Review of Palaeobotany and Palynology 152 (2008) 141–157
Contents lists available at ScienceDirect
Review of Palaeobotany and Palynology j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / r ev p a l b o
Biostratigraphic and paleogeographic significance of a palynological assemblage from the Middle Devonian Ulusubasite Formation, eastern Junggar Basin, Xinjiang, China Huaicheng Zhu a,b, Reed Wicander c,⁎, John E.A. Marshall d a
Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China State Key Laboratory of Palaeobiology and Stratigraphy, 39 East Beijing Road, Nanjing 210008, China Department of Geology, Central Michigan University, Mount Pleasant, Michigan, 48859, USA d School of Ocean and Earth Science, University of Southampton, National Oceanography Centre Southampton, European Way, Southampton, SO14 3ZH, United Kingdom b c
A R T I C L E
I N F O
Article history: Received 11 October 2007 Received in revised form 18 April 2008 Accepted 22 April 2008 Available online 10 May 2008 Keywords: Middle Devonian Junggar Basin China acritarch miospore biostratigraphy paleogeography
A B S T R A C T Abundant palynomorphs are here reported for the first time from tuffaceous siltstones of the Middle Devonian Ulusubasite Formation at the Zhifang Section, eastern Junggar Basin, Xinjiang, Northwest China. Thirty-four miospore species assigned to 23 genera (four established species, two cf., 19 sp., and nine spp.), 19 acritarch species assigned to 19 genera (10 established species, three cf., five sp., and one spp.), and scolecodonts are identified. The biostratigraphic ranges of the acritarchs and miospores indicate a Mid Devonian age, corroborating the age assignment previously based on associated corals, brachiopods, and plant fossils. Paleogeographic reconstruction for the Middle Devonian indicates that the eastern Junggar Basin was part of the Kazakhstan Plate and situated in the low-mid latitudes of the Northern hemisphere between the Euramerican landmass to the west, and the North and South China, and Gondwana landmasses to the east and south. Although there is some miospore compositional similarity between the eastern Junggar Basin and the landmasses of Euramerica, South and North China, and Gondwana, it is not particularly high, and is mainly at the generic level. The acritarch assemblage, however, consists mostly of cosmopolitan species and provides new evidence for extending the marine linkage between North America, Baltica, the Junggar Basin of Kazakhstan, and Gondwana. Paleontologic and lithologic evidence indicates that the Ulusubasite Formation was deposited in a near-shore marine environment. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Middle Devonian palynofloras from China have been mainly reported from Yunnan Province, South China where terrestrial facies are well-developed and exposed in the eastern part of that province (Lu, 1980, 1988; Gao, 1981; Zhu and Lu, 2002). Because of an unconformity between the Late Ordovician and Early Carboniferous in the North China Block, Devonian palynological data have been reported only from scattered localities on the southern margin of the block (e.g., Gao and Liu, 1988). Xinjiang is thought to be tectonically separate from South and North China during the Devonian and because of extensive oil and gas exploration since the 1980s, its geology is now becoming better known. Whereas there are several Upper Devonian palynological reports in the literature (Lu and Wicander, 1988; Zhu, 1999, 2000), there have been no reported Middle Devonian palynologic
studies from this region. This study is thus the first report of any Middle Devonian palynomorphs from the eastern Junggar Basin of China. Thirty samples were collected from the Zhifang Section of the eastern region of the Junggar Basin, North Xinjiang by the first author (H. Zhu) for palynological analysis. Eight of those samples yielded abundant palynomorphs including miospores, acritarchs, and scolecodonts. Although palynomorph preservation is less than ideal, this is the first reported Middle Devonian palynologic occurrence from this area of China. Thus, it provides information on both the terrestrial miospores and marine acritarchs for a region that is palynologically depauperate compared to the rest of the world for this time period. Furthermore, despite the generally poor preservation of the miospores, the total recovered palynomorph assemblage provides some useful biostratigraphical and paleogeographical insights during this time interval. 2. Geological setting
⁎ Corresponding author. Tel.: +1 989 774 3179; fax: +1 989 774 2142. E-mail address:
[email protected] (R. Wicander). 0034-6667/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2008.04.010
Xinjiang is the largest provincial region in China and lies in the northwest part of the country. It is geomorphologically subdivided
142
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
Fig. 1. Location of the Zhifang Section (indicated by a ▲), eastern Junggar Basin, Xinjiang, northwest China. The base map and geographic locations are modified from Li et al. (2006, Fig. 2).
into two parts by the Tianshan Mountains, i.e., South Xinjiang and North Xinjiang (Fig. 1). North Xinjiang is mainly represented by the Junggar Basin, which was part of the Kazakhstan Plate and is presently bounded by the Tianshan Mountains in the south and the Altai Mountains in the north. The eight productive palynological samples reported here were collected from the upper part of the Ulusubasite Formation of the Zhifang Section (44° 21′ 48″ N, 91° 50′ 24″ E), Barkol County, Xinjiang, Northwest China (Fig. 1). This section, which is one of two key Devonian sections in the Junggar Basin, is located in the eastern part of the basin. The Junggar Basin constitutes the main part of the Junggar– Beishan subregion of the Junggar–Xing'an stratigraphic Region (the area approximately to the north of 42°N, which includes the Junggar Basin, the Beishan Mountains in northern Gansu Province, most of Inner Mongolia, and the northern part of Heilongjiang Northeast China) of China during the Devonian (Hou et al., 2000). The majority of Devonian rocks in the Junggar area are marine volcanics and massive tuffaceous clastics containing an abundant and diverse marine fauna that includes corals, brachiopods, crinoids, and other marine invertebrates. Occasionally intercalated with these marine beds are fossil plant-bearing rocks. The Devonian in the eastern Junggar Basin is subdivided into the Lower Devonian Taherbasite and Zuomubasite formations, the Middle Devonian Ulusubasite and Zhifang formations, and the Upper Devonian Keankuduke Formation (Fig. 2). Here, the Ulusubasite Formation conformably overlies the Zuomubasite Formation and is disconformably overlain by the Zhifang Formation. At the Zhifang Section, the Ulusubasite Formation is about 117 m thick and is mainly composed of tuffs, tuffaceous sandstones, conglomeratic sandstones, marls, bioclastic limestones, and siltstones (Fig. 3). It yields an abundance of fossils, with many megafossil plants, such as Lepidodendropsis sp., from the uppermost
part of the formation and an abundant and diverse marine fauna from the middle and lower parts, including corals such as Endophyllum sp., Pachyfavosites sp., Crassialveolites sp., and Tyrganolites
Fig. 2. Devonian stratigraphy in the east Junggar Basin, Xinjiang, China, based on Hou et al. (2000). The contact between the Ulusubasite Formation and the underlying Zuomubasite Formation is conformable, whereas it is disconformable with the overlying Zhifang Formation.
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
143
Fig. 3. Stratigraphic section of the Middle Devonian Ulusubasite Formation, showing the lithofacies and location of the nine palyniferous samples (AET 19–AET 27), but only eight of which were productive (shown by a ●), and reported herein.
sp., the brachiopods Leptostrophia sp., Acrospirifer sp., and “Fimbrispirifer” sp., as well as numerous disarticulated crinoid and bryozoan fragments. The marine fauna and plant megafossils indicate an age of Mid Devonian (Eifelian) for the Ulusubasite Formation (Hou et al., 2000). 3. Materials and methods Thirty samples were collected from the siltstones of the upper part of the Ulusubasite Formation. Standard palynological techniques were used to process the samples. These included treatment in hydrochloric acid, hydrofluoric acid, and for selected samples (i.e., those with spores), nitric acid, with the samples being neutralized in distilled
water between each acid treatment. The resulting organic residue was mounted in glycerine jelly and sealed with paraffin wax. Of the 30 samples processed, nine were palyniferous, but only eight yielded abundant palynomorphs (Fig. 3). Because of the poor quality of preservation, selected miospores were reinvestigated using an infra-red microscope to image their internal structure. This is essential when attempting to investigate such opaque high maturity palynomorphs. The microscope used was described by Marshall (1995) except that image acquisition was through a Hauppage! Win TV frame grabber card (Pal-1 Model 40205 Rev B211). The acritarchs generally have thinner walls and are easier to image/observe with routine transmitted light microscopy.
