), large-scale patterns of flo- those in another, and to relate geological resources ral migration across continents, and the detailed or events to each other and to other important geo- characteristics and timing of mass extinction events logical or scientific phenomena. within the Permian and at the Permian–Triassic In the Permian, the most important reasons for boundary. correlation related to resource extraction are to Perhaps the most important generalization that describe coal and hydrocarbon resources, to place can be made about the palynostratigraphy of the them into a regional framework, and to help to Permian is that phytogeographical provinciality is understand how to find more coal and hydrocarbons. strong throughout the Permian and particularly Probably, the bulk of work that has been done on from the Guadalupian onwards, as predicted by Permian rocks in palynostratigraphy has been in palaeobotanical studies. This makes correlation this area. There have been several tens of palynos- between regional palynostratigraphic schemes diffi- tratigraphy papers published on coal measures strat- cult. These differences appear to be more than could igraphy in India, South America, Australia and be accounted for by differing taxonomic concepts, southern Africa, with the result that coal measures although these may contribute to the extent of the sequences (e.g. Foster 1979), and even individual apparent differences between phytogeographical coal seams, can be correlated (e.g. Aitken 1994). provinces. For these reasons, it is unlikely that a sin- Perhaps the work that has led to the highest-resolu- gle comprehensive palynostratigraphic scheme for tion palynostratigraphy (in terms of being able to the Permian globally will ever be developed, or at recognize and distinguish the smallest divisions of least not one that contains inherent precision that stratigraphy and time) has been for the hydro- will be valuable for resolving Earth events. This carbons industry, particularly in the Middle East paper does not, therefore, suggest such a scheme, where very large accumulations of oil and gas nor even suggest unified schemes, for the phytogeo- occur in the Permian. These schemes are capable graphical provinces of the Permian. of biostratigraphically characterizing rock units of What are likely to be possible are high-resolution only a few metres in thickness across oil and gas palynostratigraphic schemes for regions (which sub- fields or even basins (e.g. Stephenson & Osterloff stantially already exist) that can be linked by tie 2002; Penney et al. 2008; Stephenson et al. 2008a). points. These tie points will be provided by precise In Australia also, the work of Price (1983, 1997) has assemblage-level quantitative taxonomic compari- been widely used for Permian onshore oil, gas, son or by the use of single well-characterized paly- unconventional hydrocarbons and coal exploration. nological taxa (‘bridging taxa’) that occur across Although these palynostratigraphic schemes Permian phytogeographical provinces. related to resource extraction have been very suc- Perhaps the most important of these taxa for cessful, their main shortcoming has been a lack of the Permian include (Fig. 11): (1) Scutasporites correlation with schemes outside the basins, coal- spp., Vittatina spp., Weylandites spp. and Luec- fields and hydrocarbon fields that they serve, and kisporites virkkiae (for correlation between the chiefly a lack of correlation with the international Euramerican and Angaran provinces); and (2) Vitta- Permian scale. In the past, a lack of fundamental tina spp., Weylandites spp. and Lueckisporites virk- stratigraphic standards (e.g. GSSPs) for the Permian kiae (for correlation between the Euramerican hampered this kind of correlation, but these now and Gondwana provinces). Correlation between substantially exist (see Henderson et al. 2012) and the Cathaysian and other phytogeographical prov- local and regional palynostratigraphic schemes inces remains a problem because few ‘bridging should be fitted into the wider Permian scale. taxa’ have been identified. This may represent a In general, also, standards of recording of paly- lack of effort in palynological studies in linking nological data and accessibility of data are higher, Cathaysian and other phytogeographical provinces, allowing easier comparison between palynostrati- and the isolation of much Chinese palynology in graphic schemes. Some problems still exist in the that it is published only in Chinese. 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PERMIAN PALYNOSTRATIGRAPHY

None of the Permian GSSPs involve palynolog- Hamiapollenites tractiferinus (Samoilovich) Jansonius, ical definitions, which may be problematic given 1962 the importance of palynology in correlation in the Horriditriletes ramosus (Balme & Hennelly) Bharadwaj & commercial and academic worlds. However, there Salujah, 1964 appear to be taxa that occur at GSSPs or well-dated Horriditriletes tereteangulatus (Balme & Hennelly) Back- boundary sections that could be used to correlate house, 1991 those boundaries. For example, Aratrisporites spp. Kingiacolpites subcircularis Tiwari & Moiz, 1971 and Otynisporites eotriassicus may be useful to Klausipollenites schaubergeri (Potonie´ & Klaus) Janso- correlate the Permian–Triassic boundary into non- nius, 1962 marine sections or sections without radiometric Kraeuselisporites wargalensis Balme, 1970 dates. Converrucosisporites confluens may be use- Leiotriletes virkkii Tiwari, 1965 ful in correlating the Carboniferous–Permian boun- Leptolepidites jonkeri (Jansonius) Yaroshenko & Golu- dary (Fig. 11). beva, 1991 Limatulasporites fossulatus (Balme) Helby & Foster in The Director of the British Geological Survey (NERC) is Foster, 1979 thanked for permission to publish this paper. Dr Mercedes Limitisporites monstruosus (Luber & Valtz) Hart, 1965b di Pasquo is acknowledged for providing access to valu- Lueckisporites virkkiae (Potonie´ & Klaus) emend. Clarke, able reference materials. Prof. Clinton Foster and an anon- 1965 ymous reviewer provided constructive comments. Dr Lunatisporites pellucidus (Goubin) Helby ex de Jersey, Alain Le He´risse´ (University of Brest), Prof. Hans Egger (Geological Survey of Austria) and Dr Teresa Marcinkie- 1972 wicz are thanked for providing images of specimens in Lunatisporites transversundatus (Jansonius) Fischer, Figure 11. 