Earliest Triassic (Induan) spores and pollen from the Junggar Basin, Xinjiang, northwestern China

Earliest Triassic (Induan) spores and pollen from the Junggar Basin, Xinjiang, northwestern China

ELSEVIER Review of Palaeobotany and Palynology 106 (1999) 1–56 www.elsevier.com/locate/revpalbo Earliest Triassic (Induan) spores and pollen from th...

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ELSEVIER

Review of Palaeobotany and Palynology 106 (1999) 1–56 www.elsevier.com/locate/revpalbo

Earliest Triassic (Induan) spores and pollen from the Junggar Basin, Xinjiang, northwestern China Ouyang Shu a , Geoffrey Norris b,* a

Nanjing Institute of Geology and Palaeontology. Academia Sinica Chi-Ming-Ssu, Nanjing 210008, People’s Republic of China b Department of Geology, University of Toronto, Toronto, Ont. M5S 3B1, Canada Received 1 April 1996; accepted 8 December 1998

Abstract Permian–Triassic boundary strata cropping out in the southern Junggar Basin contain a continuous and varied fossil record, making this fluvio-lacustrine succession a potential candidate for a non-marine boundary stratotype. Abundant, diverse and well preserved miospores are recorded and illustrated from the upper part of the Guodikeng Formation and the basal part of the Jiucaiyuan Formation, comprising 92 species and 48 genera, as well as some acritarchs. Seven new species (Anapiculatisporites decorus Ouyang et Norris, sp. nov., Baculatisporites uniformis Ouyang et Norris, sp. nov., Kraeuselisporites varius Ouyang et Norris, sp. nov., Lapposisporites echinatus Ouyang et Norris, sp. nov., Klausipollenites angustus Shu et Norris, sp. nov., Alisporites exilis Ouyang et Norris, sp. nov., Pilasporites perreticulatus Ouyang et Norris, sp. nov.) and 8 new combinations are proposed. Pteridophytic spores and gymnospermous pollen are almost equally diverse, comprising 45 species in 20 genera, and 43 species in 25 genera, respectively. Pteridophytes are numerically more important, notably Limatulasporites, Kraeuselisporites, Lundbladispora, Anapiculatisporites and Verrucosisporites, but also prominent are bisaccate pollen of Falcisporites, Klausipollenites and Alisporites. Less abundant taeniate pollen are largely bisaccate. These palynofloras are grouped together as the Lundbladispora–Lunatisporites–Aratrisporites Assemblage, argued to be Early Triassic (early Induan), and comprising both newly evolved Mesozoic taxa (herbaceous and shrubby lycopsids, coniferous trees, pteridosperms) and relict Paleozoic taxa (sphenophyllids, cordaitaleans and possibly Paleozoic lycopods). The lower boundary of the Triassic in the section is redrawn somewhat lower than by previous authors. The transitional nature of the Permian–Triassic palynofloras and of the paleophytoprovinces is discussed; neither indicates a sudden catastrophic event affecting terrestrial floras at the P–T boundary on a global scale. The Early Triassic vegetation appears to have grown under humid sub-tropical conditions in lowland flood plains and marshes dominated by hydrophilous and mesophilous pteridophytes and pteridosperms, but with locally drier upland environments supporting xerophilous and mesophilous conifers and pteridosperms. The Induan vegetation is closely comparable to that of the Angara, especially the Subangara Province, although it contains some elements of the Euramerican Province in association with a few Gondwanan, Cathaysian and cosmopolitan taxa.  1999 Elsevier Science B.V. All rights reserved. Keywords: palynoflora; spores; pollen; biostratigraphy; phytoprovince; paleoclimate; taxonomy; Lower Triassic; Upper Permian; China

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1. Introduction The macrofossil plant record in the Upper Permian and Lower Triassic strata of the world is relatively meager. As a result, the landscape of this period was often described in paleobotanical literature as desert-like and the flora as sparse. However, palynological data accumulated in the last decade have demonstrated that such conclusions are exaggerated and possibly related to such factors as the increasing continentality, expansion of an arid climate, lowering of the groundwater table, and development of oxidizing environments that promoted taphonomic effects unfavourable to the preservation of macroscopic plant remains. Highly diverse palynofloras are now known in some areas. Nevertheless, our knowledge about the botanical background, ecosystems, phytoprovincialism (especially in the Northern Hemisphere) and paleoclimatic implications of these assemblages are still far from complete. Detailed palynological study near the Permian–Triassic boundary is urgently needed to demarcate the Permian– Triassic boundary in marine and terrestrial strata, as well as to address the problem of putative mass extinction events at the close of the Permian. Xinjiang is located in an interesting geographical position relative to Permian–Triassic phytogeographic provinces, and exhibits well-documented upper Paleozoic and Mesozoic strata. The investigation of the Permo-Triassic strata in Xinjiang, particularly in the Urumqi Jimsar area (Fig. 1), began more than fifty years ago when Professor Yuan Fu-li surveyed the area from 1928 to 1932, jointly with the Chinese–Swedish Expedition Group. The Dicynodontia and Lystrosaurus faunas were published by this expedition in 1935. In 1959, Professor Pan reported abundant fossil plants from the Upper Permian strata in the Cangfanggou Group at Dalongkou in Jimsar. The systematic and comprehensive studies made by the Institute of Geology, Chinese Academy of Geological Sciences, and the Institute of Geological Sciences, Xinjiang Bureau of Geology and Mineral Resources (Anon., 1986, 1989) as well as the Research Institute of Petroleum Exploration and Development, Xinjiang Petroleum Administration (Anon., 1990) are of particular importance, notably those from several sections at Dalongkou which include plants, miospores, megaspores, bivalves, os-

tracods, conchostracans and vertebrates. Six palynological papers are significant, covering the Upper Permian (Hou and Wang, 1986, 1990) and Triassic (Qu and Wang, 1986, 1990; Yang and Sun, 1986, 1990). Wang and Zhan (1987) reported in an abstract on the palynological study of the Carboniferous and Permian in the Junggar Basin, northern Xinjiang. In recent years, the Nanjing Institute of Geology and Palaeontology, Academia Sinica and the Institute of Exploration and Development, Xinjiang Petroleum Administration, China have undertaken a cooperative palynological study of the Carboniferous and Permian strata in northern Xinjiang. The present palynological investigation is thus a continuation and development of the work by Qu and Wang (1986, 1990) and Hou and Wang (1986, 1990) in the transition sequence in the Dalongkou area. The objectives of the present study are: (a) to assess the paleovegetation, paleoenvironments (ecosystems), and paleoclimate based on the palynofloras; (b) to determine more precisely the age of the upper part of the Guodikeng Formation and the lower boundary of the Lower Triassic for the Permian– Triassic sequence developed at Dalongkou; (c) to determine whether a transitional (or ‘mixed’) palynoflora occurs in the terrestrial basal Lower Triassic similar to those recorded from the marine Lower Triassic in Kap Stosch, East Greenland (Balme, 1980) and in Changxing, Zhejiang, eastern China (Ouyang and Utting, 1990) as well as the coastal lower Scythian Kayitou Formation in eastern Yunnan, southwestern China (Ouyang and Li, 1980; Ouyang, 1982, 1986); (d) to assess evidence for a mass extinction at the close of the Permian as some authors have advocated (e.g. Wang, 1989); and (e) to examine the extent of phytoprovincialism of the Early Triassic flora.

2. Stratigraphy 2.1. Outline of Permian–Triassic stratigraphy at Dalongkou, southern Junggar Basin Terrestrial Permian–Triassic sequences are widespread north of the Kunlun–Qinling mountain

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Fig. 1. Map of the southern Junggar Basin showing the locality of the Permian–Triassic section on the north limb of the Dalongkou Anticline (approximately 8 km southwest of the town of Santai). Inset map symbols as follows: Wide cross-hachuring D Devonian– Carboniferous; narrow cross-hachuring D Triassic; circles and dots D Permian; dots D Jurassic; oblique lines D Cretaceous; blank areas D Quaternary. Scale of inset map: 3.5 cm D 50 km.

ranges, and are best represented in the southern Junggar Basin, northern Xinjiang, especially the sections at Dalongkou, Jimsar. Here, not only are the strata beautifully exposed representing essentially continuous sedimentation with abundant fossils, but tectonically this area is very simple (Figs. 1 and 2). The long history of geological investigation of the basin provides another advantage. Therefore, it is an ideal candidate for establishing the stratotype for the terrestrial Permian–Triassic boundary in China which we believe is located near the top of the Guodikeng Formation (middle part of the

Cangfanggou Group) as explained in detail below. The coeval Permian–Triassic Shihchienfeng Group is widely distributed in northern China proper (typically in Shanxi); however, the fossil records in this unit are less continuous. The upper Upper Permian and Lower Triassic strata in the Jimsar–Fukan area are assigned to the Cangfanggou Group, which consists mainly of variegated and red clastic rocks and includes 5 formations. The stratigraphic classification of Li et al. (1986) is tabulated below in ascending order (the thickness of each formation cropping out on the

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Fig. 2. Field sketch of the presumed Permian–Triassic boundary section along the northern limb of the Dalongkou Anticline showing the locations of the samples. AEA 749 to AEA 757 refer to sample numbers for palynological study and ADI 1108 to ADI 1110 to specimen numbers for macroscopic fossils. Dip and strike are shown at each end of the outcrop as 45ºN=50º and 37ºN=75º. Tij D Jiucaiyuan Formation (4–5); P2g D Guodikeng Formation (1–3).

3. Guodikeng Fm. (P2 –T1 g, 144.17 m)

ness. Our survey in 1985 and 1986 did not measure the entire section but concentrated on the formations across the Permian–Triassic boundary, including the upper part of the Guodikeng Formation and the lower part of the Jiucaiyuan Formation. Based on the summary of Shen Yanbing (pers. commun., 1988), the lithology of the latter is as follows (see Fig. 2), sample numbers being shown in parentheses in the unit descriptions:

2. Wutonggou Fm. (P2 w, 220.64 m)

Lower Triassic Jiucaiyuan Formation (T1 j):

northern flank of the Dalongkou Anticline is based on Xiao et al., 1989): Overlying strata: Karamay Fm. (Middle Triassic) Cangfanggou Group 5. Shaofanggou Fm. (T1 s, 259.73 m) ?––––––––––? 4. Jiucaiyuan Fm. (T1 j, 220.60 m)

1. Quanzijie Fm. (P2 q, 54.46 m) Underlying strata: Hongyanchi Fm. (Upper Permian)

Principal fossils found in the uppermost Permian and lowermost Triassic of the Cangfanggou Group are listed herein in Tables 1 and 2 (compiled from Anon., 1986; Wu, 1989; Shen, pers. commun.; and the present study). 2.2. Materials The samples for the present study were collected from the north limb of the anticline at Dalongkou, Jimsar, near Urumqi in the southern Junggar Basin of northern Xinjiang (Fig. 1, 44º10 N, 88º540 E). This section is about 8 km south of the town of Santai in Jimsar County. According to Li et al. (1986), the Cangfanggou Group in this section is about 900 m in total thick-

Unit 5. Grayish-purple thin-bedded siltstones and finegrained sandstones, yielding Lystrosaurus sp. (AEA 757) 20 m Unit 4. Yellowish-gray thick-bedded and coarse sandstones with pebble conglomerates locally, being discontinuous laterally in the lower part; gray thin-bedded and fine-grained sandstones intercalated with grayish green, grayish-purple sandy mudstones in the upper part, yielding conchostracans (AEA 755–756): Sinolimnadiopsis jimsarensis, Falsisca sp., and Aquilonoglypta sp.; ostracods: Darwinula spp.; and the Lundbladispora–Lunatisporites–Aratrisporites miospore Assemblage (AEA 755) with rare acanthomorphic acritarchs 15 m Conformable contact Upper Permian–Lower Triassic Guodikeng Formation: Unit 3. Gray to dark gray mudstones and siltstones intercalated with slightly purplish-red siltstones and beds of yellowish-gray moderately thick-bedded and

Formation

Age

Jiucaiyuan

early Early Triassic

Bivalves

Ostracods

Vertebrates

Darwinula triassiana, Sinolimnadiopsis, Lystrosaurus D. rotundata Falcisca Chasmatosaurus

Guodikeng

Darwinula paraquangzijiensis, D. pararotundata late Late Permian

Conchostracans

Palaeanodonta cf. parallela, P. glossitiformis, P. ficheri, Microdontalla plotnikovskiensis

Plants

Miospores

Pecopteris sp., Lepidodendron sp.

Lundbladispora– Lunatisporites Assemblage

Callipteris sp., Cordaites sp., Zamiopteris cf. glossopteroides, Walchia? sp.

Lueckisporites virkkiae– Klausipollenites schaubergeri Assemblage

Sinolimnadiopsis– Dicynodontia– Falcisca Lystrosaurus

Dicynodontia Polygrapta, Panxiania Sinolimnadiopsis, xinjiangensis, Falcisca Vymella subglobica, Darwinula schwegeri, Darwinuloides

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Table 1 Principal fossils in the Permian–Triassic of the upper part of the Cangfanggou Group in northern Xinjiang

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fine-grained sandstones, yielding conchostracans (AEA 749, 753): Sinolimnadiopsis jimsarensis, Falsisca sp.; ostracods; bivalves and a few gastropods as well as the Lundbladispora–Lunatisporites–Aratrisporites miospore Assemblage (AEA 749, 751) 50 m Unit 2. Variegated (gray, yellowish-black and grayish-purple) mudstones and siltstones intercalated with thin-bedded or lenticular marls, yielding abundant conchostracans (ADI 1108–1110): Polygrapta xinjiangensis Liu, Tripemphigus sp., Falsisca sp., Sinolimnadiopsis jimsarensis (a few); ostracods: Darwinula spp., Darwinuloides sp., Panxiania sp., Suchonella sp.; bivalves: Palaeanodonta spp., Palaeomutella sp., Microdonta sp., Microdontella spp., Oligodon sp.; charophytes: Paracuneatochara jimsarensis, P. xinjiangensis; plants: Callipteris sp. and Cordaites sp. 50 m Unit 1. Gray, yellowish-gray interbedded with purplish-gray siltstones and fine-grained sandstones 15 m

Nine samples were collected and macerated, but only three yielded rich pollen and spore assemblages and the remaining 6 samples are barren or yielded only a few poorly preserved miospores. One sample (AEA 755) is undoubtedly Lower Triassic, while the other two (AEA 749 and 751) belong to the upper part of the Guodikeng Formation which was originally assigned to the Upper Permian. However, Li et al. (1986) have pointed out that the uppermost part (ca. 30 m) might be Early Triassic in age. On the basis of the presence of Triassic conchostracans and particularly miospores (Lundbladispora, Lunatisporites and Aratrisporites), all of Unit 3 (50 m) containing these two samples has been assigned also to the Lower Triassic. Thus the Permian–Triassic boundary drawn by us is slightly lower than that of former authors as will be discussed later.

3. Methods The 9 samples were processed using standard techniques. One hundred grams of each were cleaned, pre-treated with 10% HCl, followed by 48 hours in 42% HF and a further treatment in 36% HCl to remove fluoride by-products. Heavy liquid separation (SG D 2.0–2.1) was used to remove residual minerals. The organic residues were mounted in glycerin jelly and sealed with paraffin wax. All sam-

ples are stored in the Nanjing Institute of Geology and Palaeontology, Academia Sinica, Chi-Ming-Ssu, Nanjing 210008. Quantitative analyses were made on the 3 palyniferous samples counting the following numbers of grains: AEA 749 — 278 grains; AEA 751 — 333 grains; AEA 755 — 300 grains. The relative abundances of each taxon as well as some important genera and higher taxa are shown in Table 2 and in Fig. 3 (due to limited space, the bars indicating the highest values are not proportional but the percentages are marked on the top of each bar).

4. Characteristics of the early Early Triassic palynoflora The palynofloras found in the three samples from the Guodikeng and Jiucaiyuan formations (see Table 2) are similar in species composition. Therefore they are grouped as a single assemblage (Table 1): the Lundbladispora–Lunatisporites– Aratrisporites Assemblage. Limatulasporites and Kraeuselisporites are dominant in the assemblage, but they are not selected as indices because they are also abundant in the underlying Upper Permian strata. The assemblage contains 92 species referred to 48 genera, of which 45 species in 20 genera are pteridophytic (and possibly also a few bryophytic) spores, and 43 species in 25 genera are gymnospermous pollen. The remainder includes 3 species in 2 genera of acritarchs and Tympanicysta stoschiana — this latter being probably of algal or fungal origin. All the taxa and their relative abundance are systematically listed in Table 2. The stratigraphic distribution of the 93 species is as follows: Number of species

AEA 749

11 37 6 27 6 6

C C C

AEA 751

AEA 755

C C C C

C

C C

A significant number of species appear to be restricted to sample AEA 751, but this probably

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Fig. 3. The relative abundances of some important spore–pollen genera and higher taxa in the analyzed samples.

reflects that the palynomorph assemblage is better preserved in this sample. Many constituent species are still of Permian aspect which might represent upward extensions of range (or less probably some reworking from older strata), including Densosporites cf. D. spongeosus, Cingulizonates sp., Cordaitina rugulifera, C. gemina, Crucisaccites sp., Iunctella sp., Vesicaspora ex gr. magnalis and Florinites? sp. in AEA 751, as well as Protohaploxypinus spp., Striatopodocarpites sp., Hamiapollenites bullaeformis, Cycadopites retroflexus, Klausipollenites schaubergeri and Striatopodocarpites cf. S. pantii in AEA 751 and AEA 755. In other words, more than 15 species should occur in the underlying strata. Consequently, there are about 60 species (about 2=3 of the whole assemblage) which may range upward from the lowest (AEA 749) to highest (AEA 755) or at least to

the middle (AEA 751) sample. This indicates that the three palynofloras are not essentially different, although some compositional differences are apparent. The three palynofloras are also closely comparable at the subturmal level i.e. they are all characterized by a slight dominance of pteridophytic spores (55, 52 and 59%, respectively) and by the sub-dominance of gymnospermous pollen (38, 47 and 38% based on 105, 157 and 115 grains, respectively); only acritarch abundances vary sharply, i.e. up to 10% in the lowest horizon, decreasing upwards (Fig. 3). Among the pteridophytic spores, the relative abundances in descending order are as follows (the species in brackets indicate the principal species in a given genus): Limatulasporites (L. fossulatus) 8–11% (10% average); Kraeuselisporites (K.

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Table 2 Spore–pollen taxa and their relative abundance recorded in the upper part of the Guodikeng Formation and basal part of the Jiucaiyuan Formation, based on counts of 278, 333, and 300 grains, respectively Abundance of taxa in samples

AEA 749

Leiotriletes exiguus L. turgidus Cyathidites cf. C. breviradiatus Calamospora breviradiata Punctatisporites cf. P. asperatus P. sp. Granulatisporites sp. Cycloverrutriletes sp. Verrucosisporites jonkeri V. minicus V. sp. Acanthotriletes rostratus f. subrotundus Apiculatisporis cf. A. subtilis Apiculatasporites sp. A. spiniger Anapiculatisporites decorus sp. nov. Baculatisporites uniformis sp. nov. Dictyotriletes sp. Microreticulatisproites sp. Reticulitriletes cf. R. lucidus Retitriletes? sp. Nevesisporites sp. Lycopodiumsporites sp. Limatulasporites fossulatus L. limatulus Annulispora folliculosa Stenozonotriletes sp. Cingulizonates sp. Densosporites cf. D. spongeosus Rotaspora sp. Pterisisporites sp. Kraeuselisporites spinulosus K. varius sp. nov. K.? sp. Lapposisporites echinatus sp. nov. Lundbladispora foveata L. disparilis L. echinata L. sp. A L. sp. B L. sp. C L.? sp. Remysporites dubovii Latosporites sp. Aratrisporites sp. Cordaitina rugulifer C. abutiloides C. gemina Crucisaccites sp. Iunctella sp. Florinites sp. Samoilovitchisaccites? sp.