144
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
4. Identified palynomorphs For simplicity, the fossil prasinophycean and probable prasinophycean phycoma (e.g., Cymatiosphaera, Divietipellis, Leiosphaeridia, and Polyedryxium) are included under the informal incertae sedis “group” name Acritarcha, and are arranged alphabetically, as are the miospores. The general poor preservation and high thermal maturation of the miospore assemblage means that this assemblage is not suitable for detailed descriptive work. There are, however, specimens that, although rare, are better preserved, and it is those that were selected for illustration. Caution was also exercised in identifying the miospore species because our knowledge of miospores from the Junggar Basin is still poor. The preservation of the acritarchs is generally better than the miospores, and hence positive specific attribution was usually possible. All figured specimens are housed at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China. 4.1. Miospores Ancyrospora sp. A (Plate II,11,12) Ancyrospora spp. Remarks: Several species of Ancyrospora were recovered. However, the state of preservation was such that we prefer to retain them all in open nomenclature. Aneurospora sp. A (Plate III,10) Anulatisporites sp. Apiculiretusispora sp. Archaeoperisaccus indistinctus Lu, 1988 (Plate II,1–9) Occurrence: Archaeoperisaccus indistinctus has previously been reported from the Givetian of Yunnan, South China (Lu, 1980, 1988), the Givetian–Frasnian of the Canning Basin, Australia (McGregor and Playford, 1993), and the Eifelian–Early Frasnian of the Ardavale Basin, Australia (Hashemi and Playford, 2005). Auroraspora sp. A (Plate I,7) Auroraspora sp. B (Plate I,12) Auroraspora spp. Remarks: Several species assignable to Auroraspora, but not well enough preserved to speciate, were recovered from the Ulusubasite Formation samples. Calamospora sp. Convolutispora sp. ?Cristatisporites sp. A (Plate I,3) Cristatisporites sp. B (Plate I,10,11) Cymbosporites sp. Densosporites sp. Densosporites sp. A (Plate I,13,16,17) Dibolisporites echinaceus (Eisenack) Richardson, 1965 (Plate II,10) Occurrence: Dibolisporites echinaceus has been previously reported from the Eifelian–Givetian of Scotland (Richardson, 1965); Late Emsian–Eifelian of Gaspé, Canada (McGregor, 1977); Late Emsian–
Givetian of Ontario, Canada (McGregor and Camfield, 1976); Eifelian– Givetian of the Canadian Arctic of Canada (McGregor and Camfield, 1982); Emsian–Eifelian of the Eifel region, Germany (Riegel, 1982); Givetian of Poland (Turnau, 1996; Turnau and Racki, 1999); Late Emsian–Early Frasnian of North America (McGregor, 1979b); Givetian– Early Frasnian of Boulonnais, France (Loboziak and Streel, 1980), and the Emsian–Frasnian of Euramerica (McGregor, 1979a; Richardson and McGregor, 1986). Dibolisporites eifeliensis (Lanninger) McGregor, 1973 (Plate III,8,9,12) Occurrence: Dibolisporites eifeliensis has been previously reported from the Emsian–Eifelian of Gaspé, Canada (McGregor, 1973, 1977); Emsian of Ontario, Canada (McGregor and Camfield, 1976); Emsian of North America (McGregor, 1979b), and the Pragian (Siegenian)– Eifelian of Euramerica (Richardson and McGregor, 1986). Dibolisporites cf. D. quebecensis Occurrence: Dibolisporites quebecensis has been previously reported from the Emsian–Early Eifelian of Gaspé, Canada (McGregor, 1973, 1977), and the Emsian of North America (McGregor, 1979b). Dibolisporites spp. Remarks: The remaining species of Dibolisporites recovered were not well enough preserved to make specific identifications, but included two distinct size ranges. ?Diducites sp. Grandispora sp. A (Plate I,1,2) Grandispora sp. B (Plate I,4–6) Grandispora sp. C (Plate I,8,9,18) Grandispora sp. Hystricosporites spp. Leiotriletes spp. Remarks: Numerous species of Leiotriletes are recognized from the studied samples. However, these triangular, laevigate trilete miospores are relatively simple with long geological ranges and wide geographical distribution, hence they are regarded as of little significance in age determination and comparisons, and are not speciated, but left in open nomenclature. Perotrilites spp. Poshisporites minutus Bharadwaj, Tiwari, and Venkatachala, 1973 (Plate I,14,15) Occurrence: Poshisporites minutus has been previously reported from the Middle Devonian of Yunnan, China (Bharadwaj et al., 1973). Punctatisporites spp. (Plate III,11) Remarks: Three species of Punctatisporites are recognized from the studied samples. However, these circular, laevigate trilete miospores are relatively simple and have long geological ranges and wide geographical distribution. They are usually regarded as of little significance in age determination and correlation, and hence are not speciated herein. Retusotriletes spp. Rhabdosporites cf. R. parvulus (Plate III,3,4) Occurrence: Rhabdosporites parvulus has been previously reported from the Givetian of Scotland (Richardson, 1965), whereas Rhabdosporites cf.
Plate I. Specimens are located by sample (AET25), slide (5), and England–Finder coordinates. For example, AET25(6), K27/1 refers to sample AET25, slide 6, England–Finder coordinates K27/1. All specimens are x750 except 13, which is x300. 1, 2. 3. 4–6. 7. 8, 9. 10, 11. 12. 13, 16, 17. 14, 15. 18.
Grandispora sp. A. AET25(6), K27/1. ?Cristatisporites sp. A. AET23(2), V46/2. Grandispora sp. B. AET19(9), X34. Auroraspora sp. A. AET27(4), O42/1. Grandispora sp. C. AET27(5), M29. Cristatisporites sp. B. AET24(8), N42. Auroraspora sp. B. AET24(2), W44/1. Densosporites sp. A. AET25(5), L44. Poshisporites minutus Bharadwaj, Tiwari, and Venkatachala, 1973. AET27(1), J31/3. Grandispora sp. C. AET24(10), M51.
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
145
146
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
147
Plate III. Specimens are located by sample (AET25), slide (5), and England-Finder coordinates. For example, AET25(6), K27/1 refers to sample AET25, slide 6, England-Finder coordinates K27/1. All specimens are ×750 except 4, 7, 11, which are ×300. 1. 2. 3, 4. 5, 6. 7. 8, 9. 10. 11. 12.
Patinate miospore. AET26(4), K41/2. Patinate miospore. AET23(1), S45/3. Rhabdosporites cf. R. parvulus Richardson, 1965. AET19(8), Z39. Patinate miospore. AET22(1), P37/4. Scolecodont. AET24(3), G32. Dibolisporites eifeliensis (Lanninger) McGregor, 1973. AET23(6), D48/1–3. Aneurospora sp. A. AET27(2), Q35. Punctatisporites sp. AET26(5), Q45/3–4. Dibolisporites eifeliensis (Lanninger) McGregor 1973. AET22(9), V39.