1979 Lycospora pusilla (Ibrahim) Somers, 1972 Microbaculispora tentula Tiwari, 1965 Naumovaspora striata Jansonius, 1962 Appendix A Otynisporites eotriassicus Fuglewicz, 1977 Species names and authors Platysaccus queenslandi de Jersey, 1962 Playfordiaspora cancellosa (Playford & Dettmann) Ahrensisporites cristatus Playford & Powis, 1979 Maheshwari & Banerji, 1975 Alisporites cf. nuthallenis Clarke, 1965 Praecolpatites sinuosus (Balme & Hennelly 1956) Alisporites plicatus Jizba, 1962 Bharadwaj & Srivastava, 1969 Alisporites splendens (Jizba) Foster, 1979 Protohaploxypinus amplus (Balme & Hennelly) Hart, Anapiculatisporites concinnus Playford, 1962 1964 Aratrisporites cf. yunnanensis Ouyang & Li, 1980 Protohaploxypinus limpidus (Balme & Hennelly) Balme & Aratrisporites saharaensis Loboziak et al., 1986 Playford, 1967 Barakarites rotatus (Balme & Hennelly) Bharadwaj & Protohaploxypinus perfectus (Naumova ex Kara-Murza) Tiwari, 1964 Samoilovich, 1953 Brevitriletes cornutus (Balme & Hennelly) Backhouse, Protohaploxypinus uttingii Stephenson & Filatoff, 2000 1991 Pseudoulatispora pseudoreticulata (Balme & Hennelly) Cedripites priscus Balme, 1970 Bharadwaj & Srivastava, 1969 Circumstriatites talchirensis Lele & Makada, 1972 Psomospora detecta Playford & Helby, 1968 Cladaitina kolodae Utting, 1994 Pteruchipollenites indarraensis (Segroves) Foster, 1979 Converrucosisporites confluens (Archangelsky & Game- Rattiganispora apiculata Playford & Helby emend. rro) Playford & Dino, 2002 Playford, 1986 Cordaitina uralensis (Luber) Samoilovich, 1953 Reduviasporonites chalastus (Foster) Elsik, 1999 Corisaccites alutas Venkatachala & Kar, 1966 Scutasporites nanuki Utting, 1994 Crassispora kosankei (Potonie´ & Kremp) Bharadwaj, Scutasporites unicus Klaus, 1963 1957 Scutasporites xinjiangensis (Hou & Wang) Ouyang et al. Crinalites sabinensis Utting, 1994 2003 Crucisaccites ornatus (Samoilovich) Dibner, 1971 Spelaeotriletes triangulus Neves & Owens, 1966 Cycadopites cymbatus (Balme & Hennelly) Segroves, 1970 Spelaeotriletes ybertii (Marques-Toigo) Playford & Distriatites insolitus Bharadwaj & Salujah, 1964 Powis, 1979 Endosporites ornatus Wilson & Coe, 1940 Striasulcites tectus Venkatachala & Kar, 1968 Falcisporites zapfei (Potonie´ & Klaus) Leschik, 1956 Striatoabieites multistriatus (Balme & Hennelly) Hart, Florinites luberae Samoilovich, 1953 1964 Florinites? balmei Stephenson & Filatoff, 2000 Striatoabieites richteri (Klaus) Hart, 1964 Hamiapollenites bullaeformis (Samoilovich) Jansonius, Striatopodocarpites cancellatus (Balme & Hennelly) 1962 Hart, 1964 Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Striatopodocarpites fusus (Balme & Hennelly) Potonie´, Azcuy, C., Beri,A´ . et al. 2007. Bioestratigrafı´a del 1958 Paleozoico Superior de Ame´rica del Sur: Primera Sverdrupollenites agluatus Utting, 1994 Etapa de Trabajo Hacia una Nueva Propuesta Cronoes- Thymospora opaqua Singh, 1964 tratigra´fica. In: Comite´ Organizador de la XI Reunia˜o de Paleobotaˆnicos Paleobotaˆnicos y por el Comite´ Vittatina costabilis Wilson, 1962 Organizador del XIII Simposio Argentino de Paleobo- Vittatina saccata (Hart) Jansonius, (1962) ta´nica y Palinologı´a (Bahı´a Blanca), 2006, Argentina. Weylandites cincinnatus (Luber ex. Varyukhina) Utting, Asociacio´n Geolo´gica Argentina Serie D: Special Pub- 1994 lication, 11. Weylandites lucifer (Bharadwaj & Salujah) Foster, 1975 Backhouse, J. 1991. Permian palynostratigraphy of the Weylandites striatus (Luber) Utting, 1994 Collie Basin, Western Australia. Review of Palaeobo- tany and Palynology, 67, 237–314. Balarino, M.L. 2014. Permian palynostratigraphy of the Claromeco´ Basin, Argentina. Alcheringa, 38, 1–21. References Balme, B.E. 1970. Palynology of Permian and Triassic strata in the Salt Range and Surghar Range, West Paki- Aggarwal,N.&Jha, N. 2013. Permian palynostrati- stan. In: Kummel,B.&Teichert, C. (eds) Strati- graphy and palaeoclimate of Lingala-Koyagudem graphic Boundary Problems: Permian and Triassic Coalbelt, Godavari Graben, Andhra Pradesh, India. of West Pakistan. University Press of Kansas, Depart- Journal of Asian Earth Sciences, 64, 38–57. ment of Geology Special Publications, 4, 306–453. Aitken, G.A. 1994. Permian palynomorphs from the No. Balme, B.E. 1980. Palynology and the Carboniferous– 5 seam, Ecca Group, Witbank/Highvelt Coalfields, Permian boundary in Australia and other Gondwana South Africa. Palaeontologia Africana, 31, 97–109. countries. Palynology, 4, 43–56. Anderson, J.M. 1977. The biostratigraphy of the Permian Balme, B.E. & Playford, G. 1967. Late Permian plant and Triassic. Part 3. A review of Gondwana Permian microfossils from the Prince Charles Mountains, Ant- palynology with particular reference to the northern arctica. Revue de Micropale´ontologie, 10, 179–192. Karoo Basin of South Africa. Memoirs of the Botanical Bangert, B. 2000. Tephrostratigraphy, petrography, Survey of South Africa, 41, 1–133. geochemistry, age and fossil record of the Ganigobis Angiolini,L.&Stephenson, M.H. 2008. Early Perm- Shale Member and associated glaciomarine deposits ian brachiopods and palynomorphs from the Dorud of the Dwyka Group, Late Carboniferous, southern Formation, Alborz Mountains, north Iran: evidence Africa. Unpublished PhD thesis, Bayerischen Julius- for their palaeobiogeographic affinities. Fossils and Maximilians-Universita¨tWu¨rzburg, Germany. Strata, 54, 117–132. Barbolini,N.&Bamford, M.K. 2014. Palynology of an Archangelsky,S.&Gamerro, J.C. 1979. Palinologı´a Early Permian coal seam from the Karoo Supergroup del Paleozoico Superior en el subsuelo de la cuenca of Botswana. Journal of African Earth Sciences, 100, Chacoparanense, Repu´blica Argentina 1. Estudio siste- 136–144. ma´tico de los palinomorfos de tres perforaciones de la Bercovici, A., Diez, J.B. et al. 2009. A palaeoenviron- Provincia de Co´rdoba. Revista Espan˜ola de Micropa- mental analysis of Permian sediments in Minorca (Bal- leontologı´a, 11, 417–478. earic Islands, Spain) with new palynological and Archangelsky,S.&Vergel, M. 1996. Paleontologı´a, megafloral data. Review of Palaeobotany and Palynol- bioestratigrafı´a y paleoecologı´a. In: El sistema Pe´r- ogy, 158, 14–28. mico en la Republica Argentina y en la Republica Ori- Beri,A.&Goso, C.A. 1996. Ana´lisis palinolo´gico y estra- ental del Uruguay. Academia Nacional de Ciencias, tigra´fico de la Fm. San Gregorio (Pe´rmico Inferior) en 1996, Co´rdoba, Argentina, 40–44. el a´rea de los Cerros Guazunambi, Cerro Largo, Uru- Archangelsky,S.