C

AEA 751

AEA 755

C C C C

C

C C C C C C C C C C

C C

C C CC C C CC

C CC C? C

C CCC C? C

C

C

C C C C

C C CC CC C C C C

C C C C C C

C CC CC CCC CC C C C C C C C C

C

C CC C C CCC C C C C C

C C C? C C C C C

C? C C C

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Table 2 (continued) Abundance of taxa in samples Vesicaspora ex gr. magnalis V. acrifera Voltziaceaesporites heteromorphus Klausipollenites schaubergeri K. angustus sp. nov. Vitreisporites pallidus Pityosporites sp. Platysaccus triassicus P. alatus Falcisporites sublevis Alisporites cf. A. grauvogeliae A. exilis sp. nov. Angustisulcites gorpii Chordasporites rhombiformis Scutasporites cf. S. unicus Lueckisporites virkkiae Lunatisporites leptocorpus L. permotriassicus L. cf. L. hexagonalis L. sp. A Protohaploxypinus latissimus P. cf. P. perfectus P. cf. P. goraiensis P. cf. P. samoilovichiae Striatopodocarpites cf. P. pantii S. sp. Hamiapollenites ruditaeniatus H. bullaeformis Vittatina cf. V. subsaccata V. cf. V. striata V. sp. Cycadopites caperatus C. retroflexus C. sp. Decussatisporites? sp. Eucommiidites sp. Pilasporites trigonius P. perreticulatus sp. nov. Solisphaeridium? sp. Tympanicysta stoschiana

AEA 749

C CCC C C C CCC C C C C C C

AEA 751

AEA 755

C C C C C C C

CC C C C

C CCC CC C C C C C C C

C

C

C

C C C C C C C C C C

C

C C C C

C C C

C C C

C

CC

C

CC

The following symbols are used for relative abundances:

C CCC CC C C C C

C C

C C C CC C C

D absent; C D 1–2%; CC D 2–5%; CCC D 6–10%.

varius) 5–15% (9% average); Lundbladispora (L. foveota) 4–10% (6.5% average); Anapiculatisporites (A. decorus) 1–8% (4% average); Verrucosisporites (V. jonkeri) 2–6% (3.4% average); and Apiculatisporis (A. spiniger) 1.2–4% (2.2% average). Others are only occasionally observed. Aratrisporites sp., occurs in all the three horizons, albeit sporadically; and Kraeuselisporites spinulosus reaches 3.6% in AEA 749, but decreases markedly upward.

Of the gymnospermous pollen, monosaccate (Cordaitina, Florinites?) and monosulcate forms (Cycadopites, Eucommiidites) are both very low in numbers, whereas non-striate bisaccate forms are dominant, ca. 26–30% (28% average for the total assemblage), principal genera being Falcisporites (F. sublevis) 7–14% (10% average); Klausipollenites 3–7% (5.6% average); Alisporites 2–3%; and Vitreisporites 16% (3% average). Striate bisaccate

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forms are low in numbers totalling 5–9% (6.5% average). Protohaploxypinus, Lunatisporites, Striatopodocarpites, Hamiapollenites and Vittatina, do not exceed 1–1.5%, respectively. Lueckisporites virkkiae was only seen as single specimens in the lower and middle horizons. It is noteworthy that in addition to Tympanicysta stoschiana, some specimens of Pilasporites (1.2–4%) have been observed in the three horizons, and individual specimens of a spinate acritarch (Solisphaeridium? sp.) are found from the basal part of the Jiucaiyuan Formation (AEA 755). The main taxa and their percentages are shown in Fig. 3. These results are basically the same as those reported by Qu and Wang (1986), and Hou and Wang (1986), except that in the latter the striate bisaccate forms are slightly more numerous (9–10%).

5. Discussion 5.1. Vegetation, paleoenvironment and climate According to the present knowledge of in situ miospores (e.g. Potonie´ , 1962; Traverse, 1988; Balme, 1995), the vegetation as reflected by the palynoflora is characterized by dominant and diverse pteridophytes and subordinate but almost equally diverse conifers and pteridosperms, together with a few ginkgoaleans, cycadaleans, and possibly bryophytes. Among the pteridophytes, sphenophyllids–equisetaleans (represented by Calamospora) appear to have declined from the Permian, and Paleozoic lycopods (represented possibly by Densosporites and Cingulizonates) occur merely as relicts; on the other hand, the newly evolved Mesozoic herbaceous or shrubby lycopods (producing Lundbladispora, Kraeuselisporites, Aratrisporites) play important roles for the first time. Plants of Cordaitales, if not extinct, are much reduced. Trees of conifers and shrubs of pteridosperms constitute jungles or thin forest as discussed in a later section. Both the Guodikeng Formation and Jiucaiyuan Formation are mostly composed of argillaceous and arenaceous clastic rocks with only occasional lenticular marl horizons and pebble conglomerates. Sedimentology and paleontology suggest shallow water

lacustrine environments alternating with flood plain, river bank and paludal environments (Li et al., 1986). The vegetation possibly covers two main ecosystems: (1) one characterized by lowland environments (flood plains, riparian belts and marshes) containing lycopods and other pteridophytes in association with some hydrophilous–mesophilous pteridosperms and other plants. The assumed presence of such lycopods as Pleuromeia (producing Lundbladispora) may be interpreted as physiologically xerophilous plants in riparian niches. (2) The other ecosystem might have been in upland environments (upland levees, drier parts of floodplains, extrabasinal areas including mountainous regions) which were mainly inhabited by conifers, most of the pteridosperms, and other xerophilous–mesophilous plants. Additionally, in the lakes there were conchostracans, ostracods and a few gastropods — all being shallow water benthonic small crustaceans and molluscs — as well as algal phytoplankton. The presence of spinate acritarchs in the basal part of the Jiucaiyuan Formation may indicate possible marine or brackish water conditions. Paleomagnetic studies suggest that the Tianshan Block was just a little north of the equator, Urumqi being between 16.8 and 15.2ºN during the Late Permian and Early Triassic (Li et al., 1989). Ustritsky (1973) advocated that the north pole was located in the Tunguska Basin (near the outlet of the Lena River) during the Late Permian. The common presence of some palynomorphs both in the Lower Triassic Junggar Basin and in those from Siberia (including Tunguska), suggests that the locality under study was most likely within the subtropical belt. The rather high ratio of spores:pollen appears to indicate a warm and humid climate, at least for the time during which the spore-bearing beds accumulated. The coexistence of striate pollen does not contradict this inference because the parent plants might have been growing in localized drier, but further evidence is required for confirmation. 5.2. The age of the upper part of the Guodikeng Formation and the Lower Triassic boundary Several lines of evidence have demonstrated that the Jiucaiyuan Formation is lower Lower Triassic and the lower part of the Guodikeng Formation is late Late Permian in age (Li et al., 1986; Wu, 1989; cf.

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Table 1). The age of the upper part of the Guodikeng Formation, however, deserves further discussion. The palynology of samples AEA 749 and 751 underlying the Lystrosaurus–Dicynodontia-bearing uppermost part of the Guodikeng Formation is of significance in this regard. The miospore assemblages from these two samples are largely the same in general composition and content, and are also closely similar to that from the overlying Jiucaiyuan Formation (AEA 755) (see Table 2; Fig. 3). Compared with the spore assemblages known from the Lower Triassic in this area (Qu and Wang, 1986, 1990), they contain a number of Triassic, particularly Early Triassic species. Identical or closely comparable species to those known from the Jiucaiyuan Formation include Cyathidites cf. C. breviradiatus, Verrucosisporites jonkeri, Apiculatisporis spiniger, Dictyotriletes mediocris, Nevesisporites sp., Pterisisporites sp., Lundbladispora disparilis, Aratrisporites sp., Klausipollenites angustus and Pilasporites trigonius in addition to Lueckisporites virkkiae, Klausipollenites schaubergeri, Limatulasporites fossulatus, L. limatulus, Densosporites cf. D. spongeosus, Striatopodocarpites cf. S. pantii and Hamiapollenites bullaeformis. The following occur in the uppermost part of the Guodikeng Formation and in the upper Lower Triassic Shaofanggou Formation: Verrucosisporites minicus, Apiculatasporites sp., Dictyotriletes sp., Annulispora follioculosa, Lundbladispora foveota, Lapposisporites sp., Lunatisporites leptocorpus and Lunatisporites cf. L. hexagonalis. Such similarity and other considerations discussed in the following sections lead us to conclude that the upper part (about 50 m in this section) of the Guodikeng Formation is lower Lower Triassic although its palynoflora displays a Permian– Triassic transitional nature. According to Li et al. (1986), the upper part (about 30 m in thickness) of the Guodikeng Formation represents “a Permian–Triassic biotic transitional bed”. Further, they state that “the lithological boundary does not coincide with the biostratigraphic boundary of the Permian and Triassic, and this confirms that the biostratigraphic boundary between the two Systems is lower than the lithological boundary between the Guodikeng and the Jiucaiyuan Formation”. This is based on their study of palynological assemblages from the upper part (horizon 43, ca. 30

11

m apart from the top) of the Guodikeng Formation, in which “still occur some Late Permian elements, such as H. bullaeformis, L. virkkiae and L. singhii, but which do not occupy a dominant position. However, a large number of typical Triassic forms, such as Lundbladispora and Taeniaesporites occur which compare closely to assemblages from the Jiucaiyuan Formation. In the same horizon there are fossil ostracods represented by Darwinula pararotunda and D. paraquanzijiensis rather than the typical Permian genera Panxiania and Vymella. This reflects a transitional ostracod fauna towards the assemblages in the lower part of Jiucaiyuan Formation which comprises populations of individually small forms of Darwinula triassiana and D. rotundata. However, in the upper–middle part of the upper Guodikeng transitional bed, Dicynodon of Permian aspect possibly occurs in association with Lystrosaurus”, Li et al. thus concluded: “A broad review of the fossil data available, and the disappearance (extinction) of ancient genera and species as well as the appearance of newly arising forms suggest that the remarkable biotic replacement seems to be beneath horizon 43 in this section. Of course, in order to demarcate a unified and reliable biostratigraphic boundary between the terrestrial Permian and Triassic, a detailed and comprehensive investigation of the transitional sequence is still needed” (translated from the Chinese). Wu (1989) further demarcated the lower boundary of the Lower Triassic at the base of horizon 40 (ca. 45 m apart from the top) based on a comprehensive review of the known fossil groups. Liu (1989) suggests that the conchostracans from the middle part of the Guodikeng Formation (horizon 34) are Early Triassic in age. However, the associated presence of Zamiopteris cf. Z. glossopteroides, Walchia sp. and Samaropsis sp. indicates a probable Late Permian age. The present paper considers that the Permian– Triassic boundary should be drawn at the base of the Unit 3 (as shown in Fig. 2) which is about 20 m lower than the base of Li, Zhang and Wu’s horizon 43, and possibly equivalent to or a little lower than the base of their horizon 40 according to comparisons of the figures and topography. The reasons are as follows: (1) The palynological assemblage from sample AEA 749 (a little lower than Li et al.’s horizon 43), is

12

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similar to those from horizons AEA 751 and 755. All three belong to the Lundbladispora–Lunatispories– Aratrisporites Assemblage, and are noticeably different from the underlying assemblage (Li et al.’s horizon 36) in the middle–upper part of the Guodikeng Formation which is characterized by a remarkable dominance of gymnosperms and was named the Kraeuselisporites–Lueckiporites Assemblage (Hou and Wang, 1986). Considering the fact that the genus Kraeuselisporites remains abundant in the overlying Lower Triassic, it would be preferable to rename it the Klausipollenites schaubergeri–Lueckiporites virkkiae Assemblage for, although these two species do also occur in the earliest Triassic, they do so only sporadically or not as abundantly as in the Permian. (2) The first appearance of Aratrisporites sp. in AEA 749 is particularly important because this is just above Li et al.’s horizon 40; the species continues upwards in AEA 751 (the upper part of the Guodikeng Formation) and AEA 755 (the base of the Juicaiyuan Formation). Aratrisporites is considered to be an important index for the Triassic. For example, its first appearance in the Permian–Triassic sections in Changxing, Zhejiang (Ouyang and Utting, 1990) is just 2.7 m above the Triassic base in which occur Hypophiceras changxingensis and Otoceras? sp. in the Griesbachian Chinglung Formation. In AEA 749 there are other Triassic index forms such as Apiculatisporites spiniger, Remysporites dubovii, Lundbladispora spp., Lapposisporites echinatus and Angustisulcites gorpii. Some species with a Permian aspect also occur in this horizon. With the exception of several non-striate bisaccate forms (e.g. Falcisporites sublevis), they are all sporadically present representing remnant Permian elements. The present authors consider that the first appearances of newly evolved diagnostic species are more valuable than the remnant presence of ancient forms. (3) Horizon ADI 1110, which is ca. 7 m below sample AEA 749, contains the conchostracans Polygrapta xinjiangensis and Sinolimnadiopsis jimsarensis. The former shows a Permian aspect and the latter, although more common in the Lower Triassic, is also known in the Upper Permian. This suggests that the horizon is Late Permian and not Early Triassic as Liu (1989) advocated. (4) The presence of Dicynodon sinkiangensis in the uppermost part of the Guodikeng Formation may

be a relict species of the Dicynodontia in the Early Triassic because it has been found in association with Lystrosaurus which is a diagnostic Triassic vertebrate widely known in South Africa, Russia and Xinjiang (Li et al., 1986). 5.3. Transitional nature of the palynoflora The presence of so many taxa of Permian aspect in the Lower Triassic raises the question whether they are reworked. This is considered highly unlikely for the following reasons: (1) the sedimentation of the Permian–Triassic strata is fundamentally continuous. Therefore, the likelihood of palynomorph reworking is minimized (Boulter et al., 1988); (2) among the taxa with an older aspect, Densosporites, Cingulizonates and Cordaitina are most likely to be interpreted as reworked, but the presence of Lepidodendron? or Viatcheslavia in the basal part of the Jiucaiyuan Formation and the fact that Cordaitina-type pollen reach more than 10% in some Early Triassic assemblages from Siberia (Kara-Murza, 1960) support the interpretation of their presence in the assemblage as remnant forms; (3) palynomorphs with a more mature colour (yellow-brown) are mainly represented by striate bisaccate pollen derived from peltaspermaceous and advanced conifers that also range into the lower Mesozoic (Triassic); their maturation colour is not significantly different from those of younger aspect. Among the 88 species of spores and pollen recorded in the present paper, there are about 40 species that display a Permian aspect. More than 10 species of pteridophytic spores appear to have extended upward from the Permian or older, such as Leiotriletes exiguus, L. turgidus, Calamospora breviradiata, Verrucosisporites jonkeri, Acanthotriletes rostratus f. subrotundus, Rotaspora sp., Latosporites sp., Densosporites cf. D. spongeosus, and Cingulizonates sp. Most of these are of uncertain botanical affinity. However, Densosporites, according to in situ spore data (Balme, 1995), is probably derived from Paleozoic Selaginellales or Isoetales. Twenty-six of the species of gymnospermous pollen (a little more than half of all species) have a Permian aspect, such as Cordaitina (3 species), Crucisaccites sp., Vesicaspora (2 spp.), Scutasporites cf. S. unicus, Vittatina (3 spp.), Hamiapollenites (2

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spp.), Cycadopites (2 spp.), Chordasporites rhombiformis, Protohaploxypinus (3 spp.), Lunatisporites permotriassicus, Klausipollenites schaubergeri, Alisporites cf. A. grauvogeliae, Falcisporites sublevis, Platysaccus alatus, Striatopodocarpites cf. S. pantii, Lueckisporites virkkiae, Vittatina cf. V. subsaccata and Protohaploxypinus latissimus. Of these, L. virkkiae is one of the most important indexes of the Late Permian. It is known in situ in some Permian fossil conifers, e.g. the Majonicaceae (Majonica) from European floras (Clement-Westerhof, 1988) and Sashinia (D Quadrocladus) from Angara floras (Meyen, 1982). The similar form Scutasporites is also known from the Permian conifer Dvinostrobus of the Angara flora (Meyen, 1988). Vittatina and Protohaploxypinus s.l. type pollen have been linked to the peltaspermaceous plants which were widely distributed in the Permian of the Northern Hemisphere, although similar Protohaploxypinus-type pollen are known from Glossopterids (Arberiales) and the Triassic conifer Rissikia of Gondwanaland (Meyen, 1988). Vesicaspora, as a form genus, has a long stratigraphic range, mainly from Late Carboniferous and Permian. Morphologically comparable in situ pollen grains have been reported from the Late Carboniferous–Permian pteridosperms such as Callistophytales (Callandrium and Callistophyton) (Taylor, 1988) and the Permian pteridosperm family Cardiolepidaceae (Cardiolepis D Phylladoderma, Permotheca). According to Meyen (1988), the parent plants of Cordaitina might be related to Cordaianthales (Rufloriaceae and Vojnovskyaceae) of the Angara Province, while Florinites is related to the Cordaitales (e.g. Cordaianthus) of the Euramerican Province (Balme, 1995). All these plants are basically representatives of the late Paleozoic. Lunatisporites is mainly recorded from the Triassic, but a few species are known in the Permian. L. permotriassicus, first occurs in the Permian– Triassic transition in western Siberia and subsequently in northern Xinjiang. Similar species were already present in the Upper Permian of the northern European part of USSR. Clement-Westerhof (1974) found Lunatisporites-type in situ pollen in a Late Permian conifer from Italy. Two species of Cycadopites, C. caperatus and C. retroflexus, are commonly seen in the Permian assemblages of the Angara and Subangara

13

provinces. Cycadopites may have been derived from ginkgoaleans, cycadaleans, and peltaspermaleans (Balme, 1995) which, although more important in the Mesozoic, had evolved as early as the Carboniferous. Judging from known stratigraphic ranges and botanical comparisons, the palynological evidence discussed above suggests extensions of ranges from Permian or older across the Permian–Triassic boundary. The palynofloras with Paleophytic aspect may include Lycopsida (Densosporites, Cingulizonates), Sphenopsida (Calamospora), and some ancient Filices although these plants did not play important roles. Plant taxa extending from the Permian into Triassic appear mainly to be of gymnospermous origin, including ginkgoaleans, cydadaleans, conifers and pteridosperms, particularly peltaspermaleans. The conifers are largely represented by non-striate bisaccate pollen (partly Falcisporites, Pityosporites, Platysaccus and Klausipollenites) and some striate– taeniate bisaccate forms (e.g. Lueckisporites, Scutatasporites and Lunatisporites). In the palynofloras from the uppermost Guodikeng and basal Jiucaiyuan formations, the taxa with a Mesozoic — notably Triassic — aspect include Aratrisporites sp., Lundbladispora (6– 7 species, especially L. echinata and L. foveota), Remysporites dubovii, Voltziaceaesporites heteromorphus and Angustisulcites gorpii. In addition, Kraeuselisporites (3 spp.), Lapposisporites, Lunatisporites (3 spp.) and Eucommiidites sp., are largely Mesozoic elements. Aratrisporites is derived probably from such plants as Lycostrobus and Cylostrobus (Potonie´, 1962; Helby and Martin, 1965; Retallack, 1975) as well as possibly Annalepis (e.g. Wang and Wang, 1990), all belonging to Mesozoic lycopods. Lundbladispora (and possibly Kraeuselisporites) may be related to Pleuromeia, another diagnostic plant genus of Triassic lycopods. Voltziaceaesporites, as known in Europe and northern China, is often found in association with the Triassic conifer cone Willsiostrobus (Traverse, 1988), and Eucommiidites-type in situ pollen is known to be associated with probable Mesozoic cycads (Balme, 1995). There are about 20 species which are of Mesozoic aspect. Among the 8 species newly proposed and some undeterminable species, some might prove to be restricted to the Mesozoic. Therefore, elements