R. parvulus has been reported from the Eifelian–Givetian of the Canadian Arctic, Canada by McGregor and Camfield (1982). Verrucosisporites sp. Patinate miospores (Plate III,1,2,5,6)
Remarks: Although unidentifiable to the species level, numerous patinate miospores occurred throughout the Ulusubasite Formation, and thus several representative specimens are illustrated here.
Plate II. Specimens are located by sample (AET25), slide (5), and England–Finder coordinates. For example, AET25(6), K27/1 refers to sample AET25, slide 6, England–Finder coordinates K27/1. All specimens are ×750 except 7–9, 12, which are ×300. 1, 2, 7. 3, 4, 8. 5, 6, 9. 10. 11, 12.
Archaeoperisaccus indistinctus Lu, 1988. AET27(7), E35/1. Archaeoperisaccus indistinctus Lu, 1988. AET27(1), G43/1. Archaeoperisaccus indistinctus Lu, 1988. AET25(4), O44/4. Dibolisporites echinaceus (Eisenack) Richardson, 1965. AET25(6), U30/2. Ancyrospora sp. A. AET23(4), H39.
148
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
4.2. Acritarcha Ammonidium sp. Baltisphaeridium sp. Cymatiosphaera cornifera Deunff, 1955 (Plate IV,10) Occurrence: Cymatiosphaera cornifera has been reported from the Late Pragian (Siegenian)–Givetian of Ontario (Deunff, 1955, 1966, 1967; Playford, 1977); Lochkovian of western Newfoundland (Burden et al., 2002); Eifelian–Givetian of Ohio (Wicander and Wood, 1981; Wicander and Wright, 1983); Middle Devonian of Sichuan Province, China (Gao, 1997), and a possible Early Famennian occurrence from Belgium (Stockmans and Willière, 1969). Diexallophasis remota (Deunff) Playford, 1977 (Plate IV,3) Occurrence: Diexallophasis remota as circumscribed by Playford (1977) is widely distributed in the Northern hemisphere with a reported range of Llandoverian–Givetian (Burden et al., 2002), and is also present in Upper Givetian–Lower Frasnian strata in Argentina (Ottone, 1996). Divietipellis cf. D. ventricosa (Plate IV,13) Occurrence: Divietipellis ventricosa has previously been reported from the Late Pragian (Siegenian)–Early Eifelian of Ontario (Playford, 1977); Givetian of Ohio (Wicander and Wood, 1981), and the Late Givetian– Early Frasnian of Argentina (Barreda, 1986). Ephelopalla sp. A (Plate IV,6) Exochoderma cf. E. arca (Plate IV,1) Occurrence: Exochoderma arca has been previously reported from the Eifelian–Givetian of Kentucky, Ohio, and New York (Wicander and Wood, 1981; Wicander and Wright, 1983; Wood and Clendening, 1985; Wood, 1986; Huysken et al., 1992), and the Late Givetian–Early Frasnian of Argentina (Barreda, 1986; Ottone, 1996). Gorgonisphaeridium cumulatum Playford, 1977 (Plate IV,8) Occurrence: Gorgonisphaeridium cumulatum has only been reported from the Late Pragian (Siegenian)–Givetian of Ontario (Playford, 1977). Hapsidopalla exornata (Deunff) Playford, 1977 (Plate IV,12) Occurrence: This species has been previously reported from the Late Eifelian–Early-Mid Givetian of Ontario (Playford, 1977); probable Frasnian of southern Ontario, Canada (Deunff, 1967); Eifelian–Givetian of Ohio (Wicander and Wood, 1981; Wicander and Wright, 1983); and Late Givetian–Early Frasnian of Argentina (Barreda, 1986; Ottone, 1996). Leiosphaeridia spp. Remarks: Leiosphaeridia spp. occurs in several samples. We have chosen not to speciate these simple, psilate forms, but merely to note their occurrence. There appears to be at least two possible species based on size alone. Multiplicisphaeridium ramusculosum (Deflandre) Lister, 1970 (Plate IV,5) Remarks: Multiplicisphaeridium ramusculosum is a cosmopolitan species with a stratigraphic range of Late Ordovician–Late Devonian (Wicander and Wood, 1997).
Navifusa bacilla (Deunff) Playford, 1977 (Plate IV,15,18) Remarks: We follow the synonymy of Playford (1977) as well as his expanded diagnosis of this species. All of our specimens showed grana arranged somewhat longitudinally, producing a pseudostriate appearance. Occurrence: Navifusa bacilla as here recognized, is a cosmopolitan species previously reported from the Emsian through Famennian of North America (Deunff, 1955, 1966, 1967; Legault, 1973; Wicander, 1974; Wicander and Wood, 1981, 1997; Wicander and Wright, 1983; Wood and Clendening, 1985; Huysken et al., 1992); Devonian of Brazil (Brito and Santos, 1965; Brito, 1967); Middle Devonian of Paraguay (Pöthe de Baldis, 1974); Late Givetian–Early Frasnian of Argentina (Barreda,1986; Ottone,1996); Givetian of Libya (Moreau-Benoit,1984); Emsian to Late Devonian of North Africa (Jardiné and Yapaudjian, 1968; Jardiné, 1972; Jardiné et al., 1974; Abdesselam-Rouighi, 1987), and the Mid Devonian, Sichuan Province, China (Gao, 1997). Oppilatala cf. O. sparsa (Plate IV,4) Occurrence: Oppilatala sparsa has previously been reported from the Givetian of Ohio, Kentucky, and Iowa (Wicander and Wood, 1981, 1997; Wood and Clendening, 1985). Polyedryxium embudum Cramer, 1964 (Plate IV,16,17) Occurrence: Polyedryxium embudum has been reported from the Early Emsian–Early Eifelian of Ontario (Deunff, 1966, 1971; Playford, 1977); Eifelian–Givetian of Ohio (as Polyedryxium cf. P. embudum in Wicander and Wood, 1981; as Polyedryxium embudum? in Wicander and Wright, 1983); Late Devonian of Kentucky (as Polyedryxium cf. P. embudum in Huysken et al., 1992); Frasnian of Iowa (as Polyedryxium cf. P. embudum in Wicander and Playford, 1985); Late Givetian–Early Frasnian of Argentina (Barreda, 1986; Ottone, 1996); Pragian (Siegenian)–Emsian of northwest Spain (Cramer, 1964); Late Frasnian–Early Famennian of Belgium (Martin, 1981, 1982); Early Frasnian of Western Australia (Playford, 1981; Playford and Dring, 1981); Late Frasnian–Famennian of China (Lu and Wicander, 1988). Rugaletes sp. A (Plate IV,14) Solisphaeridium sp. A (Plate IV,7) Stellinium micropolygonale (Stockmans and Willière) Playford, 1977 (Plate IV,11) Occurrence: This is a widely distributed species in both the northern and southern hemispheres and ranges from the Pragian (Siegenian) through Famennian (see Ottone, 1996 for pertinent references as well as Playford and McGregor, 1993 and Hashemi and Playford, 1998 for additional citations), with a reported occurrence in the Tournaisian of Wales (McNestry, 1988). Tyligmasoma alargada (Cramer) Playford, 1977 (Plate IV,2) Occurrence: Tyligmasoma alargada has been previously reported from the Early Emsian–Givetian of Ontario (Playford, 1977; as Triangulina cf. T. alargada in Legault, 1973); Lochkovian of western Newfoundland (Burden et al., 2002); Eifelian–Givetian of Ohio, Iowa,
Plate IV. Specimens are located by sample (AET25), slide number (5), and England–Finder coordinates. For example, AET25(6), K27/1 refers to sample AET25, slide 6, England–Finder coordinates K27/1. All specimens are × 750 except 1–6, 8, 10–12, 14, which are × 1000. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16, 17. 18.