&Wagner, R.H. 1983. Glosspteris guay. Revista Espan˜ola de Micropaleontologı´a, 28, anatolica sp. nov. from uppermost Permian strata in 67–79. south east Turkey. Bulletin of the British Museum (Nat- Besems, R.E. & Schuurman, W.M.L. 1987. Palynostra- ural History), Geology, 37, 81–91. tigraphy of Late Paleozoic glacial deposits of the Archangelsky, S., Azcuy, C.L., Pinto, I.D., Gonza´- Arabian Peninsula with special reference to Oman. lez, C.R., Marques-Toigo, M., Ro¨sler,O.&Wag- Palynology, 11, 37–53. ner, R.H. 1980. The Carboniferous and Early Bharadwaj, D.C. 1957. The spore flora of the Velener Permian of the South American Gondwana area: a Schichten (Lower Westphalian D) in the Ruhr Coal summary of the biostratigraphic information. Actas Measures. Palaeontographica B, 102, 110–138. del II Congreso Argentino de Paleontologı´a y Bioes- Bharadwaj, D.C. 1969. Lower Gondwana Formations. tratigrafı´a y I Congreso Latinoamericano de Paleonto- In: 6th Congre`s International de Stratigraphie et de logı´a 1978, 4, 257–269. Ge´ologie du Carbonife`re. 1967, Sheffield. Compte Archbold, N.W. 1999. Permian Gondwana correlations: Rendu, 1, 255–274. the significance of the western Australian marine Perm- Bharadwaj, D.C. & Salujah, S.K. 1964. Sporological ian. Journal of African Earth Sciences, 29, 63–75. Study of Seam VIII in Raniganj Coalfield, Bihar, Azcuy, C.L. 1980. A review of the early Gondwana paly- India. Part I: Description of the sporae dispersae. nology of Argentina and South America. In: Proceed- Palaeobotanist, 12, 181–215. ings of the IV International Palynological Conference, Bharadwaj, D.C. & Srivastava, S.C. 1969. Some new 1976, Lucknow, India, Volume 2. Birbal Sahni Insti- miospores from Barakar Stage, Lower Gondwana, tute, Lucknow, India, 175–185. India. Palaeobotanist, 17, 220–229. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Bharadwaj, D.C. & Srivastava, S.C. 1977. Gondwana Davydov, V.I., Glenister, B.F., Spinosa, C., Ritter, palynology – a historical review. In: Venkatachala, S.C., Chernykh, V.V., Wardlaw, B.R. & Snyder, B.S. & Sastri, V.V. (eds) IV Colloquim on Indian W.S. 1998. Proposal of Aidaralash creek as Global Micropalaeontology and Stratigraphy 1974–1975. Stratotype Section and Point (GSSP) for base of the Proceedings of the Instititute of Petroleum Explora- Permian System. Episodes, 21, 11–18. tion, Oil and Natural Gas Commission, Dehradun. De Jersey, N.J. 1962. Triassic spores and pollen from the Institute of Petroleum Exploration, Oil & Natural Ipswich Coalfield. Geological Survey of Queensland Gas Commission, Dehradun, 150–156. Publications, 307, 1–18. Bharadwaj, D.C. & Tiwari, R.S. 1964. On two monosac- De Jersey, N.J. 1972. Triassic miospores from the Esk cate genera from Barakar Stage of India. Palaeobotan- beds. Geological Survey of Queensland Publications, ist, 12, 139–146. 357, 1–40. Bose, M.N. & Kar, R.K. 1966. Palaeozoic sporae disper- Dibner, A.F. 1971. Cordaitales pollen of Angaraland. sae from Zaire (Congo) I. Kindu\Kalima and Walikale Uchenye Zapiski, Nauchno Issledovatelski Instituti regions. Annales Musee Royal de L’Afrique Centrale, Geologii Arctiki, Paleontologia i Bioestratigrafia, 32, Se´rie 8, 53, 1–168. 5–66 (In Russian with English summary). Bose, M.N. & Kar, R.K. 1976. Mezozoic sporae dispersae Dino,R.&Playford, G. 2002. Miospores common to from Zaire I. Haute-Lueki Series. Annales Musee South American and Australian Carboniferous Royal de L’Afrique Centrale, Se´rie 8, 78, 1–40. sequences: stratigraphic and phytogeographic implica- Bose, M.N. & Maheshwari, H.K. 1968. Paleozoic sporae tions. In: Hills, L.V., Henderson, C.M. & Bamber, dispersae from Congo – VII. Coal measures near Lake E.W. (eds) Carboniferous and Permian of the World. Tanganyika, south of Albertville. Annales Musee Canadian Society of Petroleum Geologists, Memoirs, Royal de L’Afrique Centrale, Se´rie 8, 60, 1–116. 19, 336–359. Brugman, W.A. & Visscher, H. 1988. Permian and Tri- di Pasquo, M. 2003. Avances sobre palinologı´a, bioestra- assic palynostratigraphy of northeast Libya. In: El tigrafı´a y correlacio´n de los Grupos Macharetı´ y Man- Arnauti, A., Owens,B.&Thuso, B. (eds) diyutı´ Neopaleozoico de la Cuenca Tarija, Provincia de Subsurface Palynostratigraphy of Northeast Libya. Salta, Argentina. Ameghiniana, 40, 3–32. Garyounis University Publications, Benghazi, Libya, di Pasquo, M., Azcuy, C.L. & Souza, P.A. 2003. Palino- 157–169. logı´a del Carbonı´fero Superior del Subgrupo Itarare´ en Brugman, W.A., Loboziak,S.&Visscher, H. 1988. The Itaporanga, Cuenca Parana´, Estado de Sa˜o Paulo, Bra- problem of the Carboniferous-Permian boundary in sil. Parte 2: sistema´tica de polen y significado paleoam- north east Libya from a palynological point of view. biental y estratigra´fico. Ameghiniana, 40, 277–296. In: El Arnauti, A., Owens,B.&Thuso, B. (eds) di Pasquo, M., Grader, G.W., Isaacson, P., Souza, Subsurface Palynostratigraphy of Northeast Libya. P.A., Iannuzzi,R.&Dı´az- Martı´nez, E. 2015. Garyounis University Publications, Benghazi, Libya, Global biostratigraphic comparison and correlation of 151–155. an early Cisuralian palynoflora from Bolivia. Histori- Ce´sari, S.N. 2007. Palynological biozones and radio- cal Biology, 27, 868–897. metric data at the Carboniferous–Permian boundary Doubinger, J. 1974. E´ tudes palynologiques dans L’Autu- in western Gondwana. Gondwana Research, 11, nien. Review of Palaeobotany and Palynology, 17, 529–536. 21–38. Ce´sari, S.N. & Gutie´rrez, P.R. 2000. Palynostratigraphy Doubinger, J., Odin,B.&Conrad, G. 1987. Les associ- of Upper Palaeozoic sequences in central-western ations sporopolliniques du Permien continental du Argentina. Palynology, 24, 113–146. bassin de Lode`ve (He´rault, France): caracterisation Ce´sari, S.N., Limarino, C.O. & Gulbransen, E.L. 2011. de L’Autunien supe´rieur, du ‘Saxonien’ et du Thurin- An Upper Paleozoic biochronostratigraphic scheme gien. Annales de la Socie´te´ Ge´ologique du Nord, for the western margin of Gondwana. Earth-Science 106, 103–109. Reviews, 106, 149–160. Draxler, I. 2010. Upper Permian spore holotypes of Cisterna, G.S., Sterren, A.F. & Gutie´rrez, P. 2011. W. Klaus from the Southern Alps (Dolomites, Italy) The Carboniferous-Permian boundary in the central in the collections of the Geological Survey of Austria. western Argentinean basins: paleontological evi- Jahrbuch der Geologischen Bundesanstalt, 150, dences. Andean Geology, 38, 349–370. 