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with a Mesozoic aspect represent nearly one third of the entire palynoflora. Thus it appears highly probable that the Early Triassic palynoflora in northern Xinjiang represents a Permian–Triassic transitional or ‘mixed’ flora. In other words, the palynoflora contains a significant number of species with a Paleozoic aspect and a lesser number (ca. 1=3) of species of typical Mesozoic aspect. Similar transitional Early Triassic palynofloras have been known from different phytogeographical provinces. For instance, the lower Scythian Kayitou Formation in eastern Yunnan (Ouyang and Li, 1980; Ouyang, 1986); the lower Griesbachian Lower Chinglung Formation in Changxing, eastern China (Ouyang and Utting, 1990); the earliest Griesbachian ‘Protohaploxypinus Association’ in East Greenland (Balme, 1980); the lower Induan Subbasalt Formation in the Urals (Tuzhikova, 1985); the lower Maltsevsky Formation in Kuzbass (Kurbatova, 1966) in Russia; and to a lesser degree in the Lower Triassic strata of Central Europe (Freudenthal, 1964) as well as in the Griesbachian Bjorne Formation, Sverdrup Basin of the Canadian Arctic Archipelago (Utting, 1985, 1989). Furthermore, in Gondwanaland, to judge from the macrofossil record, the Early Triassic vegetation also contains many elements of the Glossopteris flora (Pant, 1987). Thus the palynological evidence available does not lend support to the view that the plant kingdom suffered a sudden catastrophic mass extinction at the close of the Permian (e.g. Wang, 1989). Ouyang (1991) has discussed in some detail the palaobotanical and ecological implications of these earliest Triassic transitional palynofloras. Based on the above discussion, the Junggar palynofloras from the upper part of the Guodikeng Formation and the Jiucaiyuan Formation should be early Early Triassic (early Induan) in age. Another important piece of evidence which lends support to such dating is that the overlying Shaofanggou Formation (ca. 260 m thick) in the same section, according to Qu and Wang (1986), yields a similar palynoflora, also dominated by pteridophytic spores (57% in average) with subordinate gymnospermous pollen (43%) but it shows some noticeable differences compared with the present one. The principal genera it contains are Polycingulatisporites, Annulispora, Limatulasporites, Dictyotriletes, Lundbladispora and Lu-

natisporites. Other differences are (1) the decrease of non-striate bisaccate pollen (6.5%), (2) the increase of striate bisaccate pollen (16%) and lycopod spores (Lundbladispora and Aratrisporites), and (3) the essential disappearance of Lueckisporites and Hamiapollenites. Important species for dating the Shaofanggou palynoflora include Aratrisporites parvispinosus Leschik, Lundbladispora obsoleta Balme, Kraeuselisporites spinosus Jansonius, Lunatisporites novimundi (Jansonius) and L. noviaulensis (Leschik). The present authors are in agreement with Qu and Wang in dating the Shaofanggou assemblage as late Early Triassic (late Induan). 5.4. Phytogeographical provinciality Very early Triassic macrofossil plant records so far reported are relatively meager (Barnard, 1973; Wang, 1985), and floral provincial differentiation has only been approximately assessed (Zhou and Li, 1979). Further difficulties stem from paleopalynology itself due to the fact that most parent plants of dispersed miospores are unknown, thus making the restoration of the vegetation and paleoecology very difficult; and that generally we do not know to what extent a palynological form species or form genus represents a natural taxon. Thus, palynological distribution patterns do not necessarily represent special ecological habitats as do natural taxa. For example, Lundbladispora nejburgii is known in the Lower Triassic of many localities in the Northern Hemisphere including the Jiucaiyuan Formation, Xinjiang (see Qu and Wang, 1986). It has some connection with Pleuromeia (P. rossica Neuburg) or Pleuromeiaceae (Kiuntzel, 1966; Balme, 1995). However, as in the Triassic Liujiagou Formation of Shanxi, northern China at the same locality, P. nejburgii occurs (Qu, 1982) while Pleuromeia is represented by P. jiaochengensis Wang et Wang, 1982 instead of P. rossica, so it would be unwise to emphasize its phytogeographical significance. The same can be applied to Lueckisporites virkkiae. Finally, it is worth noting that Early Triassic palynological data indicate that the phytogeographical provincialism was less well marked than in the late Paleozoic. This is due in part to increasing continentality from the tectonic assembly of Pangea and in part to a possibly less steep climatic gradient, as evidenced

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by the almost global distribution of some genera in this epoch, such as Aratrisporites, Lundbladispora and Lunatisporites as well as some common forms of acanthomorphic acritarchs. It follows, as Balme (1970) pointed out, that “many, if not most, fossil spores and pollen grains have dubious value as guides to the plant geography of the past”, and that “well-defined form species, combining unusual morphological characters, are likely to prove the most satisfactory units in studying the broad distributive patterns of past floras”. This is also the reason that in the delimitation of most species, in accordance with Halle (1927), we use closely circumscribed species to minimize “confusion and hasty conclusions in both taxonomic and phytogeographic matters”. In spite of the above-mentioned difficulties, provincial comparisons of the miospores identified will be attempted below. The data are summarized in Table 3. It is worth emphasizing that a given species (or genus) first reported in a special phytogeographical province is not necessarily diagnostic of the province. For example, Zhang (1983) cited Crucisaccites Lele et Maithy, 1964 (with the type species from the Lower Permian of Gondwanaland) as “a Gondwana palynological genus” but it occurs in both the Upper Carboniferous and Permian in northern Xinjiang (Ouyang et al., 1993), and comparable specimens are also reported from the Upper Carboniferous and Lower Permian in the Tunguska Basin (e.g. Medvedeva, 1960). 5.4.1. Comparison with the Angara and Subangara provinces The Junggar earliest Triassic palynoflora contains a large number of species which were first recorded from the Angara and Subangara provinces: Leiotriletes turgidus, Punctatisporites cf. P. asperatus, Acanthotriletes rostratus f. subrotundus, Dictyotriletes mediocris (D Reticulina rucida KaraMurza), Remysporites dubovii, Cordaitina rugulifera, C. abutiloides, C. gemina, Iunctella sp., Vesicaspora ex gr. magnalis, V. acrifera, Platysaccus alatus, Falcisporites sublevis, Lunatisporites permotriassicus, Protohaploxypinus latissimus, P. cf. P. perfectus, Hamiapollenites bullaeformis, Vittatina cf. V. subsaccata, V. cf. V. striata, Cycadopites caperatus, C. retroflexus and Pilasporites trigonius. These represent 22 species which are identical or closely

15

comparable to those from the Permian or Triassic strata of the Angara and Subangara provinces as defined by Meyen (1982) and others. In addition, the following 15 species of the Junggar palynoflora first described from other major provinces are also closely comparable to species in Angaraland: Cyathidites cf. C. breviradiatus, Verrucosisporites jonkeri, Acanthotriletes junggarensis, Apiculatasporites sp., Limatulasporites fossulatus, L. limatulus, Lundbladispora foveota, Klausipollenites schaubergeri, Alisporites cf. A. grauvogeiliae, Platysaccus triassicus, Lueckisporites virkkiae, Lunatisporites triassicus, L. cf. L. hexagonalis, Protohaploxypinus cf. P. samoilovichiae and Striatopodocarpites cf. S. pantii. Therefore (together with the acritarch Tympanicysta stoschiana), there are a total of 38 species (about 40% of the total) in northern Xinjiang that have direct or indirect connections with the Permian and Triassic floras of Angara. Some species with simple morphology (e.g. Leiotriletes turgidus), and some with a wider distribution in the Northern Hemisphere (e.g. L. virkkiae and H. bullaeformis), are less significant phytogeographically. More specific comparisons follow: (1) The Junggar palynoflora has certain similarities with those from the Permian strata in the Kuznetsk Basin (e.g. Andreeva et al., 1956) as shown by the following common species: Leiotriletes turgidus (D L. urbanus Andreeva), Limatulasporites fossulatus, Cordaitina rugulifera, C. abutiloides, C. gemina, Vesicaspora acrifera, V. ex gr. magnalis, Platysaccus triassicus (possibly Coniferaletes imperspicuus Andreeva), Cycadopites caperatus and C. retroflexus. Among these, the Cordaitina species deserve special attention because they are more common in the Siberian region than in other areas and they have been recorded by Liuber and Val’ts (1941) from the Kuznetsk Basin and Pechora Basin and by Medvedeva (1960) in the Tunguska Basin. (2) The Junggar palynoflora has some species in common with those from the Permian, especially Upper Permian in the Taymir Basin (Kara-Murza, 1952): Leiotriletes turgidus, Dictyotriletes mediocris, Cordaitina abutiloides (possibly D Circella rotata f. arctica Kara-Murza), Iunctella sp., Protohaploxypinus perfectus and Cycadopites retroflexus. The Kuznetsk, Tunguska, Taymir and Pechora basins all belong to the Warm Temperate Climate Province

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Table 3 Known phytoprovincial and stratigraphic distributions of some spore and pollen species occurring in the Lower Triassic Jiucaiyuan Formation and the upper part of the Guodikeng Formation of northern Xinjiang Taxa

Phytochoria:

Angara

Region:

K–T

Age:

(P)

(T)

C

C C C C C

Leiotriletes turgidus Punctatisporites cf. P. asperatus Acanthotriletes rostratus f. subrotundus Dictyotriletes mediocris Cordaitina abutiloides C. gemina Iunctella sp. Vesicaspora ex gr. magnalis V. acrifera Apiculatisporis cf. subtilis Apiculatasporites labrosus Lundbladispora foveota Cordaitina rugulifera Platysaccus alatus Lunatisporites permotriassicus Protohaploxypinus cf. P. perfectus Cycadopites retroflexus Verrucosisporites jonkeri Cycadopites caperatus Remysporites dubovii Protohaploxypinus latissimus Hamiapollenites bullaeformis Pilasporites trigonius Vittatina cf. V. subsaccata Lunatisporites cf. L. hexagonalis Protohaploxypinus cf. P. samoilovichiae Lapposisporites Falcisporites sublevis Vittatina cf. V. striata Klausipollenites schaubergeri Alisporites cf. grauvogeliae Cingulizonates Densosporites cf. D. spongeosus Apiculatisporites spiniger Lundbladispora echinata Calamospora breviradiata Scutasporites cf. S. unicus Voltziaceaesporites heteromorphus Angustisulcites gorpii Leiotriletes exiguus Chordasporites rhombiformis Lundbladispora subornata Polycingulatisporites rhytismoides Lueckisporites virkkiae Tympanicysta stoschiana Limatulasporites fossulatus L. limatulus Cyathidites cf. C. breviradiatus Striatopodocarpites cf. S. pantii Protohaploxypinus cf. P. goraiensis

C C

Subangara K–T (P)

Euramerica

Cathaysia

Gondwana

(P)

(P)

(T)

(P)

(T)

C

C

C

C C C C C

U and R (T)

(P)

(T)

(T)

C

C C C C C C C C

C

C C

C C

C C C C C

C C

C C C C C C

C C

C C C C C C C C C C C C C C C

C

C

C C

C C C C C

C C

C C C

C C

C C

C C C C

C C C C

C C C C C

C C C

C C C

C C C C

C C

C C C

C C

C C C C C C C C C

C C

C C

C

C C

Abbreviations in column headers are as follows: K–T D Kuznetsk–Tunguska; T–Y D Taymir–Yakutski; U and R D Urals and Russian Platform. P D Permian (mainly Late Permian); T D Triassic (mainly Early Triassic).

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(D Angara Botanic Province) dominated by a cordaitalean vegetation during the Permian as defined by Hart (1970). The present palynoflora displays a close relationship with those of the Kuznetsk and Taymir basins which belong to the Taymir–Kuznetsk subprovince of the Angara Province (Meyen, 1982). (3) The Junggar palynoflora contain the following species which have been recorded in the Permian (mainly Upper Permian) from the Russian platform or the European part of the former USSR (Liuber and Val’ts, 1941; Samoilovich, 1953; Zoricheva and Sedova, 1954; Varjukhina, 1971; Molin and Koloda, 1972; Tuzhikova, 1985): Limatulasporites limatulus, Cordaitina rugulifera, Platysaccus alatus, Falcisporites sublevis, Alisporites cf. A. grauvogeliae, Vesicaspora sp. ex. gr. V. magnalis, Scutasporites cf. S. unicus, Lunatisporites cf. L. hexagonalis, L. permotriassicus, Protohaploxypinus latissimus, P. cf. P. perfectus, Hamiapollenites bullaeformis, Vittatina cf. V. subsaccata, V. striata, Cycadopites caperatus, C. retroflexus and Pilasporites trigonius. From this list it is clear that the Early Triassic palynoflora of northern Xinjiang bears closer relationship in pollen composition to those of the Permian strata in the European part of the former USSR, although some of the species (e.g. L. virkkiae) have also been reported sporadically in the Siberian area in the upper ‘Korvunchansky’ Formation (Lower Triassic) of the Tunguska Basin (Meyen, 1982, p. 76). However, gymnospermous dominance typically with bisaccate pollen (especially striate forms) is more diagnostic in European Russia which belongs to the Permian ‘Hot Dry Climate Province’ of Hart (1970) and which was dominated by xerophytic plants, or in the Subangara Province of Meyen (1982). (4) Comparison of the palynofloras with those from the Early Triassic indicates that the Junggar palynoflora contains some elements in common with those of Yakutsk (Fradkina, 1967), the Subbasalt Formation in the Urals (Tuzhikova, 1985) and especially with Assemblage I (Induan) in the Moscow Syncline or the Timano-Pechora area (Yaroshenko and Golubeva, 1981). The common species for each area are as follows (see synonyms in the taxonomic section): — For the Yakutsk assemblage: Cyathidites sp. cf. C. breviradiatus, Verrucosisporites minicus, Acanthotriletes junggarensis, Apiculatasporites sp.,

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Dictyotriletes mediocris, Limatulasporites fossulatus, Lundbladispora foveota and Platysaccus alatus. — For the Subbasalt Formation: Verrucosisporites jonkeri, Limatulasporites fossulatus, Remysporites dubovii, Klausipollenites schaubergeri, Alisporites cf. A. grauvogeliae, Lueckisporites virkkiae, Lunatisporites cf. L. hexagonalis, L. cf. L. puntii, Protohaploxypinus cf. P. samoilovichiae, Vittatina cf. V. striata, V. subsaccata, and Pilasporites trigonius. Other common genera include Aratrisporites, Lundbladispora, Kraeuselisporites and Hamiapollenites. — A comparison with the Induan assemblage of the Moscow Syncline is very difficult because no systematic descriptions of the palynomorphs (Yaroshenko and Golubeva, 1981) were given and only a few specimens were illustrated by Kiuntzel (1966). However, from the taxa listed by these authors, it appears that the general features are very similar to the present assemblage. The common genera are: Leiotriletes, Cyathidites, Punctatisporites, Lundbladispora, Kraeuselisporites, Limatulasporites, ‘Reticulatisporites’, Aratrisporites, Cordaitina (rare), Alisporites, Platysaccus, Klausipollenites, Lueckisporites, Lunatisporites, Protohaploxypinus, Striatopodocarpites, Vittatina and Cycadopites. Identified species include Limatulasporites limatulus, L. virkkiae, Protohaploxypinus samoilovichiae, Klausipollenites schaubergeri, Tympanicysta stoschiana and possibly Verrucosisporites jonkeri (see Kiuntzel, 1966, p. 1, pl. I, fig. 7). Assemblage I of the Moscow Syncline is characterized by the abundance of bisaccate Striatiti (especially Lunatisporites), Lundbladispora, Kraeuselisporites, bisaccate non-Striatiti and occurrences of Limatulasporites and Cycadopites. With the exception of the higher percentage of bisaccate Striatiti, these features are largely the same as in our assemblage. In summary, the Junggar palynoflora from northern Xinjiang displays a close relation with the Permian–Triassic transitional palynofloras from both the Angara and Subangara provinces. In particular, in spore composition it is highly similar to the Angara Province. On the other hand, in gymnospermous pollen composition it shows a much closer relation with the Subangara Province, especially the ancient conifers (partly bisaccate non-Striatiti and Striatiti, e.g. Platysaccus and Lueckisporites) and pteridosperms (e.g. Protohaploxypinus, Vittatina) as

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well as in the moderate abundance of Lundbladispora and Kraeuselisporites. Palynological distinctions between the two provinces during the Late Permian and Early Triassic is not as great as that in the earlier part of the late Paleozoic, as evidenced by some common species. This is also in agreement with the macrofossil record because as Meyen (1982, p. 76) pointed out, the Permian–Triassic floras “may receive the epithet ‘Angara’ only due to their geographical position. Otherwise they are basically different from older Permian floras and commence a new phase in the floral history of Eurasia, when hitherto existing barriers had been broken and new migration routes appeared”. In delineating phytogeographical provinces, spores of heterosporous plants and the pollen of seed plants are highly significance. Thus, the early Early Triassic vegetation in northern Xinjiang, although containing characteristic spores of the Angara Province, bears closer resemblance to that of European Russia (roughly equivalent to the Subangara Province) for the rather abundant or diverse presence of heterosporous plants (producing Aratrisporites, Lundbladispora and Kraeuselisporites) and seed plants (conifers and pteridosperms, producing non-striate and striate bisaccate pollen). In other words, northern Xinjiang might have belonged to the Subangara Province in Meyen’s sense, or Sun’s (Sun, 1989) zone III of the Angara Province, indicating that the phytogeographical characters were largely the same and not as distinctly differentiated as in the Late Permian. 5.4.2. Comparison with the Permian–Triassic palynofloras of the Euramerican Province The present palynoflora contains some Permian or pre-Permian elements which were first recorded from the Euramerican Province, such as Calamospora cf. C. breviradiata, Densosporites cf. D. spongeosus, Klausipollenites schaubergeri, Lueckisporites virkkiae and Scutasporites cf. S. unicus, as well as Cingulizonates sp. and Rotaspora sp., although these are not restricted to the Province. Closely comparable or identical species with those first reported from the Triassic of the Euramerican Province include Verrucosisporites jonkeri, Apiculatisporites spiniger, Lundbladispora echinata, Voltziaceaesporites heteromorphus, Platysaccus tri-

assicus, Alisporites cf. A. grauvogeliae, Angustisulcites gorpii, Lunatisporites cf. L. hexagonalis, and Striatopodocarpites cf. S. pantii. In addition, some species first recorded from other provinces also occur in the Triassic of the Euramerican Province, such as Limatulasporites fossulatus (possibly D Dulhuntyspora? minuta Jansonius), or in the Permian and Triassic, such as Protohaploxypinus cf. P. samoilovichiae, Hamiapollenites bullaeformis and Vittatina cf. V. striata. In spite of the fact that the species listed above are mostly form species (Potonie´ and Klaus, 1954; Potonie´ and Kremp, 1955; Butterworth and Williams, 1958; Klaus, 1963; Barss, 1967; Visscher, 1973; Balme, 1980; Dybova-Jachowicz, 1981), and some are not restricted to the Euramerican Province, the comparison reveals that the Junggar palynoflora appears to have important connections with the Permian, and particularly the Triassic floras of the Euramerican Province. The relationship is not, however, as strong as with the Angara and Subangara provinces. In this regard, the presence of identical or closely comparable genera and species to those from the Upper Bunter (Visscher, 1966) of the Netherlands is worthy of notice, namely Lapposisporites, Lundbladispora, Kraeuselisporites, Voltziaceaesporites heteromorphus, Alisporites cf. A. grauvogeliae, and Angustisulcites gorpii. However, due to the presence of some elements with a Paleozoic affinity (e.g. Densosporites and Lueckisporites virkkiae) and the absence of Triadispora (quite diverse in the Bunter assemblage), the northern Xinjiang spore-bearing strata are obviously older than the late Early Triassic Upper Bunter. Yang and Sun (1986) recorded more than 20 species of megaspores from the Jiucaiyuan Formation of the Junggar Basin, referred mainly to Trileites, Pusulosporites, Maexisporites and Aneuletes. They correlated their megaspore assemblages with the Otynisporites eotriassicus zone and Trileites polonicus–Pusulosporites populosus zone from the Lower and Middle Buntsandstein, respectively, in Poland. A majority of species are conspecific with those from Poland, such as T. vulgaris Fuglewicz, P. inflatus Fuglewicz and Otynisporites eotriassicus Fuglewicz. Although their phytogeographical value is unclear due to the lack of widespread studies of Lower Triassic megaspores in