Exochoderma cf. E. arca Wicander and Wood, 1981. AET24(6), O40/3. Tyligmasoma alargada (Cramer) Playford, 1977. AET24(1), V31/0–4. Diexallophasis remota (Deunff) Playford, 1977, AET27(9), K32/3. Oppilatala cf. O. sparsa Wicander and Wood, 1981. AET24(4), P36/0. Multiplicisphaeridium ramusculosum (Deflandre) Lister, 1970. AET24(10), B36/2. Ephelopalla sp. A. AET24(3), U35/0. Solisphaeridium sp. A. AET25(7), B34/3. Gorgonisphaeridium cumulatum Playford, 1977. AET19(8), O32/0. Veryhachium trispinosum (Eisenack) Stockmans and Willière, 1962 “complex”. AET25(10), L44/3. Cymatiosphaera cornifera Deunff, 1955. AET27(8), T38/4. Stellinium micropolygonale (Stockmans and Willière) Playford, 1977. AET24(7), K37/0–3. Hapsidopalla exornata (Deunff) Playford, 1977. AET19(6), U36/0. Divietipellis cf. D. ventricosa Playford, 1977. AET19(6), P30/0–2. Rugaletes sp. A. AET24(6), R41/3. Navifusa bacilla (Deunff) Playford, 1977. AET26(8), O32/1. Polyedryxium embudum Cramer, 1964. AET25(8), M41/0–1. Navifusa bacilla (Deunff) Playford, 1977, AET26(3), Q45/0.
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
149
150
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
and Kentucky (Wicander and Wood, 1981, 1997; Wicander and Wright, 1983; Huysken et al., 1992); Emsian of northwest Spain (Cramer, 1964; Cramer and Díez, 1976); Silurian of France (as cf. T. alargada in Moreau-Benoit, 1974); Late Lochkovian (Gedinnian)– Pragian (Siegenian) of France (Deunff, 1976, 1980); Ludlow–Emsian of North Africa (Magloire, 1967; Jardiné and Yapaudjian, 1968; Jardiné, 1972; Jardiné et al., 1972, 1974); Emsian–Frasnian of Ghana (AnanYorke, 1974); Early Givetian of Libya (Moreau-Benoit, 1984); Late Silurian–Givetian, Brazil (Brito, 1967, 1969); Emsian of Uruguay (Pöthe de Baldis, 1978); and a reported occurrence in the Tournaisian of Wales (McNestry, 1988). Veryhachium trispinosum (Eisenack) Stockmans and Willière, 1962 “complex” (Plate IV,9) Occurrence: This is a cosmopolitan species having a range of Ordovician through Permian (Wicander and Wood, 1981). 5. Composition and age of the palynoflora The Ulusubasite Formation palynoflora constitutes a somewhat diversified, albeit poorly preserved assemblage of miospores, acritarchs, and scolecodonts. The assemblage is dominated by miospores, both in terms of abundance and diversity, followed by acritarchs, and rare scolecodonts (Fig. 4). Considering that the eight palynomorphic-productive samples are all within a 15 m interval (Fig. 3) with no marked compositional change, we regard this interval as representing one miospore assemblage. This
miospore assemblage consists of 34 species attributed to 23 genera. Four species have been previously named, two are provisionally designated, and 28 are placed in open nomenclature. The assemblage is characterized and dominated by spinose forms such as Dibolisporites cf. D. quebecensis and Poshisporites minutus, which are abundant (N20%); species of Punctatisporites are frequent (10–20%); Apiculiretusispora sp., Auroraspora spp., Dibolisporites echinaceus, D. eifeliensis, Grandispora sp. B, Leiotriletes spp., and the patinate miospores, are all common (5–9%). The rest of the miospore species are rare (b5%) (Fig. 4). In general terms, there are many examples of complex camerate sculptured miospores that are placed in open nomenclature within Grandispora (e.g., Plate I, figs. 1, 2, 4–6, 8, 9, 18). Other camerate miospores include those that have laevigate (Auroraspora, Plate I, fig. 7) or finely sculptured (Rhabdosporites, Plate III, figs. 3, 4) exoexines. Also present is a form with a thickened internal ring structure that can be placed within Densosporites (Plate I, figs. 13, 16, 17) as it is defined within the Devonian. Other zonate forms present are generally attributed to Cristatisporites (Plate I, figs. 3, 10, 11). Ancyrospora (Plate II, figs. 11, 12), a genus with bifurcate sculpture, was also identified. The diversity of camerate and zonate miospores recovered indicates that the miospore assemblage is certainly no older than Eifelian (Richardson and McGregor, 1986). Furthermore, Densosporites first appears within the Eifelian. The patinate miospore group is well-known in the Devonian, and their presence in the Ulusubasite Formation at this location helps
Fig. 4. Distribution and abundances of miospores, acritarchs, and scolecodonts within the Ulusubasite Formation. Relative abundances are based on counts of 200 specimens per sample and converted to a percentage of the 200 specimens. A = abundant (N 20%); F = frequent (10–20%); C = common (5–9%); R = rare (b5%).
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
151
Fig. 5. Distribution and abundances of acritarchs within the Ulusubasite Formation. Relative abundances for the acritarchs in this figure are based on the number of acritarch specimens in the 200 total palynomorph specimens counted per sample in Fig. 4. A = abundant (N8 specimens); C = common (4–8 specimens); R = rare (b 4 specimens).