85–99. Clapham, W.B. 1970. Permian miospores from the Flow- Dunn, M.T. 2001. Palynology of the Carboniferous – erpot Formation of Western Oklahoma. Micropaleon- Permian boundary stratotype, Aidaralash Creek, tology, 16, 15–36. Kazakhstan. Review of Palaeobotany and Palynology, Clarke, R.F.A. 1965. British Permian saccate and mono- 116, 175–194. sulcate miospores. Palaeontology, 8, 322–354. Edwards, R.A., Warrington, G., Scrivener, S.C., Clayton, G., Coquel, R., Doubinger, J., Gueinn, K.J., Jones, N.S., Haslam, H.W. & Ault, L. 1997. The Loboziak, S., Owens,B.&Streel, M. 1977. Car- Exeter Group, south Devon, England: a contribution boniferous miospores of Western Europe: illustration to the early post-Variscan stratigraphy of northwest and zonation. Mededilingen Rijks Geologische Dienst, Europe. Geological Magazine, 134, 177–197. 29, 1–71. Elsik, W.C. 1999. Reduviasporonites Wilson 1962: syn- Cleal, C.J. & Thomas, B.A. 1991. Carboniferous and onymy of the fungal organism involved in the Late Permian palaeogeography. In: Cleal, C.J. (ed.) Plant Permian crisis. Palynology, 23, 37–41. Fossils in Geological Investigation: The Palaeozoic. Eshet, Y. 1990. Paleozoic–Mezozoic Palynology of Ellis Horwood, Chichester, 154–181. Israel. I: Palynological Aspects of the Permo-Trias Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Succession in the Subsurface of Israel. Geological Sur- Barbwire Terrace, Canning Basin, Australia. Austra- vey of Israel Bulletin, 81, 1–57. lian Journal of Earth Sciences, 35, 135–157. Evans, P.R. 1969. Upper Carboniferous and Permian pal- Fuglewicz, R. 1977. New species of megaspores from the ynological stages and their distribution in eastern Aus- Trias of Poland. Acta Palaeontologica Polonica, 22, tralia. In: Gondwana Stratigraphy: IUGS Symposium, 405–431. 1976, Argentina. UNESCO, Earth Sciences, 2, 41–54. Glenister, B.F., Baker, C., Furnish, W.M. & Dickens, Faddeeva, I.Z. 1980. Regularities in the changes of mio- J.M. 1990. Late Permian ammonoid cephalopod from spore complexes in stratotype sections of the East Western Australia. Journal of Palaeontology, 64, European platform and the Urals. In: Proceedings of 399–402. the Fourth International Palynological Conference, Gomankov, A.V. 1992. The interregional correlation of 1976–1977, Lucknow, Volume 2. Birbal Sahni Insti- the Tatarian and the problem of the Permian upper tute of Palaeobotany, Lucknow, India, 844–848. boundary. International Geology Review, 34, Falcon, R.M.S. 1975. Palynostratigraphy of the Lower 1015–1020. Karroo sequence in Sebungwe District, Mid Zambezi Gomankov, A.V., Balme, B.E. & Foster, C.B. 1998. Basin, Rhodesia. Palaeontologia Africana, 18, 1–29. Tatarian palynology of the Russian Platform: a review. Falcon, R.M.S., Pinheiro,H.&Sheperd, P. 1984. The Proceedings of the Royal Society of Victoria, 110, palynobiostratigraphy of the major coal seams in the 115–135. Witbank Basin with lithostratigraphic, chronostrati- Go¨tz, A.E. & Ruckwied, K. 2014. Palynological records graphic and palaeoclimatic implications. Comunica- of the Early Permian postglacial climate amelioration c¸o˜es dos Servic¸os Geolo´gicos de Portugal, 70, (Karoo Basin, South Africa). Palaeobiodiversity and 215–243. Palaeoenvironments, 94, 229–235. Farabee, M.J., Taylor, E.L. & Taylor, T.N. 1990. Cor- Gradstein, S.R. & Kerp, H. 2012. A brief history of relation of Permian and Triassic palynomorph assem- plants on Earth. In: Gradstein, F.M., Ogg, J.G., blages from the central Transantarctic Mountains, Schmitz,M.&Ogg, G. (eds) The Geologic Time Antarctica. Review of Palaeobotany and Palynology, Scale 2012. Elsevier, Amsterdam, 233–237. 65, 257–265. Grebe,H.&Schweitzer, H.-J. 1962. Die sporae disper- Fielding, C.R., Frank, T.D., Birgenheier, L.P., Rygel, sae des niederrheinischen Zechsteins. Fortschritte der M.C., Jones, A.T. & Roberts, J. 2008. Stratigraphic Geologie von Rheinland und Westfalen, Vorausdruck, record and facies associations of the late Paleozoic 10, 1–24. ice age in eastern Australia (New South Wales and Guerra-Sommer, M., Cazzulo-Klepzig, M., Menegat, Queensland). In: Fielding, C.R., Frank, T.D. & R., Formoso, M.L.L., Basei, M.A.S., Barboza, E.G. Isbell, J.L. (eds) Resolving the Late Paleozoic Ice & Simas, M.W. 2008. Geochronological data from Age in Time and Space. Geological Society of Amer- the Faxinal coal succession, southern Parana Basin, ica, Special Papers, 441, 41–57. Brazil: a preliminary approach combining radiometric Fischer, M.J. 1979. The Triassic palynofloral succession U–Pb dating and palynostratigraphy. Journal of South in the Canadian Arctic Archipelago. American Associ- American Earth Sciences, 25, 246–256. ation of Stratigraphic Palynologists Contributions Hart, G.F. 1964. A review of the classification and dis- Series, 5B, 83–100. tribution of the Permian miospore: Disaccate Striatiti. Foster, C.B. 1975. Permian plant microfossils from the 5th International Congress of Stratigraphy and Geol- Blair Atholl Coal Measures Central Queensland, Aus- ogy of the Carboniferous, Comptes Rendus, Paris tralia. Palaeontographica B, 154, 121–171. 1963, 3, 1171–1199. Foster, C.B. 1979. Permian Plant Microfossils of the Hart, G.F. 1965a. Microflora from the Ketewaka–Mchu- Blair Atholl Coal Measures, Baralaba Coal Measures chuma Coalfield, Tanganyika. Bulletin of the Geologi- and Basal Rewan Formation of Queensland. Geologi- cal Survey of Tanganyika, 36, 1–27. cal Survey of Queensland Publications, 372. Hart, G.F. 1965b . The systematics and distribution of Foster, C.B. & Afonin, S.A. 2005. Abnormal pollen Permian miospores. Witwatersrand University Press, grains: an outcome of deteriorating atmospheric condi- Johannesburg. tions around the Permian–Triassic boundary. Journal Hart, G.F. 1969. The stratigraphic subdivision and equiv- of the Geological Society, London, 162, 653–659, alents of the Karroo sequence as suggested by palynol- https://doi.org/10.1144/0016-764904-047 ogy. In: IUGS Gondwana Symposium Proceedings, Foster, C.B. & Afonin, S.A. 2006. Syndesmorion gen. 1967, Buenos Aires, 1, 23–35. nov. – a coenobial alga of Chlorococcalean affinity Hart, G.F. 1970. The biostratigraphy of Permian palyno- from the continental Permian–Triassic deposits of floras. Geoscience and Man, 1, 89–131. Dalongkou section, Xinjiang Province, China. Review Henderson, C.H., Davydov,V.