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other provinces, this similarity reinforces the impression that the vegetation in the early Early Triassic in northern Xinjiang had important connections with Europe. 5.4.3. Comparison with the Gondwana Province In addition to the presence of Aratrisporites, Lundbladispora, Kraeuselisporites and Lunatisporites which have almost global distribution in the Early Triassic, the Junggar palynoflora contains several species which were first reported from the Permian in Gondwanaland. These are Protohaploxypinus cf. P. goraiensis and Striatopodocarpites cf. S. pantii (Potonie´ and Lele, 1961; Goubin, 1965; Balme, 1970). Also Cyathidites cf. C. breviradiatus Helby, 1967 was first reported from the Triassic in Gondwanaland. Furthermore, Limatulasporites fossulatus and L. limatulus were also first recorded from the Permian and Triassic of Gondwanaland (Playford, 1965; Balme, 1970; Helby, 1973; Foster, 1979) but they have also been found in the Angara and Subangara provinces. A comparison between the present assemblage and that from the Lower Triassic Lower Mariokeni Formation (Karoo) of Kenya (Hankel, 1990) indicates that there are quite a number of common genera in both assemblages; however, at the species level, only Cyathidites cf. C. breviradiatus, Protohaploxypinus cf. P. samoilovichiae and Striatopodocarpidites cf. S. pantii are the same. Thus it appears that the Early Triassic vegetation in northern Xinjiang seems to have less connections with Gondwana than with the Angara and Euramerican provinces. In dealing with the similarity of Permian (and Early Triassic) miospore assemblages between the northern hemisphere and Gondwanaland, as shown by the common presence of abundant and almost indistinguishable bisaccate striate= taeniate pollen genera, Meyen (1982) assumed that presently we have enough grounds to state that the similarity “... is essentially related to parallelism”. 5.4.4. Comparison with the Cathaysian Province The present palynoflora contains a few species which were first recorded from the Cathaysian Province in the Upper Permian and=or Triassic, namely Leiotriletes exiguus and Chordasporites rhombiformis (Ouyang and Li, 1980; Zhou, 1980; Li, 1988; Ouyang and Utting, 1990). In addition,

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Qu and Wang (1986) reported the presence of Lundbladispora subornata Ouyang et Li and Polycingulatisporites rhytismoides Ouyang et Li in the Jiucaiyuan Formation; both were first recorded from the lower Scythian Kayitou Formation in Yunnan. A number of species first recorded from other provinces also occur in the Permian and=or Triassic in the Cathaysian Province, including: L. virkkiae, K. schaubergeri, F. sublevis, C. caperatus and Tympanjicysta stoschiana in the Permian and Triassic; Angustisulcites gorpii and Voltziaceaesporites hetermorphus in the upper Lower Triassic Heshanggou Formation of Shaanxi (Ouyang and Norris, 1988); and Alisporites cf. A. grauvogeliae and Vittatina striata in the Griesbachian of Zhejiang (Ouyang and Utting, 1990). In the early Griesbachian assemblage of Zhejiang and the early Scythian assemblage in Yunnan, some important genera also occur which have been found in the present palynoflora: Densosporites, Cordaitina, Aratrisporites, Lundbladispora, Lunatisporites and Eucommiidites. However, as a whole, the palynoflora of northern Xinjiang is considerably different from those known in the Lower Triassic of the Cathaysian Province. For instance, the early Scythian assemblage in eastern Yunnan is characterized by (1) the abundant presence of many typical Paleozoic forms, and (2) the rare presence of bisaccate Striatiti. In contrast, the coeval assemblages from northern China (Henan and Shanxi) are characterized firstly by the greater abundance of bisaccate Striatiti (including Lunatisporites), Lundbladispora or Aratrisporites, and Cycadopites; and, secondly, by Paleozoic species, even in the lowermost Triassic, which are less abundant than in Yunnan but more abundant than in northern Xinjiang. This suggests that the climate in northern Xinjiang during the Early Triassic was less humid than in eastern Yunnan (southwest China) and less arid than in northern China. Comparisons between the major provinces are summarized in Table 3.

6. Conclusions Palynomorphs obtained from the upper part of the Guodikeng Formation and the basal part of the Jiucaiyuan Formation in the southern Junggar Basin

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near Jimsar, Xinjiang are very abundant, diverse and well preserved, in contrast to the poor record of macrofossil plants. They provide valuable information for age determination and correlation, reconstruction of paleovegetation, and paleoenvironment. The palynoflora contains both allochthonous and autochthonous or para-autochthonous miospores which might have been derived mainly from two ecosystems, viz. (a) lowland environments (flood plains, riparian belts and marshes) inhabited largely by lycopods and other pteridophytes in association with some hydrophilous–mesophilous pteridosperms and other plants, and (b) upland environments (upland levees, drier parts of floodplains, extrabasinal areas including mountains) inhabited mainly by conifers, most pteridosperms and other xerophilous– mesophilous plants. The higher ratio of spores:pollen and other considerations indicate that the paleoclimate was probably subtropical, warm and humid (at least during sedimentation of the spore-bearing strata) although not as humid as in eastern Yunnan (southwestern China). Furthermore, lacustrine conditions are indicated by occurrences of algal acritarchs. The presence of a few acanthomorphic acritarchs in the basal Jiucaiyuan Formation suggests the occasional influence of marine or brackish water. Palynofloras from the three horizons in the studied section are similar in species composition and percentages, and are tentatively grouped as the Lundbladispora–Lunatisporites–Aratrisporites Assemblage. This assemblage is characterized by dominant and diverse pteridophytic spores and subordinate but almost equally diverse gymnospermous pollen. Among the spores, the newly evolved lycopods are relatively important. Permian-type coniferous and pteridospermous species are also prominent components. Comparisons with the known Permian–Triassic assemblages of the world indicates that the Lundbladispora–Lunatisporites– Aratrisporites Assemblage is early Early Triassic (early Induan) in age. Based on palynomorph and conchostracan evidence in the studied section, the lower boundary of the Triassic may be drawn at the base of Unit 3 in the uppermost Guodikeng Formation, being slightly lower than the traditional boundary. The section may be considered as one of the candidates for the Permian–Triassic boundary stratotype for terrestrial

strata, due to the well-documented stratigraphy and the rather continuous record of various fossils in this section. The possibility of recycling of some pre-Triassic miospores cannot be excluded. However, it is more probable that the present palynoflora represents a Permian–Triassic transitional vegetation in the Early Triassic. Similar, roughly contemporaneous, transitional palynofloras are known in other areas in China and elsewhere, suggesting that no sudden catastrophic mass extinction of terrestrial floras occurred on a global scale at the close of Permian. Comparisons with the major Permian–Triassic phytogeographical provinces indicate that the Junggar palynoflora most closely compares with the Angara, and especially the Subangara Province, although it also contains a significant number of species common to the Euramerican Province and a few Cathaysian and Gondwana elements. Combined with the relevant paleobotanical data available, the area under study possibly belongs to the Subangara Province in Meyen’s (1988) sense during the Early Triassic, although the provincial differentiation was not as distinct as in the earlier period.

7. Descriptions of new species and new combinations All the taxa recorded and illustrated in the present paper are listed in Table 2. However, due to limited space, only new species and new combinations are described, and a few previously described species are briefly mentioned with synonyms or remarks. Genus Cyathidites Couper, 1953 Cyathidites sp. cf. C. breviradiatus Helby, 1967 (Plate I, 3–5) 1967 Cyathidites breviradiatus Helby, p. 63, pl. 1, fig. 4. 1967 Coniopteris sp., Fradkina, pl. I, fig. 3 (without description). 1977 ‘Acanthotriletes’ (levigate forms), Anderson, p. 29, pl. 41, figs. 1 and 3. 1986 Dictyophyllidites sp., Hou and Wang, pl. 25, fig. 6 (without description).

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1986 Cyathidites breviradiatus auct. non Helby, Qu and Wang, p. 133, pl. 31, fig. 7 (without description).

Punctatisporites sp. (Plate I, 7, 11)

Remarks: The specimens observed are closely similar to those cited above, namely, from the Lower– Middle Triassic in northeastern Siberia (Fradkina, 1967), and from the upper Upper Permian to Lower Triassic in Xinjiang (Hou and Wang, 1986; Qu and Wang, 1986). However, the species C. breviradiatus, described by Helby (1967, p. 63, pl. 1, fig. 4) from the Triassic of Australia is characterized by short trilete rays (< 1=2R) and thicker exine (2 µm) and is thus distinct from the species from the Junggar Basin.

Remarks: The specimens observed are identical to those identified as P. triassicus Schulz by Qu and Wang (1986) from the same formation and locality. The species described by Schulz (1964, p. 598, pl. 1, fig. 1) from the middle Buntsandstein of Germany is characterized by rather shorter trilete rays with distinct imperfect curvaturae and a punctate exine that is not differentiated into two layers. The latter features differentiate the present species from e.g. Punctatisporites pistilus Ouyang, 1986 (p. 38, pl. III, figs. 1 and 2).

Genus Punctatisporites Ibrahim, 1933 emend. Potonie´ and Kremp, 1954 Punctatisporites cf. P. asperatus Kara-Murza, 1952) Ouyang et Norris, comb. nov. (Plate I, 8, 9) Basionym: Leiotriletes asperatus Kara-Murza, 1952, Tr. Inst. Geol. Arktiki 31, 38, pl. 8, figs. 3, 8–10. Description: Amb subcircular, 33–40 µm in diameter (3 specimens). Trilete mark distinct, with labra 1 µm in breadth and 3 µm in height, extending nearly to the margin. Exine thin, <1 µm thick, levigate= chagrenate outside the contact areas where the exine is slightly thickened with scabrate-subgranulate ornament. Proximally a concentric fold is often present along the periphery, being almost confluent around the ends of the trilete rays. Comparison: The present specimens are somewhat similar to Leiotriletes asperatus Kara-Murza, 1952 from the Permian (P1 –P2 ) in the Taymir Basin except that in the latter the trilete rays are a little shorter (1=2–2=3R) and the exine appears somewhat thicker. We transfer the species to Punctatisporites because of its round shape and ’uneven’ exine. Todisporites concentricus Li (in Liu et al., 1981, p. 136, pl. I, figs. 14 and 15) first recorded from the Upper Triassic Yenchang Formation in Shaanxi differs in having a larger diameter (46–58 µm), and a distal concentric peripheral fold. Punctatisporites fissus Leschik (1955, p. 20, pl. 2, fig. 14) is also comparable, but it differs in having a microgranulate exine and an unclosed distal(?) fold.

1986 Punctatisporites triassicus auct. non Schulz, 1964; Qu and Wang, p. 135, pl. 31, fig. 6.

Genus Verrucosisporites Ibrahim, 1933 emend. Potonie´ and Kremp, 1954 Verrucosisporites jonkeri (Jansonius, 1962) Ouyang et Norris, comb. nov. (Plate II, 6–8) Basionym: Tsugaepollenites jonkeri Jansonius, 1962, Palaeontographica B 110, 51, pl. 12, figs. 4–6. 1985 Tsugaepollenites jonkeri Jansonius, Tuzhikova, pl. XIX, fig. 17; pl. VII, figs. 8 and 9. 1985 Tsugaepollenites (?) sp., Tuzhikova, pl. LII, fig. 7. 1986 Verrucosisporites sp. 1, Hou and Wang, p. 79, pl. 25, fig. 21. 1986 Tsugaepollenites jonkeri Jansonius, Qu and Wang, p. 159, pl. 33, figs. 21 and 22. Description: Amb circular to subcircular, 29 (34) 39 µm in diameter (6 specimens). Trilete mark generally not discernible, but occasionally distinctly seen (Plate II, 6), ca. 1=3–1=2 radius, sometimes with imperfect curvaturae. Exine ca. 2.0–2.5 µm in thickness, surface (distal and equatorial in particular) provided with irregular low and rather planar verrucae, very variable in size, from 0.5 to 6 µm (average 3–5 µm) in diameter, 1–1.5 µm in height, generally spaced 1 µm apart. The most diagnostic feature is that their uneven margins in surface view are like those of a cauliflower. Comparison and discussion: The present specimens are almost identical to those listed above. Hou and Wang (1986) reported this species in the upper Upper Permian at the Dalongkou section, and the others are

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PLATE I

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recorded from the Lower Triassic in Xinjiang, China and the Russian Urals. They are also indistinguishable from Tsugaepollenites jonkeri Jansonius, 1962 from the Lower Triassic of Canada. Tsugaepollenites Raatz, 1938 is characterized by closely spaced verrucae that are concentrated in a zone encircling the pollen. Jansonius interpreted the peripheral ornament as an ‘equatorial velum’, and noted that “on distal face ornaments strongly reduced leaving a large tenuitas” although he also mentioned “occasionally a faint reflection of a full Y mark has been observed”. Due to the presence of imperfect curvaturae and a trilete mark (Jansonius, 1962, pl. 12, fig. 5) which do

PLATE I Each entry comprises the following: sample number (e.g. AEA 749); slide number (e.g. -1); coordinates (e.g. 30.7=104). All figures ð600. 1. Leiotriletes exiguus Ouyang et Li, 1980. AEA 749-1, 30.7=104. 2. Leiotriletes turgidus Kara-Murza, 1952. AEA 751-5, 35=99. 3-5. Cyathidites sp. cf. C. breviradiatus Helby, 1967. 3. AEA 749-5, 33=103.6. 4. AEA 749-3, 42.2=105.7. 5. AEA 755-2, 29=101. 6. Calamospora breviradiata Kosanke, 1950, AEA 749-8, 35=104. 7, 11. Punctatisporites sp. 7. AEA 751-1, 39=101.1. 11. AEA 751-3, 36=104.1. 8a, b–9. Punctatisporites sp. cf. P. asperatus (Kara-Murza) Ouyang et Norris, comb. nov. 8. AEA 751-3, 42.1=103.7. 9. AEA 751-8, 35.2=100. 10. Cycloverrutriletes sp., AEA 755-1, 29.7=104.7. 12, 13. Granulatisporites sp. 12. AEA 755-1, 34=109. 13. AEA 755-1, 30.1=107.4. 14a, b. Verrucosisporites sp. AEA 755-1, 34.5=110. 15. Verrucosisporites minicus Qu et Wang, 1986. AEA 749-5, 35.1=96 16. Verrucosisporites sp. cf. V. minicus Qu et Wang, 1986. AEA 749-4, 36=106.6. 17–23. Anapiculatisporites decorus Ouyang et Norris, sp. nov. 17. AEA 755-1, 31=104.7. 18. Holotype, AEA 755-2, 36.3=111.7. 19. AEA 755-2, 40.5=100. 20. AEA 751-1, 40=98.1. 21. AEA 755-2, 36.9=107. 22a, b. AEA 755-2, 35=101. 23. AEA 755-2, 33=95.1.

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not accord with the definition of Tsugaepollenites, this species is transferred to Verrucosisporites. Verrucosisporites minicus Qu et Wang, 1986 (Plate I, 15, 16) 1986 Verrucosisporites minicus Qu et Wang, p. 139, pl. 31, figs. 24, 25. 1967 Lophotriletes sp. Fradkina, pl. I, fig. 20 (without description). Remarks: The specimen illustrated by Fradkina (1967, pl. I, fig. 20) under the name Lophotriletes sp. from the Lower–Middle Triassic of northeastern Siberia seems to be indistinguishable from V. minicus. Genus Acanthotriletes Naumova, 1939 ex 1949 emend. Potonie´ and Kremp, 1954 Acanthotriletes rostratus f. subrotundus (KaraMurza, 1952) Ouyang et Norris, comb. nov. (Plate II, 9) Basionym: Spinosella rostrata f. subrotunda KaraMurza, 1952, Tr. Inst. Geol. Arktiki 31, 13, pl. 1, fig. 15. Description: Amb subcircular, 35 µm in diameter. Exine 1.5–2 µm in thickness, surface with rather closely spaced spinae with basal diameter 1–1.6 µm and height 2–3 µm, gradually tapering distally to bluntly pointed apices, spaced 1–3 µm apart, ca. 36 in number along the periphery. Trilete rays simple, just discernible, extending to the equator. Comparison: This specimen is close to the type material illustrated and described by Kara-Murza (1952) from the Permian of the Taymir basin except that in the latter the spinae are slightly fewer in number although the author mentioned “a great number of long, relatively thin and curved spinae” (p. 13) occur on the surface. She pointed out that her new species and forma differs from Spinosella rostrata in having a smaller size and a rounded triangular outline. Spinosella Liuber, 1939 is a nomen nudum (see Jansonius and Hills, 1972–1976, Card 2653); its valid name Spinosisporites Liuber, 1966 (Jansonius and Hills, 1972–1976, Card 2655) is typified by a triangular amb. Spinososporites Knox, 1950 is a nomen nudum. Horriditriletes Bharadwaj et Salujka, 1964 is comprehensively sculptured with baculae, spinae, or coni (Foster, 1979). Therefore we tentatively use the

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PLATE II

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form genus Acanthotriletes and a new combination for the present species. In size and ornamentation, this species is also somewhat similar to Azonotriletes

PLATE II Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1–3. Anapiculatisporites decorus Ouyang et Norris, sp. nov. 1. AEA 755-1, 42.3=111.2. 2. AEA 755-1, 39=96. 3. AEA 749-6, 38=108. 4, 5. Apiculatasporites sp. 4. AEA 749-2, 33.2=106.9. 5. AEA 749-4, 36=107.6. 6–8. Verrucosisporites jonkeri (Jansonius) Ouyang et Norris, comb. nov. 6a, b. AEA 755-2, 39=110. 7. AEA 755-2, 33=95.1. 8. AEA 749-3, 31.8=101.3. 9. Acanthotriletes rostratus f. subrotundus (Kara-Murza) Ouyang et Norris, comb. nov. AEA 749-4, 34.9=100.2. 10–12. Apiculatisporis cf. A. subtilis Qu et Wang, 1990. 10. AEA 755-2, 39.5=101.3. 11. AEA 755-2, 40.1=107.2. 12. AEA 755-2, 36.3=112. 13–16. Apiculatisporis spiniger (Leschik) Qu, 1980. 13. AEA 751-8, 39.4=99.7. 14. AEA 755-2, 41.2=104.2. 15. AEA 755-1, 33.6.=96. 16. AEA 755-2, 36.1=104.8. 17–19. Baculatisporites uniformis Ouyang et Norris, sp. nov. 17. Holotype, AEA 749-2, 39.5=102.7. 18. AEA 749-7, 33.8=108.9. 19. AEA 749-2, 35=94.6. 20. Microreticulatisporites sp., AEA 755-1, 32=97. 21–22. Dictyotriletes mediocris Qu et Wang, 1990. 21. AEA 751-8, 43=104.4. 22. AEA 749-3, 30=104. 23a, b. Dictyotriletes sp. AEA 751-4, 31.1=109. 24, 25. Lycopodiumsporites sp. 24. AEA 755-2, 42.1=106. 25. AEA 751-6, 37.2=98.3. 26–28. Retitriletes? sp. 26. AEA 755-2, 37.1=112.1. 27. AEA 755-1, 38.9=110. 28. AEA 755-2, 42.5=102.3 29, 30. Annulispora folliculosa (Rogalska) De Jersey, 1959. 29. AEA 755-2, 38.3=110. 30. AEA 755-1, 431=107.4. 31–33. Limatulasporites limatulus (Playford) Helby et Foster, 1979. 31. AEA 751-1, 43.3=103.7. 32. AEA 751-5, 36.2=94.7. 33. AEA 751-2, 40=107.9.