constrain its age. This miospore group had its inception within the Givetian stage in Euramerica (Turnau and Racki, 1999), thus indicating that the Junggar miospore assemblage is at least Givetian in age. Unfortunately, patinate miospores do not image well in thermally mature assemblages, but the examples shown (Plate III, figs. 1, 2, 5, 6) are clearly trilete miospores with a thickened distal surface that has become modified to give an irregular sculpture with individual elements often connected by irregular muri. Lastly, the assemblage contains the camerate monolete miospore Archaeoperisaccus indistinctus. This distinctive miospore is wellknown from a number of localities in China (Lu, 1980, 1988), as well as the Canning and Adavale basins in Australia (McGregor and Playford, 1993; Hashemi and Playford, 2005) where its geologic range in these areas is Eifelian–Frasnian (Fig. 6). Archaeoperisaccus is a widely distributed genus in China (Eifelian–Famennian) and the northern Euramerican region (Frasnian–Famennian) where it is considered a useful biostratigraphic marker for the Frasnian (Gao, 1988a, 1989; Hashemi and Playford, 2005). The acritarch assemblage comprises 19 species assigned to 19 genera. Three species are provisionally identified (Divietipellis cf. D. ventricosa, Exochoderma cf. E. arca, and Oppilatala cf. O. sparsa), five are placed in open nomenclature (Ammonidium sp., Baltisphaeridium sp., Ephelopalla sp. A, Rugaletes sp. A, and Solisphaeridium sp. A), and 10 are assigned to previously established species. Leiosphaeridia is not speciated, but left in open nomenclature as Leiosphaeridia spp. Although not as abundant or diverse as the miospores, acritarchs were recovered from seven of the eight studied samples, with the greatest diversity occurring in sample AET24 (Fig. 5). The most abundant acritarch is Hapsidopalla exornata, followed in decreasing abundance by Solisphaeridium sp. A, Ephelopalla sp. A, and Navifusa bacilla. The remainder of the acritarch assemblage is represented by only one or a few specimens. Fig. 6 shows the known stratigraphic ranges for the miospore and acritarch species recovered from the Ulusubasite Formation. Excluding Leiotriletes and Punctatisporites, the six named or provisionally named (cf.) miospore species all occur in the Middle Devonian. Two species (one of which is provisionally identified herein — Dibolisporites cf. D. quebecensis, and the other, D. eifeliensis, is positively identified), however, are presently not known to extend beyond the Eifelian (Fig. 6). Except for the provisionally identified Oppilatala cf. O. sparsa, none of the recovered acritarch species are restricted to the Middle
Devonian (Fig. 6), although most were common during that time interval (Wicander, 1984; Molyneux et al., 1996; Le Hérissé et al., 2000). Faunal evidence from the Ulusubasite Formation also corroborates a Middle Devonian age assignment. According to Hou et al. (2000) and Liao (personal communication), corals and brachiopods recovered mainly from the lower and middle parts of the formation indicate an Eifelian age. The palynologic evidence, however, is not sufficient to delineate to the stage level of the Middle Devonian. Because the palynologic assemblage comes from the uppermost part of the Ulusubasite Formation, this stratigraphic interval could be considered Late Eifelian– Givetian in age. 6. Comparison with other palynologic assemblages Paleogeographic reconstructions indicate that during the Middle Devonian, the studied area, as part of the eastern Junggar region, was part of the terrane belonging to the Kazakhstan Plate (Scotese and McKerrow, 1990; Li and Powell, 2001; Blakey, 2003) (Fig. 7). The Kazakhstan Plate consisted of arc islands near a subduction zone during the late Early Devonian and early Mid Devonian. Tuffaceous sediments developed in the Ulusubasite Formation support the existence of these arc islands near the subduction zone. Furthermore, the paleolatitude of the Junggar Terrane is consistent with the latitudinal requirements needed by corals found in the Ulusubasite Formation. As previously noted, the palynomorph-bearing strata are all within the upper part of the Ulusubasite Formation and are assigned a Middle Devonian age based on palynologic, megafaunal, and megaplant fossil evidence (Hou et al., 2000). Therefore, comparison of the Ulusubasite palynoflora is herein made to similar age palynomorph assemblages. 6.1. Miospore assemblage 6.1.1. China There are a few published reports on Devonian miospores from China before 1980 (Fig. 7). Nevertheless, based on the published and unpublished data available at that time, Gao (1981) proposed a preliminary Devonian miospore zonation scheme for China. His scheme included eight zones, two of which (Zones V and VI) are equivalent to the Eifelian and Givetian stages respectively. Zone V is based on material from Yunnan and Guizhou provinces, Southwest
152
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
Fig. 6. Stratigraphic ranges of previously described miospore and acritarch species occurring in the Ulusubasite Formation (for source of data, see Identified palynomorphs section). Question marks indicate uncertainty of range limits; dashes indicate probable occurrence; arrows indicate ranges older or younger than Devonian.
China (Gao and Hou, 1975). However, only one species from this zone is present in the Ulusubasite Formation miospore assemblage, and that is Poshisporites minutus. Ancyrospora furcula and A. langii were reported from Zone VI of Gao (1981). Although we do not speciate the specimens of Ancyrospora recovered, it should be noted that some of
our specimens do resemble A. furcula and A. langii, but are not well enough preserved to make a positive attribution. Subsequently, Lu (1980, 1988) and Gao and Ye (1987) have described Middle Devonian palynofloras from South China and the West Qinling Mountains, respectively.
Fig. 7. Paleogeographic map for the Middle Devonian (based mainly on Blakey, 2003; the positions of the Mongol–Kazakhstan arc and Tarim block are modified from Scotese and McKerrow, 1990, and Li and Powell, 2001). NAM = North America; SIB = Siberia; BAL = Baltica; GON = Gondwana; T = Tarim. Numbers refer to selected published records of Devonian miospores and acritarchs discussed in the text. (1) Scotland (Richardson, 1960, 1962, 1965; Marshall and Allen, 1982); (2) Ardene–Rhenish regions (Streel, 1964, 1967; Stockmans and Willière, 1969; Riegel, 1973, 1974, 1982; Martin, 1981, 1982; Streel et al., 1987; Loboziak and Streel, 1989); (3) Midcontinent, United States (Wicander and Wood, 1981, 1997; Wicander and Wright, 1983; Wood and Clendening, 1985; Huysken et al., 1992); (4) Southeast Canada (Deunff, 1955, 1966, 1967, 1971, 1976, 1980; Legault, 1973; McGregor, 1973, 1977; McGregor and Camfield, 1976; Playford, 1977); (5) northwestern Newfoundland, Canada (Burden et al., 2002); (6) Canadian Arctic (Owens, 1971; McGregor and Camfield, 1982); (7) Svalbard (Allen, 1965, 1967); (8) Eastern Europe (Naumova, 1953; Avkhimovitch et al., 1993; Turnau, 1996; Turnau and Racki, 1999); (9) Junggar, NW China (Lu and Wicander, 1988; present study); (10) Tarim, NW China (Zhu, 1999, 2000); (11) South China (Bharadwaj et al., 1973; Gao and Hou, 1975; Lu, 1980, 1988; Gao, 1997, 1999; Zhu and Lu, 2002); (12) North China (Gao and Ye, 1987; Gao and Liu, 1988); (13) Western Australia (Balme and Hassell, 1962; McGregor and Playford, 1993; Playford, 1981; Playford and Dring, 1981); (14) Tibet (Gao, 1986, 1988b); (15) North Africa (Magloire, 1967; Jardiné and Yapaudjian, 1968; Jardiné, 1972; Jardiné et al., 1972, 1974; Moreau-Benoit, 1984; Abdesselam-Rouighi, 1987; Loboziak and Streel, 1989); (16) West Africa (Anan-Yorke, 1974); (17) South America (Brito and Santos, 1965; Brito, 1967, 1969; Pöthe de Baldis, 1974, 1978; Barreda, 1986; Ottone, 1996).