&Wardlaw, B. 2012. of Palaeobotany and Palynology, 6, 1–8. The Permian period. In: Gradstein, F.M., Ogg, J.G., Foster, C.B. & Archbold, N.W. 2001. Chronologic Schmitz,M.&Ogg, G. (eds) The Geologic Time anchor points for the Permian Early Triassic of the Scale 2012. Elsevier, Amsterdam, 653–679. eastern Australian basins. In: Weiss, R.H. (ed.) Contri- Hermann, E., Hochuli, P.A., Bucher,H.&Roohi,G. butions to Geology and Palaeontology of Gondwana in 2012. Uppermost Permian to Middle Triassic palynol- Honour of Helmut Wopfner. Geological Institute, Uni- ogy of the Salt Range and Surghar Range, Pakistan. versity of Cologne, Cologne, Germany, 175–199. Review of Palaeobotany and Palynology, 169, 61–95. Foster, C.B. & Waterhouse, J.B. 1988. The Granulatis- Hochuli, P.A. 1985. Palynostratigraphische gleiderung porites confluens Oppel Zone and Early Permian und korrelation des Permo-Karbon der Nordoschweiz. marine faunas from the Grant Formation on the Eclogae Geologicae Helvetiae, 78, 719–831. Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Inosova, K.I., Krusina, A.K. & Shvartsman, E.G. 1975. Southern Tunisia – Biostratigraphy and palaeoenvir- The scheme of distribution of characteristic genera and onment. Review of Palaeobotany and Palynology, 66, species of microspores and pollen in Carboniferous and 273–291. Lower Permian deposits. In: Field Excursion Guide- Klaus, W. 1963. Sporen aus dem su¨dalpinen Perm. Jahr- book for the Donets Basin, Ministry of Geology of buch der Geologischen Bundesanstalt, Wien, 106, the Ukrainian SSR. Nauka, Moscow [in Russian & 229–363. English]. Kyle, R.A. 1977. Palynostratigraphy of the Victoria Group Isbell, J.L., Miller, M.F., Wolfe, K.L. & Lenaker, P.A. of south Victoria Land, Antarctica. New Zealand 2003. Timing of late Paleozoic glaciation in Gond- Journal of Geology and Geophysics, 20, 1081–1102. wana: was glaciation responsible for the development Larrsson, K., Lindstro¨m,S.&Guy-Ohlson, D. 1990. of northern hemisphere cyclothems? In: Chan, M.A. An early Permian palynoflora from Milorgfjella, Dron- & Archer, A.W. (eds) Extreme Depositional Environ- ning Maud Land, Antarctica. Antarctic Science, 2, ments: Mega End Members in Geologic Time. Geolog- 331–344. ical Society of America, Special Papers, 370, 5–24. Lei, Y., Servais,T.&Feng, Q. 2013. The diversity of the Jan, I.U., Stephenson, M.H. & Khan, F.R. 2009. Paly- Permian phytoplankton. Review of Palaeobotany and nostratigraphic correlation of the Sardhai Formation Palynology, 198, 145–161. (Permian) of Pakistan. Review of Palaeobotany and Lele, K.M. & Makada, R. 1972. Studies in the Talchir Palynology, 156, 402–442. flora of India – 7. Palynology of the Talchir Formation Jansonius, J. 1962. Palynology of Permian and Triassic in the Jayanti coalfield, Bihar. Geophytology, 2, 41–73. sediments, Peace River Area, Western Canada. Palae- Leonova, T.B. 1998. Permian ammonoids of Russia and ontographica B, 110, 35–98. Australia. Proceedings of the Royal Society of Victoria, Jerzykiewicz, J. 1987. Latest Carboniferous (Stephanian) 110, 157–162. and Early Permian (Autunian) palynological assem- Leschik, G. 1956. Sporen aus dem Salzton des Zechsteins blages from the Intrasudetic Basin, southwestern von Neuhof (bei Fulda). Palaeontographica B, 100, Poland. Palynology, 11, 117–131. 122–142. Jha, N., Sabina, K.P., Aggarwal,N.&Mahesh,S. Lindstro¨m, S. 1995a. Early Late palynostratigraphy and 2014. Late Permian Palynology and depositional envi- palaeo-biogeography of Vestfjella, Dronning Maud ronment of Chintalapudi sub basin, Pranhita–Godavari Land, Antarctica. Review of Palaeobotany and Paly- basin, Andhra Pradesh, India. Journal of Asian Earth nology, 86, 157–173. Sciences, 79, 382–399. Lindstro¨m, S. 1995b. Early Permian palynostratigraphy Jin, Y., Wardlaw, B.R., Glenister, B.F. & Kotlyar, of the Northern Heimefrontfjella Mountain Range, G.V. 1997. Permian chronostratigraphic subdivisions. Dronning Maud Land, Antarctica. Review of Palaeo- Episodes, 20, 10–15. botany and Palynology, 89, 359–415. Jizba, K.M.M. 1962. Late Palaeozoic bisaccate pollen Lindstro¨m,S.&McLoughlin, S. 2007. Synchronous from the United States midcontinent area. Journal of palynofloristic extinction and recovery after the end- Paleontology, 36, 871–887. Permian event in the Prince Charles Mountains, Ant- Jones, M.J. & Truswell, E.M. 1992. Late Carboniferous arctica: implications for palynofloristic turnover across and Early Permian palynostratigraphy of the Joe Joe Gondwana. Review of Palaeobotany and Palynology, Group, southern Galilee Basin, Queensland, and impli- 145, 89–122. cations for Gondwana stratigraphy. Bureau of Mines Liu, F., Zhu,H.&Ouyang, S. 2008. Late Carboniferous– and Mineral Resources Journal of Australian Geology Early Permian palynology of Baode (Pao-te-chou) and Geophysics, 13, 143–185. in Shanxi Province, North China. Geological Journal, Kaiser, H. 1976. Die permische Mikroflora der Cathaysia- 43, 487–510. schichten von nordwest-Schansi, China. Palaeontog- Liu, F., Zhu,H.&Ouyang, S. 2011. Taxonomy and raphica B, 159, 83–157. biostratigraphy of Pennsylvanian to Late Permian Kar, R.K. & Jain, K.P. 1975. Libya, a probable part of megaspores from Shanxi, North China. Review of Gondwanaland. Geophytology, 5, 72–80. Palaeobotany and Palynology, 165, 135–153. Kemp, E.M. 1973. Permian flora from the Beaver Loboziak, S., Clayton,G.&Owens, B. 1986. Aratris- Lake area, Prince Charles Mountains, Antarctica 1. porites saharaensis sp. nov.: a characteristic Lower Palynological examination of samples. Bulletin of the Carboniferous miospore species of North Africa. Geo- Bureau of Mineral Resources, Geology and Geophys- bios, 19, 497–503. ics, Australia, 126, 7–12. Looy, C.V., Twitchett, R.J., Dilcher, D.L., Van Kemp, E.M. 1975. Palynology of Late Palaeozoic glacial Konijnenburg-Van Cittert, J.H.A. & Visscher, deposits of Gondwanaland. In: Campbell, K.W. H. 2001. Life in the end-Permian dead zone. Proceed- (ed.) Gondwana Geology. Australian National Univer- ings of the National Academy of Sciences of the United sity Press, Canberra, 125–134. States of America, 98, 7879–7883. Kemp, E.M., Balme, B.E., Helby, R.J., Kyle, R.A., Love, C.F. 1994. The palynostratigraphy of the Haushi Playford,G.