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aggestus Andreeva, 1956 (in Andreeva et al., 1956, p. 244, pl. XLVII, fig. 34) from the upper Upper Permian of the Kuznetsk Basin but the latter is more triangular in outline. Genus Apiculatasporites Ibrahim, 1933 Apiculatasporites sp. (Plate II, 4, 5) ?1967 Acanthotriletes sp., Fradkina, pl. I, fig. 6. Remarks: The specimen shown on Plate II, 5 has been affected by pyrite growth. The species differs from other known species of Apiculatasporites or Acanthotriletes by its well-developed labra and dense, very small spinules and subcircular outline. It appears to be similar to Acanthotriletes sp. illustrated by line drawings by Fradkina (1967) from the Lower–Middle Triassic of northeastern Siberia, but this can not be determined with confidence. It differs from Apiculatisporis xiaolongkouensis Hou et Wang, 1986 (p. 81, pl. 21, figs. 20, 21, 30) from the Upper Permian in the Junggar basin in having a smaller diameter and smaller spines. Genus Apiculatisporis Potonie´ et Kremp, 1956 Apiculatisporis cf. A. subtilis Qu et Wang, 1990 (Plate II, 10–12) Remarks: The present specimens (31–36 µm) are comparable with the Upper Permian Apiculatisporites xiaolongkouensis Hou et Wang, 1986 (p. 81, pl. 21, figs. 20, 21, 30; 45–60 µm) which is larger and with the Middle Triassic (Karamay Formation) Apiculatisporis subtilis Qu et Wang, 1990 (p. 50, pl. 11, figs. 31–33) which is smaller, both from the Junggar Basin. The species is similar to Spinosella densa Kara-Murza, 1952 (p. 49, pl. 10, fig. 12) from the Taymir Basin (P21 –P12 ), typically triangular in outline. Some specimens of Lophotriletes sp. Fradkina, 1967 (pl. 1, fig. 12) from the Lower– Middle Triassic of northeastern Siberia are closely similar but the line drawings are inadequate and there is no description. Apiculatisporis spiniger (Plate II, 13–16)

(Leschik)

Qu,

1980

1955 Apiculatisporites spiniger Leschik, p. 18, pl. 2, figs. 6, 7.

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1980 Apiculatisporis spiniger (Leschik) Qu, p. 130, pl. 68, fig. 15; pl. 70, fig. 9. 1986 Apiculatisporis spiniger (Leschik) Qu, in Qu and Wang, p. 143, pl. 31, figs. 38, 44; pl. 37, fig. 12. 1986 Acanthotriletes sp., Qu and Wang, pl. 31, fig. 40 (without description). ?1986 Anapiculatisporites cooksonae auct. non Playford; Qu and Wang, p. 143, pl. 39, fig. 29. ?1986 Apiculatisporis cf. spiniger (Leschik) Qu, in Hou and Wang, p. 81, pl. 25, fig. 22. Remarks: The forms recorded as Anapiculatisporites spiniger (Leschik) Reinhardt by Reinhardt and Schmitz (1965) from the Upper Buntsandstein of Germany represent another species whose spinae are longer, more regularly distributed and do not show very swollen bases. On the other hand, Daneopsites parvispinellatus Maljavkina, 1953 as described and illustrated by Varjukhina (1971, p. 48, pl. I, fig. 4) from the Triassic of northeast European Russia appears to be closely related, if not conspecific, with A. spiniger. Originally Leschik (1955) compared his species with a living species of Danea. Genus Anapiculatisporites Potonie´ et Kremp, 1954 Anapiculatisporites decorus Ouyang et Norris, sp. nov. (Plate I, 17–23; Plate II, 1–3; Fig. 4)

torial diameter 30 (36) 56 µm, including ornament (10 specimens); holotype 34 µm. Trilete rays simple or with thin labra, occasionally open, about 1=2–2=3 radius in length, sometimes with imperfect to nearly perfect curvaturae which may be coalescent with exine subequatorially, resulting in a pseudocingulate appearance. Exine 1.5–4 µm thick, distally and equatorially the surface covered with rather dense warts or conical tubercles, 1–3 µm in basal diameter, generally 1–1.5 µm in height, with obtuse or rarely acute extremities spaced 0.5–2 µm apart, about 30–40 in number along the periphery. Contact areas levigate. Remarks: The specimens shown on Plate II, 1–3 represent rather poorly preserved specimens of the new species with more circular and rounded warts and indistinct trilete rays. Comparison: The species is somewhat similar to Anaplanisporites stipulatus Jansonius, 1962 (p. 45, pl. 11, figs. 17, 18) from the Lower Triassic of Canada, but differs in having a thicker exine and more prominent ornamentation as well as arcuate ridges. The specimens recorded from the upper Buntsandstein of Germany and identified as Anapiculatisporites sp. by Reinhardt and Schmitz (1965, p. 19, pl. 1, figs. 9, 10) are also similar, but they have mostly conical ornamental elements with pointed ends. Genus Baculatisporites Pflug et Thompson, in Thompson and Pflug, 1953

Holotype: Sample-slide=coordinate: AEA 755-2= 36.3 ð 111.7 (Plate I, 18; Fig. 4). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province. Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: Latin, decorus, decorative, beautiful. Description: Amb circular to subcircular or rounded triangular, nearly plano-convex in lateral view. Equa-

Holotype: Sample-slide=coordinate: AEA 749-2= 39.5 ð 102.7 (Plate I, 17). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province.

Fig. 4. Drawing of the holotype of Anapiculatisporites decorus Ouyang et Norris, sp. nov. ð600.

Fig. 5. Drawing of Baculatisporites uniformis Ouyang et Norris, sp. nov. ð600.

Baculatisporites uniformis Ouyang et Norris, sp. nov. (Plate II, 17–19; Fig. 5)

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Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: Latin, uniformis, uniform. Description: Amb subcircular, 29–35 µm in diameter (5 specimens), holotype 33 µm. Trilete rays simple, slightly open, nearly extending to equator. Exine ca. 1.5–2 µm in thickness, surface provided with dense and rather uniform bacula, 1–1.5 µm in basal diameter, 2–3 µm in height, spaced 0.5–1 µm (occasionally 2.5 µm) apart, with often slightly swollen ends, or obtusely pointed or uneven to faintly bifurcated extremities, about 40–60 in number along the periphery. The ornament becomes diminished and scattered toward the proximal surface. Comparison: This new species is similar to Baculatisporites comaumensis (Cookson) Potonie´ (see Cookson, 1953, p. 470, pl. 2, figs. 27, 28) but differs in having a thicker exine and denser bacula often with swollen apices. Neoraistrickia amblyeformis Hou et Wang, 1986 (p. 82, pl. 21, figs. 11, 12) from the Upper Permian in the Dalongkou section is distinguished by its larger diameter (40–45 µm) with stronger and irregular baculate ornamentation. Azonotriletes selaginelliformis Samoilovich, 1953 (p. 51, pl. XI, fig. 7) from the Kungurian of the Cis-Urals is characterized by the presence of sparsely spaced bacula or spinae (“irregularly scattered spines with blunt apices”). Genus Dictyotriletes Naumova, 1939 ex Ischenko, 1952 emend. Potonie´ and Kremp, 1954 Dictyotriletes mediocris Qu et Wang, 1990 (Plate II, 21, 22) 1990 Dictyotriletes mediocris Qu et Wang, p. 50, pl. 9, figs. 13, 23, 24. ?1952 Reticulina lucida Kara-Murza, p. 64, pl. 12, fig. 27. ?1967 Dityotriletes sp. Fradkina, pl. I, fig. 13 (without description). Remarks: The present specimens are very close to D. mediocris Qu et Wang, 1990 from the Lower Triassic Shaofanggou Formation in northern Xinjiang; they are also comparable to Reticulina lucida KaraMurza, 1952 from the Upper Permian of the Taymir Basin in size and general aspect, which, however, shows rather flat muri without projections at the nodes. Reticulina as used by Kara-Murza (1952, pp.

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63, 64, 110) is a nomen nudum because its authorship is not mentioned and a type species was not designated. The specimen illustrated and named Dictyotriletes sp. by Fradkina (1967) from the Lower– Middle Triassic of northeast Siberia is probably conspecific but is larger (65 µm?). Dictyotriletes sp. (Plate II, 23) ?1986 Maculatasporites sp., Qu and Wang, pl. 33, fig. 25 (without description). Genus Microreticulatisporites Knox, 1950 emend. Potonie´ and Kremp, 1954 Microreticulatisporites sp. (Plate II, 20) Remarks: The lumina of the reticulum in Lycopodiumsporites antiquus de Jersey 1964 (p. 5, pl. 1, fig. 5) from the Triassic of Australia are somewhat larger (mostly 2–3 µm). Genus Limatulasporites Helby et Foster, in Foster, 1979 Limatulasporites fossulatus (Balme) Helby et Foster, 1979 (Plate III, 14–18) 1970 Nevesisporites fossulatus Balme, p. 335, pl. 3, figs. 1–5. 1966 Limbella ovaliformis Maljavkina, Kurbatova, pl. I, fig. 10. 1967 Chomotriletes sp. (in part), Fradkina, pl. I, figs. 16–21. 1971 Discisporites colliculiniformis (Maljavkina) Varjukhina, p. 80, pl. 24, fig. 15. 1973 Nevesisporites fossulatus Helby, p. 154, pl. 1, fig. 4. 1973 Nevesisporites sp., Obonitsakaya, pl. IV, figs. 40–43. 1979 Limatulasporites fossulatus (Balme) Helby and Foster, Foster, p. 51, pl. 13, figs. 1–3. 1985 Discisporites colliculiniformis (Maljavkina) Varjukhina, Tuzhikova, p. 157, pl. XXVII, figs. 21, 21. 1986 Limatulasporites fossulatus (Balme) Helby et Foster, Hou and Wang p. 86, pl. 25, figs. 32–34. 1986 Limatulasporites fossulatus (Balme) Helby et Foster, Qu and Wang, p. 149, pl. 31, figs. 53, 54; pl. 37, fig. 30; pl. 39, fig. 46.

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PLATE III

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Remarks: L. fossulatus shows variation in the proximal ornamentation, diameter of distal crassitude and thickness of cingulum. The principal features of our specimens are identical to those listed above, including those by Balme (1970) and Foster (1979). Transitional forms between L. fossulatus and L. limatulus (Playford) do occur in our samples which makes the separation of the two species somewhat difficult. Specimens with rather regular prominent proximal grana are assigned to L. limatulus. This species has a wide distribution. It is known from Pakistan (Permian), Australia (Permo-Triassic), the former U.S.S.R. (Tunguska, Kuznetsk, Urals, north-

PLATE III Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1. Remysporites dubovii Tuzhikova, 1985. AEA 749-5, 35=105. 2. Lundbladispora disparilis Qu et Wang, 1986. AEA 751-8 38.1=107. 3. Cingulizonates sp., AEA 751-8, 38.1=108.8. 4, 7. Densosporites sp. cf. C. spongeosus Butterworth et Williams, 1958. 4. AEA 751-4, 37.9=100.9. 7. AEA 751-5, 36.1=94.4. 5. Rotaspora sp., AEA 749-6, 27.2=104.3. 6. Nevesisporites sp., AEA 479-6, 27.1=105.2. 8. Stenozonotriletes sp., AEA 751-5, 38.3=102.8. 9. Lundbladispora sp., AEA 751-2, 37.1=103.8. 10. Pterisisporites sp., AEA 755-1, 42.4.=109. 11. Lundbladispora sp. A, AEA 751-5, 30.7=101. 12. Lundbladispora sp. B, AEA 749-3, 34.6=105. 13. Kraeuselisporites? sp., AEA 751-1, 30.1=100. 14–18. Limatulasporites fossulatus (Balme) Helby et Foster, 1979. 14. AEA 751-6, 27.6=102. 15. AEA 749-2, 31=95.2. 16. AEA 751-4, 41.8=100.9. 17. AEA 751-1, 39=105. 18. AEA 749-6, 32=104. 19, 20. Lundbladispora sp. C. 19. AEA 751-4, 45=105. 20. AEA 749-8, 30.2=108.1 21–23. Lundbladispora foveota Qu et Wang, 1986. 21. AEA 749-4, 38=103.3. 22. AEA 749-1, 39.2=108.1. 23. AEA 751-8, 40.6=106.4 24–26. Lapposisporites echinatus Ouyang et Norris, sp. nov. 24. AEA 749-3, 43=105.1. 25. Holotype, AEA 749-6, 37.3=103.9. 26. AEA 749-8, 42=105.

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eastern Siberia, mainly Lower Triassic) and northwestern China (Xinjiang, Lower Triassic). In addition, Dulhuntyspora? minuta Jansonius, 1962 (p. 48, pl. 11, figs. 1–5) from the Lower Triassic of Canada seems also to belong to Limatulasporites, and may possibly be conspecific with L. fossulatus. Genus Densosporites Berry, 1937 emend. Butterworth, Jansonius, Smith and Staplin, 1964 Densosporites sp. cf. D. spongeosus Butterworth et Williams, 1958 (Plate III, 4, 7) 1958 Densosporites spongeosus Butterworth et Williams, p. 380, pl. 3, figs. 40–41. 1986 Densosporites spongeosus Butterworth et Williams, Qu and Wang, p. 150, pl. 32, fig. 43. Remarks: In size and general morphology, the present specimen is close to D. spongeosus recorded by Qu and Wang (1986) from the same formation and locality except that in the latter the cingulum is somewhat thicker. Among the known species of Densosporites (e.g. Kosanke, 1950; Potonie´ and Kremp, 1955; Butterworth and Williams, 1958; Smith and Butterworth, 1967), the present specimen most closely resembles D. spongeosus. However, the trilete mark in the latter is not so prominent as in ours. Smith and Butterworth (1967, p. 244) treated D. spongeosus as a junior synonym of D. triangularis Kosanke, 1950 (p. 34, pl. 7, fig. 1), but we use the former name because our specimen has a narrower cingulum and lacks the granulose to vermiculate ornament as well as ‘a few spines’ along the periphery of the cingulum characterizing D. triangularis. Genus Pterisisporites Song, 1976 Pterisisporites sp. (Plate III, 10) 1986 Lycospora imperialis auct. non Jansonius, 1962; Qu and Wang, p. 151, pl. 32, figs. 14, 15. Remarks: Morphologically, the species is closer to Pterisiporites Song than to Lycospora even though the type species of the former is derived from the Tertiary.

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PLATE IV

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Genus Kraeuselisporites Leschik, 1955 emend. Scheuring, 1974 Kraeuselisporites spinulosus Hou et Wang, 1986 (Plate IV, 6–8) 1986 Kraeuselisporites spinullous Hou et Wang, p. 85, pl. 21, figs. 21, 23. Remarks: The orthography of the specific epithet ‘spinullous’ is here corrected to its proper Latin spelling spinulosus, in accordance with the ICBN. Comparison: The present specimens closely resemble K. spinulosus Hou et Wang, 1986 from the Upper Permian in the Dalongkou section, but the trilete mark may be shorter than in the latter. K. argutus Hou et Wang, 1986 (p. 85, pl. 21, fig. 24), which was said to be distinguished from K. spinulosus in having thicker endexine and comparatively stronger coni may be a conspecific taxon. K. spinulosus is similar to K. spinosus Jansonius, 1962 (p. 47, pl. 11, figs. 22, 23) from the Lower Triassic of Canada, but has a less distinct and darker inner zone of the cingulum and a shorter trilete mark. Kraeuselisporites varius Ouyang et Norris, sp. nov. (Plate IV, 1–3, 9–14; Fig. 6) ?1986 (pars) Kraeuselisporites disparilis Qu et Wang, p. 155, pl. 32, fig. 41 (not the holotype fig.

PLATE IV Each entry coordinates. 1–3, 9–14. 1a, b. 2. 3. 9. 10. 11. 12. 13. 14. 4, 5. 4. 5. 6–8. 6. 7. 8.

comprises the sample number, slide number and All figures ð600. Kraeuselisporites varius Ouyang et Norris, sp. nov. Holotype, AEA 749-5, 34=103.9. AEA 751-3, 33.7=99. AEA 751-2, 35=98. AEA 749-4, 30.1=104. Paratype, AEA 751-8, 31.8=104.2. AEA 749-8, 36=107. AEA 749-5, 31=103.1. AEA 749-3, 39.2=100. AEA 751-5, 30=104.3. Kraeuselisporites echinatus Reinhardt et Scho¨n, 1967. AEA 749-1, 38=109. AEA 749-2, 38=101.1. Kraeuselisporites spinulosus Hou et Wang, 1986. AEA 749-2, 32.5=108. AEA 749-2, 33.2=100.5. AEA 749-4, 33.2=100.

Fig. 6. Drawing of Kraeuselisporites varius Ouyang et Norris, sp. nov. ð600.

40). ?1986 Kraeuselisporites sp., Qu and Wang, pl. 33, figs. 6, 7 (without description). Holotype: Sample-side=coordinate: AEA 749-5= 34 ð 103.9 (Plate IV, 1). Paratype: Sample-slide=coordinate: AEA 751-8= 31.8 ð 104.2 (Plate IV, 10). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalonkou, Junggar Basin, Xinjiang Province. Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: Latin, varius, various. Description: Zonate spores, amb rounded triangular to subcircular, lenticular in lateral view, with distal face more convex. Diameter 44 (64) 82 µm (10 specimens), central body 38 (50) 64 µm, occasionally larger; holotype 62 µm, paratype 65 µm. Outline of body basically conforms with that of spore. Exine moderately thick, spongy-scabrate, thickened distally (ca. 2 µm) and equatorially in particular, which results in a cuneiform cingulum with a width of usually 10–14 µm. The cingulum is differentiated into two zones, a darker inner zone (covering the body) and a lighter outer zone with a width of ca. 3–5 µm beyond the body, although the boundary between the outer and the inner zone of the cingulum is sometimes unclear. Distally, and to a lesser degree equatorially, the surface of the exine may be provided with moderately densely spaced spinulae and=or coni. Spinulae ca. 1–1.5 µm in basal diameter, 1 µm in height, spaced 1–4 µm apart and coni 1.5–2.5 µm in basal diameter, spaced 2–4 µm apart, with blunt or acute apices. Minute spinulae or coni

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Fig. 7. Drawing of Lapposisporites echinatus Ouyang et Norris, sp. nov. (a) Tetrad. (b) Single spore. ð600.

may occur along the periphery of the spore. Proximal surface without distinct ornamentation. Trilete rays often with strong labra, 2–4 (6) µm in breadth and height, straight or somewhat flexuous, extending onto the cingulum or to the amb. Remarks: This species may be subdivided into three groups according to the size, pattern of spines, and type of sculpture viz. spinules (Plate IV, 1–3), spinae (Plate IV, 9–11) and rather circular warts (Plate IV, 12). We believe all these forms represent a rather wide range of variation of one single species. Features shared amongst the 3 groups include size, basically spinate ornamentation, crassitude-like and cuneiform cingulum, and usually strong labra. The specimens on Plate IV, 11 and 14, respectively, are extremes of the morphologic variation. Comparison: This new species differs from the other species of Kraeuselisporites in its strong labra, cuneiform cingulum and characteristic ornamentation (e.g. Leschik, 1955; Balme, 1970). Qu and Wang (1986) reported and illustrated some 14 species of Kraeuselisporites from the Triassic, including 6 species from the Jiucaiyuan Formation. Among them, however, none is definitely conspecific with the present new species with the exception of those listed questionably in the synonymy. Our species is somewhat similar to K. spinosus Jansonius, 1962 (p. 47, pl. 11, figs. 22–24) and K. apiculatus Jansonius, 1962 (p. 47, pl. 11, figs. 25, 26) from the Lower Triassic of Canada, but differs in having much stronger labra and less distinct differentiation of the dark inner zone of the cingulum as well as a narrower and relatively irregular outer zone. Furthermore, K. apiculatus has much longer spines. K. hoofddijkensis Visscher, 1966 from the upper Lower

Triassic is distinguished by a narrower cingulum, labra with expanded ends, and broader-based warts. Genus Lapposisporites Visscher, 1966 Lapposisporites echinatus Ouyang et Norris, sp. nov. (Plate III, 24–26; Fig. 7a, b) Holotype: Sample-slide=coordinate: AEA 749-6= 37.3 ð 103.9 (Plate III, 25) Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province. Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: echinatus, spinulate, from Greek, echinos (Latin, echinus), sea urchin. Description: Cingulate, trilete spores, often united into tetrads. Tetrads subcircular to sub-trilobate in outline, size 57 (74) 100 µm (10 tetrads), holotype 67 µm. Amb of individual spores subcircular, 54– 60 µm in diameter. Exine 1–3 µm in thickness, equatorially thickened into a cingulum, generally 3– 5 µm in breadth or thickness; surface spongy and provided with relatively and uniformly dense spinulate ornamentation that may vary between specimens, 1–3 µm in basal diameter, 1.0–1.5 (3) µm in length, spaced 1–3 µm apart, some bases swollen or even connected into cristae-like groups, gradually or abruptly (near the upper part) sharpened with acute or obtuse ends; occasionally with grana-spines. Trilete rays distinct with labra, extending to equator, but generally not easily discernible due to overlap in the tetrad.