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
Lu (1980, 1988) systematically studied the Middle Devonian miospores from Zhanyi, Yunnan Province of South China. He described 173 mega- and microspore species assigned to 57 genera from the Middle Devonian (Givetian) Haickou Formation at Shijiapo in Zhanyi of Yunnan, South China (Lu, 1988). However, only Archaeoperisaccus indistinctus is recognized in the present assemblage. From the Lower–Middle Devonian Xujiachong and Haickou formations of the Longhuashan section, Zhanyi, Yunnan, Lu (1980) identified 92 miospore species assigned to 33 genera. Only one species, Archaeoperisaccus indistinctus (reported as Archaeoperisaccus cf. A. scabratus), also occurs in the Ulusubasite Formation miospore assemblage. Of the total miospore flora reported by Lu (1980, 1988), approximately a third of the flora are endemic species (proposed as new species), which may be why there are so few common miospore species between South and Northwest China. In the West Qinling Mountains, Gao and Ye (1987) reported 210 miospore species assigned to 47 genera from the Late Silurian and Devonian and proposed six miospore assemblage zones ranging from Late Silurian to Late Devonian. These zones, the Rhabdosporites langii– Grandispora velata Assemblage Zone (Zone III), the Densosporites devonicus Assemblage Zone (Zone IV), and the Convolutispora crenata Assemblage Zone (Zone V) span the Eifelian to Early Frasnian. However, there are no common species between the Qinling Mountains and the Xinjing region. Thus, there is little similarity between the Middle Devonian Ulusubasite Formation miospore assemblage and those reported from elsewhere in China. This is probably due, in part, to the generally poor preservation of the Ulusubasite miospore assemblage, as well as to the paleogeographic position of the Junggar Basin in relation to South China during the Middle Devonian (Fig. 7). 6.1.2. Extra-China Middle Devonian miospores have been extensively studied in Europe and North America and miospore zonal schemes for the entire Devonian are well-established and extensively used for local, regional, and global correlation (e.g., McGregor and Owens, 1966; McGregor, 1979a,b; Richardson and McGregor, 1986; Streel et al., 1987; McGregor and Playford, 1993; Avkhimovitch et al., 1993). Compared with known Devonian miospore assemblages from Euramerica, the Ulusubasite miospore assemblage contains many genera in common, but only a few co-occurring species. The Middle Old Red Sandstone miospore assemblages from Scotland (Eifelian–Givetian), described by Richardson (1960, 1962, 1965), have only two miospore species or comparable miospore species in common with the Ulusubasite Formation of North Xinjiang (Dibolisporites echinaceus and Rhabdosporites cf. R. parvulus). Including the two cooccurring miospores, the miospores from the Ulusubasite assemblage are generally smaller in size than those from Scotland. Allen (1965,1967) reported Early and Middle Devonian miospores from Svalbard (Vestspitsbergen), Norway, none of which are recognized at the species level in the present Ulusubasite Formation assemblage. However, the Givetian material from Svalbard was, in addition to Geminospora spp., characterized by a diversity of variable patinate spores as recognized in this new material from China. The Middle Devonian palynofloras from Canada, the Eifel region of Germany, and Poland, are also lacking in many co-occurring species with the herein described North Xinjiang miospore assemblage. Of the 38 Late Emsian and Early Eifelian miospore species described by Riegel (1973) from the Eifel region of Germany, only Dibolisporites echinaceus is present in the Ulusubasite Formation miospore assemblage. Interestingly, D. echinaceus is also the only miospore in common between the Ulusubasite Formation miospore assemblage and the Middle Devonian (Givetian) miospore assemblages described by Turnau (1996) and Turnau and Racki (1999) from Poland. The Middle Devonian palynoflora of Canada has been wellinvestigated by McGregor (1973, 1977), McGregor and Camfield
153
(1976, 1982), and McGregor and Playford (1993). Only four miospore species or comparable miospore species (excluding species of Leiotriletes and Punctatisporites) from the Middle Devonian assemblages in Canada also occur in the present assemblage. These include Dibolisporites echinaceus, D. eifeliensis, D. cf. D. quebecensis, and Rhabdosporites cf. R. parvulus. Two other species left in open nomenclature are noted to be similar to species reported from Canada, but because of their preservation, they could not be definitely or provisionally assigned to those species. Those similar taxa are Ancyrospora langii and Calamospora atava. In summary, whereas there are numerous genera in common, at the species level, there are only several co-occurring species between the Ulusubasite Formation miospore assemblage and miospore assemblages of comparable age reported from Canada (McGregor, 1973; McGregor and Camfield, 1976, 1982), the Eifel region of Germany (Riegel, 1973), or elsewhere. This is probably due, to a large extent, to the poor preservation of the Ulusubasite Formation miospore assemblage, but may also be due to the isolated nature of the Junggar Basin region in relation to the Euramerica continent during the Middle Devonian (Fig. 7). 6.2. Acritarch assemblage The acritarch assemblage from the Ulusubasite Formation, although not as diverse, is better preserved than the miospore assemblage, and consists mainly of cosmopolitan taxa. 6.2.1. China To date, 12 papers have been published on Chinese Devonian acritarchs (Li et al., 2002), but only two are concerned with the Middle Devonian of China (Gao and Liu, 1988; Gao, 1999). In Gao and Liu's (1988) one page paper, 10 genera are listed, but only one species is named and that is the long-ranging Veryhachium trispinosum. Gao (1999) described two acritarch assemblages from Shidian County, West Yunnan Province, and interpreted the paleoenvironment of the Emsian–Eifelian Xibiantang Formation and the Givetian Malutang Formation based on these assemblages. Only Navifusa bacilla occurs in both the Ulusubasite Formation acritarch assemblage and the acritarch assemblages from the Xibiantang and Malutang formations, whereas Cymatiosphaera cornifera occurs in our assemblage and the Xibiantang Formation, and Hapsidopalla exornata is common to both our assemblage and that of the Malutang Formation. In regard to the other 10 Chinese Devonian acritarch papers, only the assemblages reported by Gao (1986, 1988b, 1997) and Lu and Wicander (1988), have species in common with those reported here from the Ulusubasite Formation. Gao (1986, 1988b) recognized 48 acritarch species from Upper Devonian and Lower Carboniferous rocks in Nyalam, Xizang (Tibet), but only the cosmopolitan Navifusa bacilla occurs in both our assemblage and the Upper Devonian Zhangdon Formation. Lu and Wicander (1988) described an acritarch and miospore assemblage from the Upper Frasnian–Famennian Hongguleleng Formation, Hefeng District, Xinjiang, in which 34 acritarch species were identified, four of which occurred in both the Hongguleleng and Ulusubasite formations. Two of the species are restricted to the Devonian (Polyedryxium embudum and Stellinium micropolygonale), whereas the other two have long stratigraphic ranges (Multiplicisphaeridium ramusculosum and Veryhachium trispinosum). All four, however, are cosmopolitan in distribution. Gao (1997) reported on a well-preserved acritarch assemblage comprised of 33 species and varieties from the Lower Devonian Ganxi and Xiejiawan formations from Longmenshan Mountains, Sichuan Province. The assemblage consists mostly of long-ranging and cosmopolitan species, including Cymatiosphaera cornifera, Diexallophasis remota, Multiplicisphaeridium cf. M. ramusculosum, and Navifusa bacilla, which also occur in the Ulusubasite Formation.
154
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
6.2.2. Extra-China Most previous reports of Middle Devonian acritarchs from outside China are primarily from North America, with scattered reports from elsewhere in the world (Fig. 7). Fig. 8 shows those taxa our acritarch assemblage has in common with other published Middle Devonian acritarch assemblages. We have only listed those assemblages where the taxa are well-preserved and illustrated, and there is good stratigraphic control. Accordingly, only 11 Middle Devonian assemblages are listed, although all reported published occurrences of each Ulusubasite species are cited in the Identified palynomorphs section. Discussion of specific acritarch assemblages follows, and for purposes of comparison, we are equating provisionally identified (cf.) taxa the same as the species itself. In North America, there are seven Middle Devonian assemblages that meet the above criteria. Of the eight named acritarch species reported by Legault (1973) from the Hamilton Formation (Givetian) of southwestern Ontario, we have only two species in common. Playford (1977) reported 53 acritarch species from five lithostratigraphic units
in the Moose River Basin, northern Ontario [Upper Pragian (Siegenian) to Givetian], 10 of which also occur in the Ulusubasite Formation. Wicander and Wood (1981) record 81 acritarch species from the Silica Formation (Givetian) of Ohio, of which, 11 occur in the Ulusubasite Formation. Wicander and Wright (1983) identified 43 acritarch species from the Columbus and Delaware Limestones (Eifelian–Givetian) of Ohio. Of this total, nine are common to our assemblage from China. Although Wood and Clendening (1985) state the acritarch assemblage from the Boyle Dolomite (Givetian) of Kentucky is poorly preserved, the figured specimens are better preserved than those from the Ulusubasite Formation, and of the 23 recorded species, there are six co-occurring species. Of the 14 acritarch species reported by Huysken et al. (1992) from the Givetian-age Sellersburg Formation and Blocher Member of the New Albany Shale, Kentucky, seven species are common to the Ulusubasite Formation. Lastly, Wicander and Wood (1997) report 40 acritarch species recovered from the Rapid Member of the Cedar Valley Formation (Givetian) of Iowa, of which only six are common to the Ulusubasite Formation.