&Price, P.L. 1977. Carboniferous and Group (Westphalian–Artinskian) in Oman. In: Permian palynostratigraphy in Australia and Antarc- Simmons, M.D. (ed.) Micropalaeontology and Hydro- tica: a review. Bureau of Mines and Mineral Resources carbon Exploration in the Middle East. Chapman & Journal of Australian Geology and Geophysics, 2, Hall, London, 23–39. 177–208. MacRae, C.S. 1988. Palynostratigraphical Correlation Kilani-Mazraoui, F., Razgallah-Gargouri,S.& Between the Lower Karoo Sequence of the Waterburg Mannai-Tayech, B. 1990. The Permo-Triassic of and Pafuri Coal Basins and the Hammanskraal Plant Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Macrofossil Locality, RSA. Memoirs of the Geological Ouyang, S. 1982. Upper Permian and Lower Triassic Survey of South Africa, 75. palynomorphs from eastern Yunnan, China. Canadian Maheshwari, H.K. & Banerji, J. 1975. Lower Triassic Journal of Earth Sciences, 19, 68–80. palynomorphs from the Maitur Formation, West Ben- Ouyang,S.&Li, Z. 1980. Upper Carboniferous Spores gal, India. Palaeontographica B, 152, 149–190. from Shuo Xian, Northern Shanxi. Papers for the 5th Mangerud, G. 1994. Palynostratigraphy of the Permian International Palynological Conference, Nanjing Insti- and lowermost Triassic succession, Finnmark Plat- tute of Geology and Paleontology, Academia Sinica, form, Barents Sea. Review of Palaeobotany and Paly- Special Publication, 1–16. nology, 82, 317–349. Ouyang,S.&Utting, J. 1990. Palynology of Upper Mangerud,G.&Konieczny, R.M. 1993. Palynology of Permian and Lower Triassic rocks, Meishan, Changx- the Permian succession of Spitsbergen, Svalbard. ing County, Zhejiang Province, China. Review of Polar Research, 12, 65–93. Palaeobotany and Palynology, 66, 65–103. Marcinkiewicz, T., Fijalkowska-Mader,A.&Pien- Ouyang, S., Wang, Z., Zhan, J.Z. & Zhou, Y.X. 2003. kowski, G. 2014. Megaspore zones of the epicontinen- Palynology of Carboniferous–Permian Strata in tal Triassic and Jurassic deposits in Poland – overview. Northern Xinjiang. University of Science and Technol- Biuletyn Pan˜stwowego Instytutu Geologicznego, 457, ogy of China Press, Hefei, Anhui (in Chinese with 15–42. English summary). Marques-Toigo, M. 1974. Some new species of spores Pattison, J., Smith, D.B. & Warrington, G. 1973. A and pollens of Lower Permian age from the San Gre- review of Late Permian and Early Triassic biostratigra- gorio Formation in Uruguay. Anais da Academia Bra- phy in the British Isles. In: Logan,A.&Hills, L.V. siliera de Cieˆncas, 46, 601–616. (eds) The Permian and Triassic Systems and their Metcalfe, I., Foster, C.B., Afonin, S.A., Nicoll, R.S., Mutual Boundary. Canadian Society of Petroleum Mundil, R., Xiaofeng,W.&Lucas, S.G. 2009. Geologists, Memoirs, 2, 220–260. Stratigraphy, biostratigraphy and C-isotopes of the Penney, R.A., Al Barram,I.&Stephenson, M.H. 2008. Permian–Triassic non-marine sequence at Dalongkou A high resolution palynozonation for the Al Khlata and Lucaogou, Xinjiang Province, China. Journal of Formation (Pennsylvanian to Lower Permian), South Asian Earth Sciences, 36, 503–520. Oman. Palynology, 32, 213–231. Meyen, S.V. 1969. New data on relationship between Pe´rez Loinaze, V.S. & Ce´sari, S.N. 2004. Palynology Angara and Gondwana Late Palaeozoic floras. In: of the Estratos de Mascası´n, Upper Carboniferous. Gondwana Stratigraphy, IUGS Symposium, 1967, Paganzo Basin, Argentina: systematic descriptions Buenos Aires. UNESCO, Paris, 141–157. and stratigraphic considerations. Revista Espan˜ola de Millsteed, B.D. 1993. Palynological evidence for the age Micropaleontologı´a, 36, 407–438. of the Permian Karroo coal deposits near Vereeniging, Piasecki, S. 1984. Preliminary palynostratigraphy of the Northern Orange Free State, South Africa. South Afri- Permian-Lower Triassic sediments in Jameson Land can Journal of Geology, 97, 15–20. and Scoresby Land, East Greenland. Bulletin of the Millsteed, B.D. 1999. Palynology of the Early Permian Geological Society of Denmark, 32, 139–144. coal-bearing deposits near Vereeniging, Free State, Playford, G. 1962. Lower Carboniferous microfloras of South Africa. Bulletin of the Council for Geoscience, Spitzbergen. Palaeontology, 5, 550–618. 124, 1–77. Playford, G. 1986. Morphological and preservational Modie, B.N. & Le He´risse´, A. 2009. Late Palaeozoic variation of Rattiganispora Playford and Helby, palynomorph assemblages from the Karoo Supergroup 1968, from the Australian Carboniferous. Pollen and their potential for biostratigraphic correlation, Kal- Spores, 28, 83–96. ahari Karoo Basin, Botswana. Bulletin of Geosciences, Playford,G.&Dino, R. 2000. Palynostratigraphy of the 84, 337–358. upper Palaeozoic strata (Tapajo´s Group) Amazonas Mori, A.L.O., Souza, P.A., Marques, J.C. & Lopes, R.C. Basin, Brazil: Part Two. Palaeontographica B, 255, 2012. A new U–Pb zircon age dating and palyno- 87–145. logical data from a Lower Permian section of the south- Playford,G.&Dino, R. 2002. Permian palynofloral ernmost Parana´ Basin, Brazil: biochronostratigraphical assemblages of the Chaco-Parana´ Basin, Argentina: and geochronological implications for Gondwanan systematics and stratigraphic significance. Revista correlations. Gondwana Research, 21, 654–669. Espan˜ola de Micropaleontologı´a, 34, 235–288. Nader, A.D., Khalaf, F.H. & Hadid, A.A. 1993a. Paly- Playford,G.&Dino, R. 2005. Carboniferous and Perm- nology of the Permo-Triassic boundary in Borehole ian Palynostratigraphy. In: Koutsoukos, E. (ed.) Mityah-1, south west Mosul City, Iraq. Mu’tah Journal Applied Stratigraphy. Springer, Dordrecht, The Neth- Research and Studies, 8, 223–280. erlands, 101–121. Nader, A.D., Khalaf, F.H. & Yousif, R.A. 1993b. Paly- Playford,G.&Helby, R. 1968. Spores from a nology of the upper part of the Ga’ara Formation in the Carboniferous section in the Hunter Valley, NSW. western Iraqi desert. Mu’tah Journal Research and Journal of the Geological Society of Australia, 15, Studies, 8, 77–137. 103–119. Neves,R.&Owens, B. 1966. Some Namurian camerate Playford,G.&Powis, G.D. 1979. Taxonomy and distri- miospores from the English Pennines. Pollen Spores, bution of some trilete spores in Carboniferous strata of 8, 337–360. the Canning Basin, Western Australia. Pollen Spores, Nilsson, I., Mangerud,G.