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Comparison and discussion: All the specimens are attributed to one species, because transitional forms occur and further specific separation is difficult. This new species differs from all others in Lapposisporites in the possession of smaller spinate ornamentation and size. For instance, of the several species proposed by Visscher (1966) all have different ornamentation (L. lapposus — scabrae, grana and gemmae; L. villosus — longer spines (6–7 µm); L. armatus — scattered spinae or spines and verrucae with superposed spine-shaped differentiation; L. loricatus — polygonal verrucae with superposed spine-shaped differentiation) and larger size (90–120 µm). L. echinatus appears to be similar to the forms described and illustrated as L. lapposus Visscher by Qu and Wang (1986, p. 146, pl. 32, fig. 49) from the Jiucaiyuan Formation in the Dalongkou section. The tetrad is 75 µm in diameter and was described with “granulate or small tuberculate ornamentation” but, to judge from the figure, it seems to be spinulate-granulate, and therefore might be ascribed to L. echinatus. From our observations, it appears probable that the genus Lapposisporites represents the tetrad of cingulate Kraeuselisporites-type spores; while Paralundbladispora Visscher, 1966 is the tetrad of cavate Lundbladispora type spores. Visscher also pointed out that in the genus Paralundbladispora, single spores (albeit ‘very rare’) do occur in the same collection with tetrahedral tetrads. In living plants, similar relationships exist. For instance, the spores of Selaginella deflexa Brack generally occur in tetrads, but as Selling (1946, p. 84, pl. 1, figs. 24, 25) mentioned they also occur singly. The cingulum in the spores of Lapposisporites seems to be discernible as well in some of Visscher’s specimens (e.g. Visscher, 1966, pl. III, fig. 1A — lower left; pl. IV, fig. 2 — upper spore; pl. V, fig. 1 — lower left). If our judgement is correct, the genus should be emended. Genus Lundbladispora Balme, 1963 Lundbladispora foveota Qu et Wang, 1986 (Plate III, 21–23) 1986 Lundbladispora foveota Qu et Wang, p. 153, pl. 32, figs. 32, 33; pl. 33, fig. 15. 1967 Brochotriletes sp., Fradkina, pl. I, fig. 14 (without description). Remarks: Our specimens are similar to Brochotri-

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letes sp. described by Fradkina (1967) from the Lower–Middle Triassic of northeastern Siberia, although the cavate nature is hard to discern in the poorly sketched figure. Lundbladispora disparilis (Qu et Wang, 1986) Ouyang et Norris, comb. nov. (Plate III, 2) Basionym: Kraeuselisporites disparilis Qu et Wang, 1986, Geol. Mem. Ser. 2, 3, Geol. Publ. House, Beijing, p. 155, pl. 32, fig. 40. Description: Cavate spore, amb triangular with convex sides and rounded or slightly pointed angles, diameter 55 µm. The outline of the central body conforms with that of the amb, diameter 46 µm. Endexine levigate, equatorially slightly thickened but not forming a cingulum. Exoexine thin, spongymicroreticulate with lumina 0.5 µm across, enclosing the entire central body, equatorially extending 4– 5.5 µm beyond the body, being a little wider at the apices. Trilete rays with robust membranous labra with a broader base and narrower vertex, extending to the apices of the amb. Remarks: In general morphology and size, this specimen is closely comparable to the holotype of K. disparilis Qu et Wang, 1986 (pl. 32, fig. 40) from the Jiucaiyuan formation in the Dalongkou section. In their description, Qu and Wang (1986, p. 155) mentioned “Exine composed of two layers, cavate ...; exoexine thin, transparent, ... surface provided with coarsely granulate ornamentation, and equatorially with sparse radiate rugae”. The grana appear to occur on the other specimen (pl. 32, fig. 41) but are not typically developed on the holotype, and the ‘rugae’ may be derived from secondary folding of the exoexine. Thus we view the present specimen and the holotype of L. disparilis to be conspecific. We have reassigned these spores to the genus Lundbladispora because of the presence of a cavity. This species differs from other species of Lundbladispora in having a thin exoexine, a narrow and unequal cavate margin, strong labra and in the absence of distinct exoexinal ornamentation and an equatorial limbus (cf. Balme, 1970; Tuzhikova, 1985). Lundbladispora echinata Reinhardt et Scho¨n, 1967 (Plate IV, 4, 5) 1967 Lundbladispora echinata Reinhardt et Scho¨n, p. 749, pl. 1, fig. 11.

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?1986 Acanthotriletes sp., Hou and Wang, pl. 25, fig. 31 (without description). Remarks: These specimens are comparable to Acanthotriletes sp. (Hou and Wang, 1986) from the Upper Permian in the Dalongkou section, but the latter lacks a description. Genus Remysporites Butterworth et Williams, 1958 Remysporites dubovii Tuzhikova, 1985 (Plate III, 1) 1985 Remysporites dubovii Tuzhikova, p. 97, pl. XXI, fig. 3. Remarks: The present specimen (73 µm) is closely similar to those described by Tuzhikova (1985) from the Lower Triassic of the Urals in all salient features except that the exine in the latter appears a little thicker. She mentioned that “in the radial areas of the miospore, the trilete mark appears as if pulled to the inner part of zona, which causes the formation of concentric semicircular folds and a reduction of the zona breadth in the radial direction” (translated from Russian, p. 98). This feature, as shown in her drawing (pl. LIX, fig. 1a), is not present in our specimens. However, we believe that even if her explanation is correct it is not necessarily a diagnostic feature for the species, given that only two specimens have been measured and one specimen illustrated. Genus Cordaitina Samoilovich, 1953 Cordaitina rugulifera (Liuber) Samoilovich, 1953 (Plate V, 4, 5) 1941 Zonaletes rugulifer Liuber, in Liuber and Val’ts, p. 111, pl. XV, fig. 250a–d. 1941 Zonaletes angustelimbatus Liuber, in Liuber and Val’ts, p. 177, pl. XV, fig. 249b. 1956 Zonaletes rugulifer Liuber in Andreeva et al., p. 253, pl. LII, fig. 63a, b. 1960 Cordaitina rugulifer (Liuber) Samoilovich, Medvedeva, pl. XVI, figs. 6, 6a. Remarks: The specimen on Plate V, 4, is closer to Circella stenolimbata Liuber (in Liuber and Val’ts, 1939, pl. 4, fig. 1 and Kara-Murza, 1952, p. 79, pl. 18, figs. 1, 4–6) and Circella angustelimbatus Liuber (in Liuber and Val’ts, 1941, p. 177, pl. 15, fig. 249) than to C. rugulifera (Liuber). On the other

hand, the specimen illustrated in Plate V, 5 bears close similarity with C. rugulifera (Liuber). Considering the common features shared by both species, it seems probable that they are conspecific. Hart (1965, p. 94) believed C. stenolimbata Liuber, 1939 and C. stenolimbata in Kara-Murza etc. are synonymous with Zonaletes angustelimbatus Liuber, 1941, and placed them in Nuskoisporites based on the presumed presence of a trilete mark and particularly on the presence of an equatorial saccus limbus. Without designating a holotype and emending the diagnosis, Hart mentioned under Nuskoisporites angustelimbatus (Liuber) Hart that “the central body is circular and has a finely granulate to tuberculate sculpture”. Furthermore, Hart expressed the view that the presence of a trilete mark is not necessarily a diagnostic feature for separating monosaccate genera. Consequently, he included several species with trilete mark in the genus Cordaitina. This treatment is hard to accept, as Balme (1970, p. 355) has already pointed out. Circella Liuber, 1939 is a nomen nudum because

PLATE V Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1. Latosporites sp. AEA 749-1, 41.9=105.1 2, 3. Aratrisporites sp. 2. AEA 749-2, 37=95.3. 3. AEA 749-7, 34.7=99. 4, 5. Cordaitina rugulifera (Liuber) Samoilovich, 1953. 4. AEA 751-5, 37=95.2. 5. AEA 751-7, 36.7=106.8. 6, 7. Samoilovitchisaccites? sp. 6. AEA 751-1, 31.2=109. 7. AEA 751-1, 42.2=106.6. 8. Iunctella sp., AEA 751-3, 37.8=103.4. 9. Crucisaccites sp. AEA 751-7, 32.6=105.8. 10. Cordaitina gemina (Andreeva) Hart, 1965. AEA 751-4, 43.5=102.9. 11. Cordaitina abutiloides (Andreeva) Ouyang et Norris, comb. nov. AEA 749-3, 37.1=108. 12, 14. Vesicaspora ex gr. magnalis (Andreeva) Hart, 1965. 12. AEA 751-7, 36.1=108.9. 14. AEA 751-2, 38=96.8. 13. Vesicaspora acrifera (Andreeva) Hart, 1965. AEA 751-3, 31=103.1. 15, 16. Vitreisporites pallidus (Reissinger) Nilsson, 1958. 15. AEA 749-1, 37=103.6. 16. AEA 751-6, 30=99.3. 17. Florinites? sp., AEA 751-7, 41.4=102.

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PLATE V

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it is without a designated type species and does not have a diagnosis. Therefore, taking Circella stenolimbata Liuber (in Liuber and Val’ts, 1939, pl. 4, fig. 1) as the basionym we propose a new combination for the species under Cordaitina. Kara-Murza (1952, p. 79) also considered C. angustelimbatus to be a synonym of C. stenolimbata, but the latter has priority. We suggest that the main difference between C. rugulifera and C. stenolimbata is that the former is generally larger in size (60–80 µm) and has a wider saccus overlap, while the latter displays a narrower limbate saccus and the central body area is often provided with a finely granulate and tuberculate ornamentation. These two species were both originally described from the Permian of the Kuznetsk Basin and appear to partly overlap in morphological characters. Cordaitina duralimita Hou et Wang, 1986 (p. 87, pl. 22, figs. 4, 5) from the Upper Permian at the Dalongkou section in the Junggar Basin is probably a junior synonym of Cordaitina stenolimbata (Liuber). Cordaitina abutiloides (Andreeva) Ouyang et Norris, comb. nov. (Plate V, 11) Basionym: Zonaletes abutiloides Andreeva, in Andreeva et al., 1956, Atlas of the leading forms of fossil flora and fauna of the Permian System of the Kuznetsk Basin. Tr. VSEGEI, p. 254, pl. LIII, fig. 64a, c. 1983? Virkkipollenites sp., Zhang, pl. 2, figs. 14, 18 (without description). Description: Monosaccate pollen grains, circular to subcircular in polar view, 64–68 µm in diameter (3 specimens). Endexine dense, ca. 3 µm in thickness, equatorially thickened to 3–4 µm at the base of the saccus. Exoexine (saccus) rather loose, saccus overlap ca. 7 µm in breadth, with an intrabaculate to radially pleated texture, uneven or undulating along the periphery. Surface in apical view punctate to microreticulate with muri and lumina ca. 1 µm across, locally irregularly granulate-microtuberculate, mostly <1 µm in diameter. Comparison: the present specimens are closely similar to Zonaletes abutiloides Andreeva, 1956 from the Upper Permian of the Kuznetsk Basin, especially her fig. 64c (right) which is almost identical. Andreeva mentioned that this species differs from C. rotatus (Liuber) in having a thick fringe (saccus) and radial

folds in the saccus. Thus, the species abutiloides should be transferred to Cordaitina, and this new combination is proposed here. The Xinjiang specimens are also similar to Circella rotata Luber forma arctica Kara-Murza, 1952 (p. 27, pl. 5, figs. 1–3; p. 76, pl. 16, figs. 2, 8) from the Permian and rarely the Triassic of the Taymir Basin. However, the latter has a much larger size (with body 100–125 µm, see Kara-Murza, 1952, p. 27), although it is possibly conspecific with C. abutiloides. Cordaitina gemina (Andreeva) Hart, 1965 (Plate V, 10) 1956 Zonaletes geminus Andreeva in Andreeva et al., p. 256, pl. LIV, fig. 74a–c. 1965 Cordaitina gemina (Andreeva) Hart, p. 91, fig. 214. Remarks: This specimen is very similar to material illustrated by Andreeva (1956) from the Upper Permian of the Kuznetsk Basin, especially Andreeva’s pl. LIV, figs. b, c, which are also ovoid in outline. Hart (1965) selected her pl. LIV, fig. a (circular) specimen as the holotype. Andreeva et al. (1956, p. 257) pointed out that “the presence of a flat cylinder along the equator of the spore body is a characteristic feature for the present species. This cylinder gives an impression of a second fringe” (translated from Russian). This feature (flat cylinder D attachment zone or overlap in the present paper) is also clearly shown in our specimen (71 ð 47 µm), and the pollen size is also comparable (Andreeva reports the body to be 45–55 µm in diameter and the saccus 15–20 µm in breadth). Genus Crucisaccites Lele et Maithy, 1964 Crucisaccites sp. (Plate V, 9) Remarks: The present specimen differs from other species of Crucisaccites [e.g. Corisaccites ornatus Samoilovich, 1953 (p. 28, pl. III, fig. 1a, b) later transferred to Crucisaccites by Dibner (1971)] in having a smaller size and thinner saccus with punctate texture. It is unjustified to propose a new species because only a few poorly preserved specimens have been found. It is outside the scope of the present paper to discuss the complex problems of pollen morphology and the body-saccus construction of various species found mainly in Laurasia and assigned to

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Crucisaccites or Corisaccites by different authors, and their relation with the two genera with type species based on Gondwana material. However, according to recent observations and SEM photographs of many specimens by Wang Z. (pers. commun.) from the Permian in Xinjiang, there is little doubt that at least some of these pollen types are monosaccate rather than bisaccate (cf. the Gondwana genus Corisaccites; e.g. Balme, 1970). They seem to be also different in configuration from Crucisaccites. For the moment, we tentatively assign our specimens to the genus Crucisaccites. Genus Iunctella Kara-Murza, 1952 Iunctella sp. (Plate V, 8) Remarks: This specimen (130 ð 105 µm) is somewhat similar to Iunctella mirabilis Ouyang et Norris, 1988 (p. 206, pl. III, figs. 6–8) but differs from the latter in having a much smaller size and in the lack of a distinct tenuitas. Genus Florinites Schopf, Wilson et Bentall, 1944 Florinites? sp. (Plate V, 17) Remarks: This species (71 ð 92 µm) differs from other species of Florinites by its thick-walled central body and saccus (cf. Potonie´ and Kremp, 1956). It also differs from Vesicaspora or Striatolebachiites Varyukhina et Zauer in Varjukhina (1971) in the absence of a distal sulcus and the lack of regular and consistent proximal striae. It bears some similarity to Lebachiacites astranensis Maljavkina, 1964 (see Jansonius and Hills, 1972–1976, Card 1466) from the Permian (rarely Triassic), but in the latter “the body may be so large as to virtually make the pollen bisaccate; ... the body is not darker, sometimes even lighter in colour than the saccus”. It is also similar to Lebachiites lebachiiformis Maljavkina, 1964 (pl. 51, fig. 6) from the Lower Triassic of western Siberia, which also displays a tendency of the saccus to become bisaccate distally.

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differs from the former — e.g. S. turboreticulatus (Samoilovich, 1953, p. 53, pl. XI, fig. 13 a, b) Dibner — in the absence of typical protosaccate texture and in being larger, and from the latter in having a tendency to bisaccate differentiation distally, and in possibly developing a monolete mark proximally. Genus Vesicaspora Schemel, 1951 emend. Wilson and Venkatachala, 1963 Vesicaspora sp. ex gr. V. magnalis (Andreeva) Hart, 1965 (Plate V, 12, 14) 1956 Coniferaletes magnalis Andreeva in Andreeva et al., p. 267, pl. LIX, fig. 105. 1965 Vesicaspora magnalis (Andreeva) Hart, p. 73, fig. 171. 1980 (pars) Vesicaspora ex gr. magnalis (Andreeva) Hart, in Gomanikov and Meyen p. 118–120, figs. 2e–p, 3 and 4. Remarks: The Junggar specimens (39–56 ð 34–42 µm) are similar to Vesicaspora ex. gr. magnalis (Andreeva) as described and discussed in detail by Gomanikov and Meyen (1980) from the Permian of Angaraland. These authors pointed out that the total dimensions (57–97.5 µm) and the outline of the sulcus vary widely. Our specimens are smaller than those recorded by the Russian authors. On the other hand, the Xinjiang specimens are not unlike Vesicaspora wilsonii Schemel, 1951 (p. 479, figs. 1, 3) but differ from the latter in having a more distinct and broader sulcus. Our illustrated specimens (Plate V, 12, 14) are quite similar to those illustrated by Visscher et al. (1974, pl. III, figs. 3, 4) as cf. V. wilsonii Schemel sensu Helby, 1966 from the Upper Rotliegendes of Germany. Vesicaspora acrifera (Andreeva) Hart, 1965 (Plate V, 13)

Samoilovitchisaccites? sp. (Plate V, 6, 7)

1956 Coniferaletes acriferum Andreeva in Andreeva et al. 1956, p. 269, pl. LX, fig. 117 (upper figure). ?1956 Sulcatisporites splendens Leschik, p. 137, pl. 22, fig. 10. 1965 Vesicaspora acriferum (Andreeva) Hart p. 73, fig. 172. ?1976 Florinites duralis Inosova in Inosova et al., p. 191, pl. IX, figs. 6, 7.

Remarks: This species is transitional in character between Samoilovitchisaccites and Cordaitina. It

Remarks: This specimen (53 µm) closely resembles those described by Andreeva (1956) from the Upper

Genus Samoilovitchisaccites Dibner, 1971

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Permian of the Kuznetsk Basin, especially that on her pl. LX, fig. 117 (upper) with a size range of 35–55 µm. It is possible, however, that V. acrifera partly overlaps in circumscription with V. magnalis. Judging from the description and illustrations of Florinites duralis Inosova (in Inosova et al., 1976), we consider this species might be a junior synonym of V. acrifera. Genus Klausipollenites Jansonius, 1962 Klausipollenites schaubergeri (Potonie´ et Klaus) Jansonius, 1962 (Plate VI, 3–5) 1954 Pityosporites schaubergeri Potonie´ et Klaus, 1954, p. 536, pl. 10, fig. 7. Synonyms (see Balme, 1970, pp. 385–386, and the following additions): 1981 Klausipollenites schaubergeri (Potonie´ et Klaus) Jansonius, Dybova-Jachowicz, pl. X, figs. 8–10; pl. XI, figs. 1, 2. 1986 Klausipollenites schaubergeri (Potonie´ et Klaus) Jansonius, in Hou and Wang, p. 98, pl. 27, figs. 27, 28. 1986 Klausipollenites schaubergeri (Potonie´ et Klaus) Jansonius, in Qu and Wang, p. 166, pl. 36, figs. 5, 6. 1985 Klausipollenites schaubergeri (Potonie´ et Klaus) Jansonius, Tuzhikova, pl. XXXI, figs. 1–9 (without description). Remarks: The specimen on Plate VI, 2 .71 ð 46 µm) appears different from the specimens on Plate VI, 3–5 in having a subcircular body and thicker exoexine (including sacci), but it still shares some common features. Therefore, it is identified with reservation. K. schaubergeri is generally considered to be an Upper Permian index form, but it has also been recorded from the Lower Triassic in the Urals (Tuzhikova, 1985) and in the Dalongkou section of Xinjiang, NW China (Qu and Wang, 1986). Klausipollenites angustus Ouyang et Norris, sp. nov. (Plate VI, 10–13; Plate VII, 4, 5; Fig. 8) ?1965 Klausipollenites schaubergeri (Potonie´ and Klaus) auct. non Jansonius; Reinhardt and Schmitz, p. 24, pl. 8, fig. 11. 1984 Pityosporites nigracristatus auct. non Hennelly, 1959; Miao et al., p. 582, pl. 178, fig. 17.