Fig. 8. Comparison of the Ulusubasite Formation acritarch assemblage with other selected published Middle Devonian acritarch assemblages.
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
In South America, Barreda (1986) described an acritarch assemblage consisting of 45 species from a well in Salta Province, Argentina, corresponding to the Upper Givetian–Lower Frasnian Torono Formation. Of the 45 species reported, there are six species in common with the Ulusubasite Formation, as well as what we describe as Ephelopalla sp. A, which closely resembles Barreda's (1986) Solisphaeridium sp. 1. Ottone (1996) recorded 51 acritarch species from the Upper Givetian–Lower Frasnian Los Monos Formation in northwestern Argentina, of which eight are co-occurring in the Ulusubasite Formation. Core samples ranging in age from Givetian to Famennian from three wells in western Libya (Moreau-Benoit, 1984) yielded an assemblage of 43 acritarch species, five of which are common to the Ulusubasite Formation. The range charts and summaries for the Devonian of the Algerian Sahara (Jardiné et al., 1974) indicate four species in common with the Ulusubasite Formation. In summary, although the degree of similarity between the Ulusubasite Formation acritarch assemblage and those of comparable age reported elsewhere is not high (due, in part, to preservation, and also paleoenvironmental differences between the various assemblages), there are, nonetheless, many species in common. In fact, most of the identifiable acritarch species from the Ulusubasite Formation are both cosmopolitan and stratigraphically long-ranging. Molyneux et al. (1996) pointed out that because of the few number of Middle Devonian acritarch and prasinophyte floras reported at that time, it was difficult to assess the degree of provincialism and cosmopolitanism of many taxa. Le Hérissé et al. (2000, p. 200) reiterated this observation and also noted that most studies were concentrated on Givetian assemblages, with little being published on the Eifelian. They also indicated that it appeared that a relatively narrow and temperate marine realm had developed between the Northern Hemisphere and Gondwana during the Givetian–Frasnian, based on the recovered acritarch assemblage of Ottone (1996), Le Hérissé and Deunff (1988), and Wicander (1984). We can extend this marine linkage between North America, Baltica, the Junggar Basin of Kazakhstan, and Gondwana, based on the occurrence of six Ulsubasite Formation acritarchs. These include Cymatiosphaera cornifera, which has previously been reported from the Middle Devonian of North America and Sichuan Province, China, with a possible occurrence in the Frasnian of Belgium; Divietipellis ventricosa, Gorgonisphaeridium cumulatum, and Oppilatala sparsa, all three of which have previously been reported only from the Pragian (Siegenian)–Givetian of North America; and Exochoderma arca and Hapsidopalla exornata, both of which have been reported from North America (Eifelian–Givetian, ?Frasnian) and Argentina (Late Givetian– Early Frasnian), showing that a marine link between these two regions and China existed during the Middle Devonian (Fig. 8). This compliments the findings of Liao (1990) that the Junggar Basin Middle Devonian coral fauna is also similar to those of Europe and North American, and reinforces the presence of a marine linkage between these regions. 7. Paleoenvironmental interpretation Abundant marine fossils such as corals, brachiopods, and crinoids occur mainly in the middle and lower parts of the Ulusubasite Formation, whereas the productive palynomorph-bearing strata are in the upper part of the formation (Fig. 3). As previously stated, this region was affected by volcanic activity during the Middle Devonian as evidenced by the well-developed tuffaceous sediments interbedded throughout the Ulusubasite Formation. Such periodic volcanic activity may, in part, explain the dark color and poor preservation of the entire palynoflora due to the baking effect of volcanic sedimentation. The co-occurrence of marine acritarchs and terrestrially-derived miospores suggests that the upper part of the Ulusubasite Formation was deposited in a near-shore marine environment. The inverse relationship between abundance of land-derived spores and pollen
155
Fig. 9. Miospore to acritarch abundance ratios expressed as a percentage for the Ulusubasite Formation, eastern Junggar Basin, Xinjiang, China.
and marine microphytoplankton is well-established and accepted as a tool for determining shoreline proximity (Upshaw, 1964; Wicander and Playford, 1985; and others). Fig. 9 shows the miospore to acritarch abundance ratios in the samples studied. The miospore to acritarch ratio strongly implies near-shore conditions during deposition of the upper part of the Ulusubasite Formation (Fig. 9). This interpretation also fits the paleontologic and lithologic evidence from the Ulusubasite Formation in which an abundant and well-preserved marine fauna was recovered from the middle and lower parts of the formation, and broken crinoids and fragments of megaplant fossils occurring in the upper part of the formation. In summary, the Ulusubasite Formation in the eastern Junggar Basin represents a near-shore marine depositional sequence in the lower northern latitudes that was punctuated by nearby volcanic activity during the Middle Devonian. 8. Conclusions Abundant, although not particularly well-preserved miospores, acritarchs, and scolecodonts are first reported herein from the Middle Devonian Ulusubasite Formation of the Zhifang Section, eastern Junggar Basin, Xinjiang, Northwest China, a region in which little is known palynologically. Thirty-four miospore species assigned to 23 genera and 19 acritarch species assigned to 19 genera, as well as scolecodonts, are identified. The biostratigraphic ranges of the miospores and acritarchs indicate a Mid Devonian age. Megafossil faunal and floral evidence not only supports this age assignment, but also refines it (at least the lower and middle part of the Ulusubasite Formation) to the Eifelian. There is little similarity at the species level between the Middle Devonian Ulusubasite Formation miospore assemblage and those reported from elsewhere in China, as well as from Euramerica and Gondwana. This is probably due, to a large extent, to the poor preservation of the Ulusubasite miospore assemblage, but may also reflect the isolated nature of the Junggar Basin region in relation to South China and the Euramerica and Gondwana continents during the Middle Devonian. The acritarch assemblage consists mostly of cosmopolitan species. Furthermore, six acritarch species recovered from the Ulusubasite Formation provide additional evidence for a marine linkage between North America, Baltica, the Junggar Basin of Kazakhstan, and Gondwana. Paleontological and lithological evidence indicates that the Ulusubasite Formation was deposited in a near-shore marine environment. Acknowledgments Huaicheng Zhu thanks the Chinese Academy of Sciences for a grant and sabbatical leave to undertake this research and to the Central Michigan University Department of Geology for hospitality and use of their facilities. He also thanks Weihua Liao, Fengsheng Xia, and Jin
156
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157