&Mørk, A. 1996. Permian 21, 371–394. stratigraphy of the Svalis Dome, south-western Barents Potonie´, R. 1958. Synopsis der Gattungen der Sporae dis- Sea. Norsk Geologisk Tidsskrift, 76, 127–146. persae. II. Teil: Sporites (Nachtra¨ge), Saccites, Aletes, Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Praecolpates, Polyplicates, Monocolpates. Beihefte Basin) at Arac¸oiba da Serra, Sa˜o Paulo State, Brazil. zum Geologischen Jahrbuch, 31, 1–114. Palynology, 27, 39–74. Potonie´,R.&Klaus, W. 1954. Einige sporegattungen Spina, A., Cirilli, S., Utting,J.&Jansonius, J. 2015. des alpinen Salzgebirges. Geologisches Jahrbuch, 68, Palynology of the Permian and Triassic of the Tesero 517–546. and Bulla sections (Western Dolomites, Italy) and Price, P.L. 1983. A Permian palynostratigraphy for consideration about the enigmatic species Reduvias- Queensland. In: Foster, C.B. (ed.) Permian Geology poronites chalastus. Review of Palaeobotany and Pal- of Queensland. Geological Society of Australia, Bris- ynology, 218, 3–14. bane, 155–211. Srivastava, S.C. & Bhattacharyya, A.P. 1996. Price, P.L. 1997. Permian to Jurassic palynostratigraphic Palynofloral assemblages from Permian sediments, nomenclature of the Bowen and Surat basins. In: West Siang District, Araunachal Pradesh, India. Green, P. (ed.) The Surat and Bowen Basins, South- In: Gondwana Nine: Ninth International Gondwana east Queensland. Queensland Department of Mines Symposium, 10–14 January 1994, Hyderabad, India. and Energy, Brisbane, 137–178. A.A. Balkema, Rotterdam, The Netherlands, 261– Rawson, P.F., Allen, P.M. et al. 2002. Stratigraphical 268. Procedure. Geological Society, London, Professional Stephenson, M.H. 2006. Stratigraphic Note: update of the Handbook. standard Arabian Permian palynological biozonation; Ruckwied, K., Go¨tz, A.E. & Jones, P. 2014. Paly- definition and description of OSPZ5 and 6. GeoArabia, nological records of the Permian Ecca Group (South 11, 173–178. Africa): utilizing climatic icehouse–greenhouse sig- Stephenson, M.H. 2008a. Review of the palynological nals for cross basin correlations. Palaeogeography, biostratigraphy of Gondwanan Late Carboniferous Palaeoclimatology, Palaeoecology, 413, 167–172. to Early Permian glacigene successions. In: Fiel- Samoilovich, S.R. 1953. Pollen and spores from the ding, C.R., Frank, T.D. & Isbell, J.L. (eds) Resolv- Permian deposits of the Cherdya and Aktyubinsk ing the Late Paleozoic Ice Age in Time and Space. areas, Cis-Urals. Oklahoma Geological Survey Circu- Geological Society of America, Special Papers, lar, 56, 1–103. 441, 304–320. Schaarschmidt, F. 1963. Spores und hystricosphaerideen Stephenson, M.H. 2008b. Spores and pollen from the aus dem Zechstein von Bu¨dingen in der Wetterau. Middle and Upper Gharif members (Permian) of Palaeontographica B, 113, 38–91. Oman. Palynology, 32, 157–183. Schaarschmidt, F. 1980. Pollen flora and vegetation at Stephenson, M.H. 2009. The age of the Carboniferous– the end of the Lower Permian. In: Proceedings of the Permian Converrucosisporites confluens Oppel Bio- IV International Palynological Conference, 1976– zone: new data from the Ganigobis Shale Member, 77, Lucknow, India, Volume 2. Birbal Sahni Institute, Dwyka Group, Namibia. Palynology, 33, 55–63. Lucknow, 750–752. Stephenson, M.H. 2015. Bisaccate pollen from the Early Schweitzer, H.-J. 1986. The land flora of the English and Permian OSPZ3a Sub-Biozone of the Lower Gharif German Zechstein sequences. In: Harwood, G.M. & Member, Oman. Review of Palaeobotany and Palynol- Smith, D.B. (eds) The English Zechstein and Related ogy, 212, 214–225. Topics. Geological Society, London, Special Publica- Stephenson, M.H. & McLean, D. 1999. International tions, 22, 31–54, https://doi.org/10.1144/GSL.SP. correlation of Early Permian palynofloras from the 1986.022.01.04 Karoo sediments of Morupule, Botswana. South Afri- Segroves, K.L. 1970. Permian spores and pollen grains can Journal of Geology, 102, 3–14. from the Perth Basin, Western Australia. Grana, 10, Stephenson, M.H. & Osterloff, P.L. 2002. Palynology 43–73. of the Deglaciation Sequence Represented by the Singh, H.P. 1964. A miospore assemblage from the Perm- Lower Permian Rahab and Lower Gharif Members, ian of Iraq. Palaeontology, 7, 240–265. Oman. American Association of Stratigraphic Palynol- Smith, D.B., Brunstrom, R.G.W., Manning, P.I., Simp- ogists, Contribution Series, 40. son,S.&Shotton, F.W. 1974. A Correlation of the Stephenson, M.H. & Powell, J.H. 2013. Palynology Permian Rocks of the British Isles. Geological Society, and alluvial architecture in the Permian Umm London, Special Reports, 5. Irna Formation, Dead Sea, Jordan. GeoArabia, 18, Somers, Y. 1972. Re´vision du genre Lycospora Schopf, 17–60. Wilson & Bentall – Microfossiles Organiques du Pale´- Stephenson, M.H. & Powell, J.H. 2014. Selected spores ozoı¨que. 5, Spores. Centre National de la Recherche and pollen from the Permian Umm Irna Formation, Jor- Scientifique, 11–110. dan, and their stratigraphic utility in the Middle East Souza, P.A. 2006. Late Carboniferous palynostratigraphy and North Africa. Rivista Italiana di Paleontologia e of the Itarare´ Subgroup northeastern Parana´ Basin, Stratigrafia, 120, 145–156. Brazil. Review of Palaeobotany and Palynology, 138, Stephenson, M.H., Osterloff, P.L. & Filatoff,J. 9–29. 2003. Palynological biozonation of the Permian of Souza, P.A. & Marques-Toigo, M. 2003. An overview Oman and Saudi Arabia: progress and challenges. Geo- on the palynostratigraphy of the Upper Paleozoic strata Arabia, 8, 467–496. of the Brazilian Parana´ Basin. Revista del Museo Stephenson, M.H., Angiolini,L.&Leng, M.J. 2007. Argentino Ciencias Naturales, Nueva Serie, 5, The Early Permian fossil record of Gondwana and its 205–214. relationship to deglaciation: a review. In: Williams, Souza, P.A., Petri,S.&Dino, R. 2003. Late Carbonifer- M., Haywood, A.M., Gregory, F.J. & Schmidt, ous palynology from the Itarare´ Subgroup (Parana´ D.N. (eds) Deep-Time Perspective on Climate Change: Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

M. H. STEPHENSON