?1986 Sulcatisporites rhombicus auct. non Qu, 1984; Qu and Wang, p. 168, pl. 36, fig. 23. Holotype: Sample-slide=coordinate: AEA 755-1= 41 ð 97.6 (Plate VII, 5). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province. Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: Latin, angustus, narrow. Description: Haploxylonoid bisaccate pollen. Overall size (breadth ð length) 46–58 ð 42–48 µm; body 34–42 ð 42–48 µm; sacci 22–36 ð 34–41 µm, projecting 6–12 µm beyond the body (7 specimens); holotype 52 µm. Central body subcircular in outline with a slightly shorter transverse axis. Exine 1–2 µm in thickness, very fine intramicroreticulate-punc-

PLATE VI Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1. Voltziaceaesporites heteromorphus Klaus, 1964. AEA 751-2, 42.1=99.2 2. Klausipollenites sp. cf. K. schaubergeri (Potonie´ et Lele) Jansonius, 1962. AEA 751-7, 44=103 3–5. Klausipollenites schaubergeri (Potonie´ et Lele) Jansonius, 1962. 3. AEA 755-2, 33.4=95.8. 4. AEA 755-1, 33.1=96.1. 5. AEA 755-2, 35.9=110.1. 6. Platysaccus triassicus Ma¨dler, 1964. AEA 749-1, 40=108.7 7. Falcisporites zapfei (Potonie´ et Klaus) Leschik, 1956. AEA 749-1, 34=103.9 8, 9. Pityosporites sp. 8. AEA 755-2, 39=110.1. 9. AEA 755-2, 39=99.6. 10–13. Klausipollenites angustus Ouyang et Norris, sp. nov. 10. AEA 755-2, 33.2=111. 11. AEA 749-7, 39=107.1. 12. AEA 755-1, 35.1=97.9. 13. AEA 755-1, 38.7=112. 14–17. Falcisporites sublevis (Liuber) Ouyang et Norris, comb. nov. 14. AEA 749-2, 35=96. 15. AEA 749-2, 34.5=95.3. 16. AEA 755-2, 44=101.3. 17. AEA 749-6, 38.2=98.3.

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PLATE VI

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PLATE VII

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Fig. 8. Drawing of Klausipollenites angustus Ouyang et Norris, sp. nov. ð600.

tate with muri and lumina D 0.5 µm across. Sacci larger than semicircular, attached equatorially but strongly inclined distally. Roots of sacci straight or slightly concave to convex, distance between them very narrow (generally 2–5 µm) and demarcated

PLATE VII Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1–3. Platysaccus alatus (Liuber ex Kara-Murza) Ouyang et Norris, comb. nov. 1. AEA 749-4, 32=107.8. 2. AEA 755-2, 36.2=95.3. 3. AEA 751-7, 33.1=107.9. 4, 5. Klausipollenities angustus Ouyang et Norris, sp. nov. 4. AEA 755-1, 42.2=98. 5. Holotype, AEA 755-1, 47=97.6. 6–9. Alisporites sp. cf. A. grauvogeliae Klaus, 1964. 6. AEA 751-6, 29.6=107. 7. AEA 755-2, 36.5=110. 8. AEA 751-6, 32=109.1. 9. AEA 751-2, 42.8=108.7. 10, 11. Alisporites exilis Ouyang et Norris, sp. nov. 10. AEA 755-1, 45=100. 11. Holotype, AEA 755- 2, 37.1=107.1. 12–15. Chordasporites rhombiformis Zhou, 1980. 12. AEA 751-8, 38.3=106.3. 13. AEA 751-4, 38.1=106.3. 14. AEA 749-8, 44=102. 15. AEA 749-6, 33.2=104.7. 16. Lueckisporites virkkiae Potonie´ et Klaus, 1954. AEA 749-6, 28.2=104.7. 17–20. Scutasporites sp. cf. S. unicus Klaus 1963. 17. AEA 755-2, 43=102.3. 18. AEA 749-4, 37=102.3. 19. AEA 749-2, 27.2=97.8. 20. AEA 749-2, 35.3=106.2. 21, 22. Angustisulcites gorpii Visscher, 1966. 21. AEA 749-8, 30.3=103.2. 22. AEA 755-1, 45.1=103. 23. Lunatisporites permotriassicus (Maljavkina) var. permotriassicus (Jansonius et Hills) Ouyang et Norris, comb. nov. AEA 749-3, 36=108.7.

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often by root-exine folds of various widths, enclosing typically a narrow leptoma. Saccus wall <1 µm thick, irregularly intramicroreticulate-punctate with muri and lumina 0.5–1.5 µm across, some of the lumina enlarged due to mutual coalescence. Sacci merge with the body smoothly or at a slight angle. Comparison: The present specimens are closely comparable to those recorded by Qu and Wang (1986 p. 168, pl. 36, fig. 23) from the Jiucaiyuan Formation in the Dalongkou section and misidentified as Sulcatisporites rhombicus Qu, 1984 (in Miao et al., 1984, p. 589, pl. 177, figs. 9, 10). The holotype of the latter is characterized, however, by a much narrower central body and relatively broader (8–11 µm) fusiform saccus, thus differing significantly from Qu and Wang’s Xinjiang specimen. Reinhardt and Schmitz (1965) identified a species as K. schaubergeri from the Upper Buntsandstein of Germany, but this identification seems somewhat doubtful, for the species as illustrated by Potonie´ and Klaus has a wider distal zone between the roots. In spite of the presence of a very narrow distal zone in Reinhardt and Schmitz’s specimen, we are not sure that it is synonymous with our new species because it is possibly protosaccate. This new species differs from Alisporites australis De Jersey, 1962 (p. 8, pl. 2, fig. 14; pl. 3, figs. 3, 4) in having less expanded sacci and a narrower distal zone; and it differs from Alisporites townrowii Helby, 1967 (p. 68, pl. 2, figs. 29–32) in its much smaller size and less expanded sacci. Genus Platysaccus (Naumova, 1937) ex Potonie´ and Klaus, 1954 Platysaccus triassicus Ma¨dler, 1964 (Plate VI, 6) 1964 Platysaccus triassicus Ma¨dler, p. 71, pl. 4, fig. 13. ?1956 Coniferaletes imperspicuus Andreeva in Adreeva et al., p. 268, pl. LX, fig. 110. ?1965 Pityosporites imperspicuus (Andreeva) Hart, p. 57, fig. 131. Remarks: The specimen .81 ð 56 µm) is closely similar to P. triassicus Ma¨dler, 1964 from the Upper Buntsandstein of Germany, but the latter is slightly larger (110 µm). According to Hart’s restated diagnosis and text figure, his new combination Pityosporites imperspicuus (Andreeva) is possibly a senior synonym of Platysaccus triassicus Ma¨dler, but this is difficult to ascertain to judge from the description and

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poor drawing by Andreeva et al. (1956). Our specimen differs from Platysaccus leschikii Hart, 1960 (pl. 3, fig. 38) in having a much wider distal zone. Platysaccus alatus (Liuber ex Kara-Murza, 1952) Ouyang et Norris, comb. nov. (Plate VII, 1–3) Basionym: Protopodocarpus alatus (Liuber) ex Kara-Murza, 1952, Tr. Inst. Geol. Arktiki 31, 30, pl. 6, figs. 6, 7. 1953 Protopodocarpus alatus (Liuber) Kara-Murza, Samoilovich, pl. XII, fig. 3. 1967 Indeterminate form, Fradkina, pl. II, fig. 12 (without description). 1985 Podocarpus sp., Tuzhikova, pl. XXX, fig. 9 (without description). 1986 Podocarpidites granulatus auct. non Singh, 1971; Qu and Wang, p. 169, pl. 36, fig. 9. Description: Diploxylonoid bisaccate pollen, amb dumbbell-shaped. Overall size 45–58 ð 26–30 µm; body 23–33 ð 18–24 µm; sacci 19–25 ð 26–30 µm projecting 13–(8.5) 16 µm beyond the body (4 specimens). Central body oval-subcircular in outline with longer transverse axis. Exine rather thick, with proximal cap up to 1.5–3 µm thick, punctateintramicroreticulate with muri and lumina 0.5 µm across. Sacci larger than semi-circular to sub-circular, larger than body; proximal roots of sacci attached subequatorially; distal roots convex or nearly straight separated by ca. 1=4–1=6 body breadth; saccus wall moderately thick, intramicroreticulate with muri and lumina 0.5–1 µm across. Comparison: There is no essential difference between the present specimens and those described from the Permian of the Taymir Basin by KaraMurza (1952) under the name Protopodocarpus alatus (Liuber, 1940) both in size and in general morphology. Our specimens are also very similar to those listed in the synonyms, found in the Upper Permian of the Cis-Urals (Samoilovich, 1953), and the Lower–Middle Triassic of northeastern Siberia (Fradkina, 1967), the Urals (Tuzhikova, 1985) and Xinjiang (Qu and Wang, 1986). Protopodocarpus Bolchovitina, 1956 is a haploxylonoid, monosulcate taxon (see Jansonius and Hills, 1972–1976, Card 2171). Consequently the species is transferred to Platysaccus.

Genus Falcisporites Leschik, 1956 emend. Klaus, 1963 Falcisporites sublevis (Liuber, 1941) Ouyang et Norris, comb. nov. (Plate VI, 14–17) Basionym: Pemphygaletes sublevis Liuber, in Liuber and Val’ts, 1941, Trans All-Un. Sci.Res. Inst. Geol. 139, 57, pl. XIII, fig. 219. 1965 Piceapollenites sublevis (Liuber et Val’ts) Hart, p. 62, fig. 142. 1978 Alisporites sublevis (Liuber) Chen, p. 436, pl. 125, fig. 16. 1986 Alisporites sublevis (Liuber) Chen, in Hou and Wang, p. 97, pl. 22, fig. 16; pl. 27. figs. 17–19. 1986 Alisporites parvus auct. non de Jersey; Qu and Wang, p. 167, pl. 36, figs. 13, 14; pl. 40, fig. 10. Description: Haploxylonoid bisaccate pollen, amb oval-subcircular. Overall size (breadth ð length) 44 (48) 53ð33 (37) 41 µm; body 21 (27) 31ð32 (37) 41 µm; sacci 19–20ð33 (37) 40 µm, projecting (8) 10– 13 µm beyond the body (6 specimens). Central body oval-subcircular with shorter transverse axis; exine often thin, <1 µm thick, only slightly thickened up to 1–1.5 µm along the proximal roots of the sacci. Sacci larger than or equal to semicircular in outline, smaller than or nearly equal to the size of body. Proximal roots of sacci disposed equatorially; distal roots distinct, often concave (occasionally straight) enclosing a fusiform leptoma ca. 1=3 body breadth. Saccus wall also thin, intramicroreticulate-punctate with muri and lumina 0.5 µm across, more distinct than in central body. Comparison: The present specimens are close to Pemphygaletes sublevis Liuber first described from the Permian of the western Pre-Urals of Russia, and those listed in the synonyms mostly from the Upper Permian of China, and the Lower Triassic in Xinjiang. The species is similar to Alisporites parvus De Jersey, 1962 (p. 9, pl. 4, figs. 1–4), but differs from the latter in having a clearly delineated fusiform leptoma and thickened exine at the proximal roots. The reason we transfer the species to Falcisporites is that the genus Alisporites comprises species having sacci joining the body with re-entrant outline and=or with a distinct leptoma; and the genus Piceaepollenites Potonie´, 1931 embraces larger (>70 µm) pollen with a more or less well developed cap.

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diploxylonoid outline, very thin exine and narrow leptoma; due to its diploxylonoid configuration and distinct leptoma it is assigned to Alisporites. Genus Scutasporites Klaus, 1963 Fig. 9. Drawing of the holotype of Alisporites exilis Ouyang et Norris, sp. nov. ð600.

Genus Alisporites Daugherty, 1941 emend. Jansonius, 1971 Alisporites exilis Ouyang et Norris, sp. nov. (Plate VII, 10, 11; Fig. 9) Holotype: Sample-slide=coordinate: AEA 755-2= 37.1 ð 107.1 (Plate VII, 11; Fig. 9). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province. Age: Early Triassic. Formation: Jiucaiyuan Formation. Etymology: Latin, exilis, small, delicate. Description: Slightly diploxylonoid bisaccate pollen, amb oval to slightly dumbbell-shaped. Overall size 39–43 ð 31–38 µm; body 28–29 ð 29–35 µm; sacci 19–20 ð 32–38 µm projecting 6–10 µm beyond the body (4 specimens). Holotype 39 ð 31 µm. Central body subcircular in outline with a slightly longer transverse or longitudinal axis, exine thin, <1 µm, very fine intramicroreticulate with muri and lumina <0.5 µm across. Sacci larger than semicircular, nearly equal in size to body. Sacci disposed equatorially, but strongly inclined distally, distal roots straight and close to each other, delineating a very narrow leptoma, 1–4 µm broad. Saccus wall thin, <1 µm across. Comparison: The new species is somewhat similar to Klausipollenites angustus sp. nov. described above, but differs in being smaller, having a diploxylonoid tendency, more expanded sacci and a thinner and more closely circular central body. Pityosporites lucidus (Leschik, 1956) Hart, 1965 (p. 59, fig. 135) also has a very narrow distal zone, but it differs in being much larger (90 µm). The species is distinguished from other species of Alisporites, Sulcatisporites, or Vesicaspora (cf. e.g. Hart, 1965; Foster, 1979; Tuzhikova, 1985) by its small size, slightly

Scutasporites sp. cf. S. unicus Klaus, 1963 (Plate VII, 17–20) cf. 1963 Scutasporites unicus Klaus, p. 290, pl. 7, figs. 30–32. 1972 Taeniaesporites ortsei auct. non Klaus; in Molin and Koloda, p. 32, pl. 21, fig. 1. 1980 Scutasporites sp. cf. S. unicus Klaus, Balme, p. 32, pl. 3, fig. 5. Remarks: The present specimens are undoubtedly conspecific with those from the Tatarian of the Russian platform (Molin and Koloda, 1972) and the Upper Permian (Dzhulfian?) of Greenland (Balme, 1980). Genus Lueckisporites Potonie´ et Klaus, 1954 emend. Klaus, 1963 Lueckisporites virkkiae Potonie´ et Klaus, 1954 (Plate VII, 16) Remarks: L. virkkiae has been considered to be an index species of the Upper Permian (e.g. Visscher, 1973). However, it has also been recorded from the Lower Triassic in Madagascar (Goubin, 1965), in Austria (Singh, 1965), in the Urals of U.S.S.R. (Tuzhikova, 1985), in Xinjiang (Qu and Wang, 1986), in Zhejiang (Ouyang and Utting, 1990) and in East Greenland (Balme, 1980) where it was considered to be reworked. We consider that it is likely that the parent plants might have persisted into the Early Triassic as relicts in refugia. Genus Lunatisporites Leschik, 1955 emend. Scheuring, 1970 Lunatisporites leptocorpus (Qu, 1984) Ouyang et Norris, comb. nov. (Plate VIII, 3) Basionym: Taeniaesporites leptocorpus Qu in Miao et al., 1984, Paleontological Atlas of North China, III. Geol. Publ. House, Beijing, p. 571, pl. 172, figs. 10–11. 1986 Taeniaesporites leptocorpus Qu in Qu and Wang, p. 162, pl. 35, fig. 18.

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PLATE VIII

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Description: Haploxylonoid bisaccate pollen. Overall size (breadth ð length) 73–91 ð 43–56 µm; body 53–57 ð 40–51 µm; sacci 27–31 ð 40–56 µm projecting 11–17 µm beyond the body (2 specimens). Central body subcircular to broadly oval in outline, with longer transverse axis, exine thin, often <1 µm thick, proximally with 4–5 transverse taeniae, (5.5) 8.5–13 µm in breadth, spaced 1–3 (4.5) µm apart. Sacci somewhat larger than semicircular, disposed equatorially but inclined distally, intramicroreticulate; muri and lumina 0.5–1.5 µm across, often elongated due to coalescence, being smaller on the body (and ribs), ca. 0.5 µm across. Distal roots straight or slightly concave separated by a distance of about 1=2 body breadth. Comparison: The present specimens are quite close to those reported from the upper Lower Triassic Shaofanggou Formation in the Dalongkou section by Qu and Wang (1986) except that in the latter the ribs on the central body are more widely spaced. The species differs from other species of Lunatisporites

PLATE VIII Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1, 4, 5. Lunatisporites permotriassicus (Maljavkina) var. permotriassicus (Jansonius et Hills) Ouyang et Norris, comb. nov. 1. AEA 749-1, 43=103.5. 4. AEA 755-2, 28.2=100.4. 5. AEA 751-4, 34.3=103.7. 2. Lunatisporites sp. cf. L. hexagonalis (Jansonius) Ouyang et Norris, 1988, AEA 751-4, 43=106. 3. Lunatisporites leptocorpus (Qu) Ouyang et Norris, comb. nov. AEA 751-4, 43=106. 6–8. Protohaploxypinus sp. cf. P. perfectus (Naumova ex Kara-Murza) Samoilovich, 1953. 6. AEA 751-1, 34=106.5. 7. AEA 751-1, 36.1=110.1. 8. AEA 751-1, AEA 36=109.3 9. Lunatisporites sp. A. AEA 751-6, 29.8=106.8 10–12. Protohaploxypinus latissimus (Liuber) Samoilovich, 1953. 10. AEA 749-8, 31=106. 11. AEA 749-2, 30=95.3. 12. AEA 755-2, 40.8=109. 13, 14. Protohaploxypinus sp. cf. P. goraiensis (Potonie´ et Lele) Hart, 1964. 13. AEA 751-6, 34.1=100. 14. AEA 751-7, 30=103.8

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(Taeniaesporites) by its thin body wall and indistinct distal saccus root (e.g. Hart, 1965; Balme, 1970). Here we use the name Lunatisporites instead of Taeniaesporites for the reasons stated by Scheuring (1970); thus a new combination is proposed. Lunatisporites permotriassicus (Maljavkina, 1964) var. permotriassicus (Jansonius et Hills) Ouyang et Norris, comb. nov. (Plate VII, 23; Plate VIII, 1, 4, 5) Basionym: Striatipites permotriassica var. parvistriata sp. et var. nov., Maljavkina, 1964, Spores and Pollen from the Triassic Sediments of West Siberian Depression. Nedra, Leningrad, p. 197, pl. 4, fig. 9. 1954 Striatopiceipites sp., Zoricheva and Sedova, pl. IX, fig. 1. 1986 Taeniaesporites pellucidus auct. non Goubin; Hou and Wang. p. 107, pl. 30, fig. 5. (without description). 1986 Protohaploxypinus sp., Qu and Wang, pl. 34, fig. 20 (without description). Description: Haploxylonoid to somewhat diploxylonoid bisaccate pollen, amb oval to dumbbellshaped. Overall size (breadth ð length) 77–87 ð 43– 52 µm; body 39–57 ð 40–43 µm; sacci 30–32 ð 43– 52 µm projecting 11–19 µm beyond the body (4 specimens). Central body subcircular to oval in outline with longer transverse or longitudinal axis, exine ca. 1 µm in thickness proximally with 4–5 low and planar taeniae, 7–13 µm in breadth, unevenly intramicroreticulate-punctate on the taeniae, with muri and lumina 0.5 µm. Taeniae usually quite closely spaced with cleft 1–1.5 µm wide. Sacci disposed equatorially but inclined distally, larger than semicircular, distal roots straight or slightly concave, distance between the roots ca. 1=2–2=3 body breadth. Saccus exoexine ca. 1 µm thick, imperfectly microreticulate-punctate, with muri and lumina often 1 µm, or elongated radially. Comparison: The present specimens are closely similar to cited above which are from the Upper Permian and Lower Triassic of Xinjiang, China and western Siberia, and the Upper Permian of north European Russia. This species is characterized by closely spaced taeniae, and, according to our observation, by the almost equal thickness of body and saccus wall. By these features it is distinguished from the other species of Lunatisporites (D Taeniae-