Marrying the Signal from Computer Models and Bio- Utting, J. 1978. Lower Karroo pollen and spore assem- logical Proxies. The Micropalaeontological Society, blages from the coal measures and underlying sedi- Special Publications. Geological Society, London, ments of the Siankondobo Coalfield, Mid Zambezi 103–122. Valley, Zambia. Palynology, 2, 53–68. Stephenson, M.H., Al Rawahi,A.&Casey, B. 2008a. Utting, J. 1989. Preliminary palynological zonation of Correlation of the Al Khlata Formation in the surface and subsurface sections of Carboniferous, Mukhaizna Field, Oman, based on a new downhole, Permian and lowest Triassic rocks, Sverdrup Basin, cuttings-based palynostratigraphic scheme. GeoAra- Canadian Arctic Archipelago. Geological Survey of bia, 13, 45–75. Canada Paper, 89–1G, 233–241. Stephenson, M.H., Angiolini,L.et al. 2008b. Abrupt Utting, J. 1994. Palynostratigraphy of Permian and Lower environmental and climatic change during the deposi- Triassic Rocks, Sverdrup Basin, Canadian Arctic Archi- tion of the Early Permian Haushi limestone, Oman. pelago. Geological Survey of Canada Bulletin, 478. Palaeogeography, Palaeoclimatology, Palaeoecology, Utting,J.&Piasecki, S. 1995. Palynology of the Perm- 270, 1–18. ian of northern Continents: a review. In: Scholle, Stephenson, M.H., Jan, I.U. & Al-Mashaikie, S.Z.A.K. P.A., Peryt, T.M. & Ulmer-Scholle, P.A. (eds) 2013. Palynology and correlation of Carboniferous– The Permian of Northern Pangea, Volume 1. Palaeo- Permian glacigene rocks in Oman, Yemen and Paki- geography, Palaeoclimates, Stratigraphy. Springer, stan. Gondwana Research, 24, 203–211. Berlin, 236–261. Stolle, E. 2007. Regional Permian Palynological correla- Utting, J., Esaulova, N.K., Silantiev, V.V. &. Makar- tions: southeast Turkey–northern Iraq. Comunicac¸o˜es ova, O.V. 1997. Late Permian palynomorph assem- Geolo´gicas, 94, 125–143. blages from Ufimian and Kazanian type sequences in Stump, T.E. & van der Eem, J.G. 1995. The stratigraphy, Russia and comparison with Roadian and Wordian depositional environments and periods of deformation assemblages from the Canadian Arctic. Canadian of the Wajid outcrop belt, southwestern Saudi Arabia. Journal of Earth Sciences, 34, 1–16. Journal of African Earth Sciences, 21, 421–441. Van der Zwan, C.J. 1981. Palynology, phytogeography Sullivan, H.J. 1965. Palynological evidence concerning and climate of the Lower Carboniferous. Palaeogeogra- the regional differentiation of Upper Mississippian flo- phy, Palaeoclimatology, Palaeoecology, 33, 279–310. ras. Pollen Spores, 7, 539–563. Venkatachala, B.S. & Kar, R.K. 1966. Corisaccites gen. Tewari, R., Ram-Awatar, et al. 2015. The Permian– nov., a new saccate pollen genus from the Permian of Triassic palynological transition in the Guryul Ravine the Salt Range, West Pakistan. Palaeobotanist, 15, section, Kashmir, India: implications for Tethyan– 107–109. Gondwanan correlations. Earth-Science Reviews, Venkatachala, B.S. & Kar, R.K. 1968. Palynology of 149, 53–66. the Kathwai Shales, Salt Range, West Pakistan. 1. Tiwari, R.S. 1965. Miospore assemblage in some coals Shales 25ft above the Talchir Boulder Bed. Palaeo- of the Barakar Stage (Lower Gondwana) of India. botanist, 16, 156–166. Palaeobotanist, 13, 168–214. Vergel, M. del M. 1993. Palinoestratigrafı´a de la secuen- Tiwari, R.S. 1996. Palynoevent stratigraphy in Gondwana cia Neopalaeozoica de la Cuenca Chacoparanense, sequence of India. In: Gondwana Nine: Ninth Interna- Argentina. In: 12th Congre`s International de la Strat- tional Gondwana Symposium, 10–14 January 1994, igraphie et Ge´ologie du Carbonife`re et Permien. Hyderabad, India. A.A. Balkema, Rotterdam, The Compte Rendus, 1, 201–212. Netherlands, 3–19. Vijaya 1996. Advent of Gondwana deposition on Indian Tiwari, R.S. & Moiz, A.A. 1971. Palynological study of Peninsula: a palynological reflection and relationship. Lower Gondwana (Permian) coals from the Godavari In: Gondwana Nine: Ninth International Gondwana Basin, India. Palaeobotanist, 19, 95–104. Symposium, 10–14 January 1994, Hyderabad, India. Tiwari, R.S. & Tripathi, A. 1992. Marker assemblage A.A. Balkema, Rotterdam, The Netherlands, 283–298. zones of spore and pollen species through the Gond- Visscher, H. 1971. The Permian and Triassic of the King- wana Palaeozoic and Mezozoic sequence in India. scourt Outlier, Ireland. Geological Survey of Ireland, Palaeobotanist, 40, 194–236. Special Papers, 1. Truswell, E.M. 1980. Permo-Carboniferous palyno- Visscher, H. 1973. The Upper Permian of Western logy of Gondwanaland: progress and problems in the Europe – a palynological approach to chronostratigra- decade of 1980. Bureau of Mines and Mineral phy. In: Logan,A.&Hills, L.V. (eds) The Permian Resources Journal of Australian Geology and Geo- and Triassic Systems and their Mutual Boundary. physics, 5, 95–111. Canadian Society of Petroleum Geologists, Memoirs, Tschudy, R.H. & Kosanke, R.M. 1966. Early Permian 2 , 200–219. vesiculate pollen from Texas, USA. Palaeobotanist, Visscher, H. 1980. Aspects of a palynological charac- 15, 59–71. terisation of Late Permian and Early Triassic ‘stan- Turnau, E. 1978. Spore zonation of uppermost Devo- dard’ units of chronostratigraphical classification in nian and Lower Carboniferous deposits of western Europe. In: Proceedings of the IV International Paly- Pomerania. Mededelingen rijks geologische dienst, nological Conference, 1976–77, Lucknow, India, Vol- 30, 1–35. ume 2. Birbal Sahni Institute, Lucknow, 236–244. Twitchett, R.J., Looy, C.V., Morante, R., Visscher, Warrington, G. 1996. Palaeozoic spores and pollen H. & Wignall, P.B. 2001. Rapid and synchronous col- (Chapter 18E) Permian. In: Jansonius,J.&Mc- lapse of marine and terrestrial ecosystems during the Gregor, D.C. (eds) Palynology: Principles and end-Permian biotic crisis. Geology, 29, 351– 354. Applications, Volume 3. American Association of Downloaded from http://sp.lyellcollection.org/ by guest on October 1, 2021

PERMIAN PALYNOSTRATIGRAPHY

Stratigraphical Palynologists Foundation, Dallas, TX, Syncline. Trydy Geologicheskogo Instituta Akademii 607–619. Nauk SSSR, Moscow (in Russian). Wilson, L.R. 1962. Permian Plant Microfossils from the Yin, H., Zhang, K., Tong, J., Yang,Z.&Wu, S. 2001. Flowerpot Formation, Greer County, Oklahoma. The Global Stratotype Section and Point (GSSP) Oklahoma Geological Survey, Circular, 49. of the Permian–Triassic Boundary. Episodes, 24, Wilson, L.R. & Coe, E.A. 1940. Descriptions of some 102–114. unassigned plant microfossils from the Des Moines Zhu, H.-C., Ouyang, S., Zhan, J.-Z. & Wang, Z. 2005. Series of Iowa. American Midland Naturalist, 23, Comparison of Permian palynological assemblages 182–186. from the Junggar and Tarim Basins and their phyto- Yaroshenko, O.P. & Golubeva, L.P. 1991. Miospores provincial significance. Review of Palaeobotany and and stratigraphy of the lower Triassic of Pechora Palynology, 136, 181–207.