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sporites) and Protohaploxypinus, in addition to the smaller number of taeniae. Lunatisporites pellucidus (Goubin) differs in having a rather transparent distal zone and wider cleft between the ribs, and L. acutus (Leschik) Scheuring, 1970 differs in having a round central body and monolete or bilete mark between the central ribs. Remarks: Jansonius and Hills stated (1976, card 2741) “the change of name from parvistriatus to permotriassicus is obligate according to the International Code”, i.e. they expressed the name as S. permotriassicus Malj. var. permotriassicus Malj. Judging from the figure of the type specimen (ca. 70 µm), the pollen appears to have only 5 taeniae with somewhat broader clefts (thus Striatipites Maljavkina, 1964 is probably a junior synonym of Lunatisporites Leschik, 1955, which is the basis for our new combination), has a slightly diploxylonoid outline, and the distance between the distal roots is about 1=3 the body breadth. Therefore we consider it indistinguishable from other striate bisaccate genera, especially Lunatisporites. Maljavkina mentioned “ribs broad or narrow, numbering up to 20”; obviously this description is based on heterogeneous material. According to the only figure (type), we prefer to consider that her genus is a synonym of Lunatisporites. Consequently a new combination is proposed. Genus Protohaploxypinus Samoilovich, 1953 Protohaploxypinus latissimus (Liuber) Samoilovich, 1953 (Plate VIII, 10–12) 1941 Pemphygaletes latissimus Liuber in Liuber and Val’ts, p. 158, pl. XIII, fig. 221. 1965 Protohaploxypinus latissimus (Liuber) Samoilovich, Hart, p. 26, fig. 53. 1972 Striatohaploxypinites latissimus (Liuber) Sauer (sic), Molin and Koloda, pl. VI, fig. 1. Remarks: In general aspect, the present specimens are more or less midway between P. latissimus and P. pennatulus (Andreeva) Hart, 1964, but differ from the latter in having comparatively smaller sacci (not larger than the body) and a broader distal zone. Hart (1965, p. 26, fig. 53) defined P. latissimus as having ten ribs, but judging from the figure given by Liuber and Val’ts (1941, pl. XIII, fig. 221) it is probable that ribs in the holotype may reach 14 in number. Thus

our specimens are closest to P. latissimus. The type material is from the Permian of the Pre-Urals. Protohaploxypinus sp. cf. P. perfectus (Naumova ex Kara-Murza 1952) Samoilovich, 1953 (Plate VIII, 6–8) 1952 Platysaccus perfectus ex Kara-Murza, p. 99, pl. 22, fig. 5. Remarks: The present specimens are comparable to Platysaccus perfectus (Naumova) var. substriatus Kara-Murza (1952) from the Upper Permian of the Taymir Basin in overall size, outline, and the obliquely arranged ribs which, however, are more indistinct in the Russian material. Hart (1965, p. 27) defined the species as having 6–8 ribs. Protohaploxypinus sp. cf. P. goraiensis (Potonie´ et Lele) Hart, 1964 (Plate VIII, 13, 14) 1961 Lunatisporites goraiensis Potonie´ et Lele, p. 32, pl. 3, figs. 70–73. 1964 Protohaploxypinus goraiensis (Potonie´ et Lele) Hart, p. 1180, fig. 13. 1965 Protohaploxypinus goraiensis (Potonie´ et Lele) Hart, p. 29, fig. 61. 1970 Protohaploxypinus goraiensis (Potonie´ et Lele) Hart, Balme, p. 362, pl. 11, figs. 1–3. Remarks: Among the known species of Protohaploxypinus or Striatoabieites Sedova emend. Hart, 1964 (cf. e.g. Hart, 1965; Balme, 1970; Scheuring, 1970, 1978), the present specimens are more or less comparable to P. goraiensis, Striatiabieites brickii Sedova, 1956 (see Hart, 1965, p. 40, fig. 87) and P. pennatulus (Andreeva, 1956) Hart, 1964 which Balme (1970) considers “may be indistinguishable from P. goraiensis”. However, they differ from P. goraiensis and P. pennatulus in having a broader distal zone (cappula in Balme’s sense) and from S. brickii in having larger sacci and a haploxylonoid outline (the sacci in S. brickii join the body with an angle). We tentatively assign our specimens to P. goraiensis because they are most similar to those from the Permian of West Pakistan described by Balme (1970) as P. goraiensis in such common features as haploxylonoid outline, larger overall size, relatively larger number of taeniae and particularly the saccus wall being thicker than that of the body. Balme (1970, p. 364) described the “cappula ... less than 1=5 that of corpus, usually very nar-

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row, ... ” (but judging from his photographs, the cappula seems to be ca. 1=3 breadth of the body). Consequently, P. goraiensis is somewhat different compared with our specimens. Protohaploxypinus sp. cf. P. samoilovichiae (Jansonius) Hart, 1964 (Plate IX, 1–3) 1962 Striatites samoilovichii Jansonius, p. 67, pl. 14, fig. 9. Remarks: The present specimens, especially that on Plate IX, 2, differ from the known species of Protohaploxyinus in certain details. They are similar in some features to P. samoilovichiae (Jansonius) Hart, 1965 (p. 31, fig. 66) such as in general outline, size and rib number. However, the type material described by Jansonius (1962, p. 64, pl. 14, figs. 9–11) from the Lower Triassic of Canada displays broader ribs and rib spacing, and sacci with larger microreticulation, each saccus being smaller than the body. P. pennatulus (Andreeva) Hart has a much narrower distal zone. Genus Striatopodocarpites Sedova, 1956 emend. Hart, 1964 Striatopodocarpites sp. cf. P. pantii (Jansonius) Balme, 1970 (Plate IX, 5, 8) 1962 Striatites samoilovichii var. pantii Jansonius, p. 68, pl. 14, figs. 14, 15. ?1965 Stroterisporites pantii (Jansonius) Goubin, p. 1424, pl. 2, figs. 7, 8. ?1970 Striatopodocapites pantii (Jansonius) Balme, p. 368, pl. 12, figs. 7–9. ?1986 Striatopodocarpites sp., Hou and Wang, pl. 29, fig. 7 (without description). ?1986 Striatoabieites duivenii (Jansonius) Hart, in Qu and Wang, pl. 34, fig. 6 (without description). Remarks: The present specimens are comparable with those described by Balme (1970, p. 368, pl. 12, figs. 7, 8 and especially fig. 9) from the Permian of West Pakistan although in the latter the saccus wall is thicker (ca. 3 µm) and is “fairly coarsely intrareticulate”, and the distal zone is somewhat narrower (1=3 body breadth). Our specimens are also similar to Striatoabieites brickii Sedova, 1956 (see Hart, 1965, p. 40, fig. 87) but differ in having fewer taeniae and larger sacci. Hou and Wang (1986) and Qu and Wang

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(1986) reported the presence of Striatopodocarpites sp. and Striatoabieites brickii from the upper Upper Permian and the Lower Triassic in the Dalongkou section, respectively; judging from the figures listed in the synonymy, they seem to be synonymous with our present specimens. Hou and Wang (1986, p. 104, pl. 29, fig. 13) described a specimen under the name Striapodocarpites pantii (Jansonius) Balme which shows 12–14 taeniae and appears closer to Striatoabieites brickii than to S. pantii. Genus Hamiapollenites Wilson, 1962 Hamiapollenites bullaeformis (Samoilovich) Jansonius, 1962 (Plate X, 1–3) 1953 Protodiploxypinus bullaeformis Samoilovich, p. 33, pl. IV, fig. 1a, b. 1962 Hamiapollenites bullaeformis (Samoilovich) Jansonius, p. 71. 1965 Hamiapollenites bullaeformis (Samoilovich) Jansonius, Hart, p. 49, fig. 110. 1981 Striatodiploxypinites bullaeformis (Samoilovich) Zauer in Varjukhina, pl. XVII, fig. 5a, b. 1983 Protowelwitschiapollis exilis Zhang, p. 334, pl. 5, fig. 5, 6, 10. 1983 Protowelwitschiapollis exolescus Zhang, p. 334, pl. 5, figs. 4, 7, 9, 12. 1986 Hamiapollenites bullaeformis (Samoilovich) Jansonius in Hou and Wang, p. 105, pl. 24, fig. 17; pl. 30, figs. 12, 13. 1986 Hamiapollenites bullaeformis (Samoilovich) Jansonius in Qu and Wang p. 159, pl. 34, fig. 11. Remarks: The specimen on Plate X, 3 .65 ð 45 µm, with 7 proximal ribs) is transitional between H. bullaeformis and H. tractiferinus (Samoilovich) Jansonius, 1962 emend. Hart, 1964. Samoilovich (1953) mentioned that bullaeformis and tractiferinus are very similar, and “they differ in the shape of the body (spherical or almost spherical in P. bullaeformis and ellipsoidal in P. tractiferinus)”. Thereafter, Hart selected pl. 12, fig. 2b of Samoilovich as the holotype of H. tractiferinus and redefined it to embrace forms with 6–8 proximal ribs, by which the species differs from H. bullaeformis (10–12 ribs). However, in our experience, neither the body shape nor rib number can be used to separate the two species.

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PLATE IX

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Therefore, we have tentatively identified the specimen on Plate X, 3 as bullaeformis. Zhang (1983) established a new genus Protowelwitchiapollis (with type species P. exilis Zhang) without mentioning its difference from Hamiapollenites Wilson, 1962. Judging from her description and photographs, the type species and another species are most likely synonymous with P. bullaeformis. This species occurs in the Upper Permian (Zhang, 1983; Hou and Wang, 1986) and Lower Triassic in Xinjiang.

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Vittatina subsaccata Samoilovich, 1953 (p. 44, pl. IX, fig. 4a, b) by different authors (e.g. Barss, 1967, pl. 37, fig. 13; Dybova-Jachowicz, 1981, pl. X, figs. 1–3; Balme, 1980 p. 28, pl. 2, fig. 7), but close comparison and identification is very difficult due to the fact that various authors have their own understandings of the circumscription and line-drawings by Samoilovich (cf. discussion by Wilson, 1962, p. 24) and of the separation of several similar species. Thus we use the name with reservation.

Genus Vittatina Liuber, 1940 emend. Wilson, 1962

Vittatina sp. cf. V. striata (Liuber) Samoilovich, 1953 (Plate IX, 6)

Vittatina sp. cf. V. subsaccata Samoilovich, 1953 (Plate IX, 10–12)

1941 Azonaletes striatus Liuber, in Liuber and Val’ts, p. 156, pl. XIII, fig. 218.

1953 Vittatina subsaccata Samoilovich, p. 44, pl. IX, fig. 4a. Remarks: The present specimens, especially that on Plate IX, 11, are comparable to those identified as

PLATE IX Each entry comprises the sample number, slide number and coordinates. All figures ð600. 1–3. Protohaploxypinus sp. cf. P. samoilovichiae (Jansonius) Hart, 1964. 1. AEA 751-1, 36.8=109.9. 2. AEA 749-5, 36.6=99.2. 3. AEA 749-5, 34.3=105.6. 4. Striatopodocarpites sp. AEA 751-1, 34=96.8. 5, 8. Striatopodocarpites sp. cf. S. pantii (Jansonius) Balme, 1970. 5. AEA 755-1, 32=109. 8. AEA 751-3, 33.1=107.9. 6. Vittatina sp. cf. V. striata (Liuber) Samoilovich, 1953, AEA 751-2, 38.2=101.3. 7. Lunatisporites sp. A, AEA 751-5, 33.5=104. 9. Vittatina sp., AEA 749-3, 33.3=107.8. 10–12. Vittatina sp. cf. V. subsaccata Samoilovich, 1953. 10. AEA 751-2, 42=99. 11. AEA 749-8, 39.5=102.8. 12. AEA 751-2, 31.2=107.2. 13, 14. Hamiapollenites ruditaeniatus Qu et Wang, 1986. 13. AEA 751-4, 36.1=109.4. 14. AEA 749-5, 33=105.8. 15, 16. Cycadopites caperatus (Liuber) Hart, 1965. 15. AEA 749-2, 33.6=105.3. 16. AEA 751-6, 37=109.5. 17, 18. Pilasporites trigonius (Djupina) Tuzhikova, 1985. 17. AEA 755-1, 40.1=110.2. 18. AEA 755-1, 39.2=111.8.

Remarks: This specimen .43 ð 41 µm) is more or less comparable to the species striata as identified by different authors (Varjukhina, 1971, p. 88, pl. VII, fig. 5a–c; pl. 14, fig. 4; Molin and Koloda, 1972 pl. XVI, fig. 2; Tuzhikova, 1985, pl. XXV, fig. 14; Balme, 1980, p. 28, pl. 2, fig. 11). Pollen grains of this type are abundant in the Upper Permian of European U.S.S.R., and are rarely found in the Lower Triassic (Tuzhikova, 1985). Genus Cycadopites (Wodehouse, 1933) ex Wilson and Webster, 1946 Cycadopites retroflexus (Liuber) Hart, 1965 (Plate X, 6, 7) 1941 Azonaletes retroflexus Liuber, in Liuber and Val’ts, p. 179, pl. XVI, fig. 252a, b. ?1952 Subsacculifer obliquus Kara-Murza, p. 20, pl. 3, figs. 1, 2. 1953 Ginkgocycadophytus retroflexus (Liuber) Samoilovich, pl. III, fig. 7a, b (without description). 1956 Azonaletes retroflexus Liuber, in Andreeva et al., p. 265, pl. LVIII, figs. 102c (in part), 102d. 1965 Cycadopites retroflexus (Liuber) Hart, p. 107, fig. 258. Remarks: The present specimens are close to the type material described by Liuber (in Liuber and Val’ts, 1941) except that in the latter “the body ... is smooth, while the lateral crumplings are often reticulate-chagrenate” (the author assumed them to be sacci of conifers). Judging from the figures, it seems that the

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PLATE X

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pollen exine (‘body’) shows some kind of texture or sculpture and as for the ‘sacci’ (crumplings), we think they are actually parts of the distal exine flanking the sulcus. The specimens described by Kara-Murza (1952) under the name Subsacculifer obliquus appear to be conspecific with C. retroflexus (Liuber) although she mentioned that her new species differs from the latter in having less distinct development of ‘sacci’ and especially in having levigate exine. C. retroflexus is abundant in the Permian of Russia (e.g. Kuznetsk and Minusinsk basins). Genus Pilasporites Balme et Hennelly, 1956 emend. Jain, 1968 Pilasporites trigonius (Djupina) Tuzhikova, 1985 (Plate IX, 17, 18) 1974 Urmites trigonius Djupina, pl. VI, figs. 13–17. 1985 Pilasporites trigonius (Djupina) Tuzhikova, p. 115, pl. XXVI, figs. 27, 28.

PLATE X Each entry comprises the sample number, slide number and coordinates. All figures ð600 unless otherwise indicated. 1–3. Hamiapollenites bullaeformis (Samoilovich) Jansonius, 1962. 1. AEA 751-5, 40=102.3. 2. AEA 751-5, 36.2=96.4. 3. AEA 751-2, 36.6=103.8. 4, 5. Decussatisporites? sp. 4. AEA 749-4, 39.1=106.9. 5. AEA 749-1, 31.2=107. 6, 7. Cycadopites retroflexus (Liuber) Hart, 1965. 6. AEA 751-1, 36.1=105.7. 7. AEA 751-3, 41.8=104.8. 8, 9. Cycadopites sp. 8. AEA 749-2, 29=100.9. 9. AEA 755-2, 45=105. 10–12. Eucommiidites sp. 10. AEA 755-2, 38=102. 11. AEA 749-6, 34.2=100.6. 12. AEA 749-1, 39.2=108. 13, 14. Pilasporites perreticulatus Ouyang et Norris, sp. nov. 13. AEA 749-4, 35=109.3. 14. Holotype, AEA 749-3, 30=104.1. 15a, b. Solisphaeridium? sp. AEA 755-1, 34=104. 16–19. Tympanicysta stoschiana Balme, 1980. 16. AEA 749-2, 37.9=102.3. ð500. 17. AEA 749-5, 34.1=102.8. 18. AEA 749-2, 29.1=99. 19. AEA 749-1, 42.5=104.3.

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1986 Pilasporites crateraformis auct. non Jain, 1968; Qu and Wang, p. 172, pl. 36, fig. 29. Remarks: The present specimens (that on Plate IX, 18 is obliquely compressed) are closely similar to those described and illustrated by Tuzhikova (1985) from the Lower Triassic (Induan) of the Urals except that the exine is slightly thicker and the exine folds along the leptoma edges are broader in the latter. They are also similar to P. crateraformis Jain, 1968 (p. 42, pl. 12, figs. 17–20) from the Middle Triassic of Argentina, but the latter is larger (65–85 µm) and sometimes has a larger irregular ‘pore’. Pilasporites perreticulatus Ouyang et Norris, sp. nov. (Plate X, 13, 14; Fig. 10) Holotype: Samples slide=Coordinate: AEA 749-3, 30=104.1 (Plate X, 14; Fig. 10). Repository: Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, People’s Republic of China. Type locality: Dalongkou, Junggar Basin, Xinjiang Province. Age: Early Triassic Formation: Jiucaiyuan Formation. Etymology: Latin, perreticulatus, very finely reticulate. Description: Monoaperture pollen (?) or acritarch cyst, circular–subcircular to oval in outline. Overall size 36 (47) 54 ð32 (38) 41 µm (6 specimens); holotype 54 ð 41 µm. Exine thin, <1 µm in thickness, very finely reticulate-punctate with muri and lumina 0.5 µm wide. Distally (?) a large leptoma (aperture?) or depression is clearly visible, subcircular–oval in outline and conforming with that of the whole grain. The leptoma is demarcated by a zone of concentric thickenings or folds around its outer margin, up to

Fig. 10. Drawing of the holotype of Pilasporites perreticulatus Ouyang et Norris, sp. nov. ð600.

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2–3 µm broad, and occupies more than 4=5 the area of the distal surface. Comparison: The new species distinguishes itself from other species of Pilasporites (cf. Jain, 1968; Tuzhikova, 1985) by its very thin and finely reticulate-punctate exine. It is somewhat similar to Pilasporites chajcerii Tuzhikova, 1985 (p. 113, pl. XXVI, figs. 20–22; pl. LXII, fig. 1) from the Lower Triassic of the Urals, but the latter has a thicker exine (2 µm) and different sculpture (“with scarcely perceptible fine granulate sculpture or levigate”). Genus Solisphaeridium Pocock, 1965

Staplin,

Jansonius

et

Solisphaeridium? sp. (Plate X, 15) Remarks: Most acanthomorphic acritarchs are generally believed to be of marine origin. Thus the occurrence of Solisphaeridium? sp. in the ‘continental strata’ of Early Triassic age deserves special mention. A reworked origin is possible. Genus Tympanicysta Balme, 1980 Tympanicysta stoschiana Balme, 1980 (Plate X, 16– 19) 1980 Tympanicysta stoschiana Balme, p. 24, pl. 1, fig. 3–7. Remarks: The present specimens are almost exactly the same as those described by Balme (1980) from the Lower Triassic of Greenland except for the occasional occurrence of more prominent folds in the former. As Balme pointed out, Tympanicysta is widespread in Upper Permian and Lower Triassic strata. T. stoschiana has also been recorded from the marine Upper Permian and basal Triassic in the Changxing stratotype sections in Zhejiang Province, eastern China (Ouyang and Utting, 1990). Whether the genus name Tympanicysta Balme, 1980 is a junior synonym of Chordecystia Foster, 1979 or not requires further work.

Acknowledgements Funding for the investigation in the Department of Geology, University of Toronto, Canada was provided by a Natural Sciences and Engineering Re-

search Council of Canada research grant to G. Norris. The authors are indebted to Dr. Liao Zhuo-ting and Dr. Zhou Yu-xing for their help with the field work, and especially to Dr. Shen Yan-bin for supplying the relevant stratigraphic data. Many thanks are also due to Drs. Qu Li-fang, Hou Jing-peng and Wang Zhi for providing copies of their papers. Help by the following colleagues is also acknowledged: Dr. Zhou Yu-xing for sample preparation; the late Mr. Brian O’Donovan for his final effort in printing the photographs; Mr. Zhu Xiao-xing for drafting the figures; Ms. B. Laurent for assistance with the word processing. We should like to thank the reviewers (Drs. B.E. Balme, J.H.A. van Konijnenburg-Van Cittert and J. Jansonius) for their carefully reading of the manuscript and many valuable constructive comments.

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