Zhang for their assistance during the field investigation in 1988. We thank Cuiling He for her laboratory preparation for the studied samples, and Qifei Wang and Jun Li for procuring copies of needed Chinese references. We also thank two anonymous reviewers for their critical reading of our manuscript and their various suggestions that have improved this paper. This study was financially supported by the Chinese Academy of Sciences (Nos. KZCX2-SW-130 and KZCX2-YW105) and the National Natural Science Foundation of China (Nos. 40523004 and 40072005). References Abdesselam-Rouighi, F., 1987. Premiers resultants biostratigraphiques (miospores, acritarches et chitinozoaires) concernant le Dévonien moyen et supérieur du mole d'Ahara (basin d'Illizi, Algérie). Revue de Micropaléontologie 29 (2), 87–92. Allen, K.C., 1965. Lower and Middle Devonian spores of north and central Vestspitsbergen. Palaeontology 8 (4), 687–748. Allen, K.C., 1967. Spore assemblages and their stratigraphical application in the Lower and Middle Devonian of north and central Vestspitsbergen. Palaeontology 10 (2), 280–297. Anan-Yorke, R., 1974. Devonian Chitinozoa and Acritarcha from exploratory oil wells on the shelf and coastal region of Ghana, West Africa. Ghana Geological Survey, Bulletin 37, 1–216. Avkhimovitch, V.I., Tchibrikova, T.G., Obukhovskaya, T.G., Nazarenko, A.M., Umnova, V.T., Raskatova, L.G., Mantsurova, V.N., Loboziak, S., Streel, M., 1993. Middle and Upper Devonian miospore zonation of eastern Europe. Bulletin de Centre Recherches Exploration-Production, Elf Aquitaine 17 (1), 79–147. Balme, B.E., Hassell, C.W., 1962. Upper Devonian spores from the Canning Basin, Western Australia. Micropaleontology 8 (1), 1–28. Barreda, V.D., 1986. Acritarcos Givetiano-Frasnianos de la Cuenca del Noroeste, Provincia de Salta, Argentina. Revista Española de Micropaleontología 18 (2), 229–245. Bharadwaj, D.C., Tiwari, R.S., Venkatachala, B.S., 1973. A Devonian microflora from P'oshi district (Yunnan), China. Palaeobotanist 20 (2), 152–169. Blakey, R., 2003. Paleogeography Through Geologic Time. 1st Order Global Tectonic Features. Late Devonian. http://jan.ucc.nau.edu/~rcb7/1storder.html [viewed August 14, 2003]. Brito, I.M., 1967. Silurian and Devonian Acritarcha from Maranhão Basin, Brazil. Micropaleontology 13 (4), 473–482. Brito, I.M., 1969. Un nouveau sous-groupe d'Acritarche. Universidade de Rio de Janeiro, Instituto de Geociências, Boletim 4, 11–13. Brito, I.M., Santos, A.S., 1965. Contribução ao conhecimento dos microfosseis Silurianos e Devonianos da Bacia do Maranhão. Parte I. Os Netromorphitae (Leiofusidae). Divisão de Geologia y Mineria do Brasil, Notas Preliminares e Estudos 129, 1–23. Burden, E.T., Quinn, L., Nowlan, G.S., Bailey-Nill, L.A., 2002. Palynology and micropaleontology of the Clam Bank Formation (Lower Devonian) of western Newfoundland, Canada. Palynology 26, 185–215. Cramer, F.H., 1964. Microplankton from three Palaeozoic formations in the Province of Leon, NW-Spain. Leidse Geologische Mededelingen 30, 253–361. Cramer, F.H., Díez, M. del C.R., 1976. Acritarchs from the La Vid Shales (Emsian to lower Couvinian) at Colle, Leon, Spain. Palaeontographica Abteilung B 158, 72–103. Deunff, J., 1955. Un microplancton fossile Dévonien à hystrichosphères du continent Nord-Américain. Bulletin de microscopie appliqué, Série 2, 5 (11–12), 138–149. Deunff, J.,1966. Recherches sur les microplanctons du Dévonien (Acritarches et Dinophyceae). Rennes, the author, pp. 1–168. Deunff, J., 1967. Présence d'acritarches dans une série Dévonienne du lac Huron (Canada). Compte Rendu Sommaire des Séances de la Société Géologique de France 6, 258–260. Deunff, J., 1971. Le genre Polyedryxium Deunff-révision et observations. In: Jardiné, S. (Ed.), Microfossiles organiques du Paleozoique, 3. Acritarches. Commission International de Microflore du Paléozoique. Editions du Centre National de la Recherché Scientifique, Paris, pp. 7–49. Deunff, J., 1976. Les Acritarches. In: Lardeaux, H. (Ed.), Les Schistes et calcaires éodévoniens de Saint-Céneré (Massif Armoricain, France): sédimentologie, paleontology, stratigraphie. Mémoire de la Société géologique et mineralogique de Bretagne, vol. 19, pp. 59–77. Deunff, J., 1980. Le paléoplancton des Grès de Landévennec (Gédinnien de la Rade de Brest-Finstère) etude biostratigraphique. Geobios 13 (4), 483–539. Gao, L., 1981. Devonian spore assemblages of China. Review of Palaeobotany and Palynology 34 (1), 11–23. Gao, L., 1986. Late Devonian and Early Carboniferous acritarchs from Nyalam County, Xizang (Tibet), China. Review of Palaeobotany and Palynology 47 (1–2), 17–30. Gao, L., 1988a. Archaeoperisaccus in China: its stratigraphic and palaeogeographic significance. Pollen et Spores 30 (3–4), 461–470. Gao, L., 1988b. Late Devonian and Early Carboniferous acritarchs from Nyalam County, Xizang (Tibet). Professional Papers of Xizang (Tibet), Paleontology. Chinese Academy of Geological Sciences Geological Publishing House, Beijing, pp. 384–423. In Chinese with English abstract. Gao, L., 1989. Geological range and geographical distribution of Archaeoperisaccus. Acta Micropalaeontologica Sinica 6 (2), 197–206. In Chinese with English summary. Gao, L., 1997. Acritarchs from Lower Devonian Ganxi and Xiejiawan Formations in Longmenshan Mountain, Sichuan. Acta Geologia Gansu 6, 16–24. Gao, L., 1999. Acritarchs of Devonian from Shidian, West Yunnan and their palaeoenvironments. Journal of Palaeogeography 1 (4), 35–44. In Chinese with English abstract.
Gao, L., Hou, J., 1975. Characteristics of the Early and Middle Devonian spore assemblages from Dunshan and Dunyun counties, Guizhou Province, and their stratigraphic significance. Professional Papers of Stratigraphy and Palaeontology 1, 170–232. In Chinese. Gao, L., Ye, X., 1987. Late Silurian and Devonian spores from west Qinling Mountains, China. Late Silurian–Devonian Strata and Fossils from Luqu-Tewo Area of West Qinling Mountains, China, vol. 2. Nanjing University Press, Nanjing, pp. 379–450. In Chinese with English abstract. Gao, L., Liu, Z., 1988. Discovery of microfossils from Nanwan Formation of Xinyang Group in Henan and its geological significance. Geological Review 34, 421. In Chinese. Hashemi, H., Playford, G., 1998. Upper Devonian palynomorphs of the Shishtu Formation, Central Iran Basin, east-central Iran. Palaeontographica Abteilung B 246, 115–212. Hashemi, H., Playford, G., 2005. Devonian spore assemblages of the Adavale Basin, Queensland (Australia): descriptive systematics and stratigraphic significance. Revista Española de Micropaleontología 37, 317–417. Hou, H., Cao, X., Wang, S., Xian, S., Wang, J., 2000. Stratigraphic Lexicon of China: Devonian System. Geological Publishing House, Beijing. 119 pp. In Chinese. Huysken, K.T., Wicander, R., Ettensohn, F.R., 1992. Palynology and biostratigraphy of selected Middle and Upper Devonian black-shale sections in Kentucky. Michigan Academician 24, 355–368. Jardiné, S., 1972. Microplancton (Acritarches) et limites stratigraphiques du Silurien terminal au Dévonien supérieur. VII Congrès International de Stratigraphie et de Géologie du Carbonifère, Krefeld (1971). Compte Rendu 1, 313–323. Jardiné, S., Yapaudjian, L., 1968. Lithostratigraphie et palynologie du DévonienGothlandien gréseux du bassin de Polignac (Sahara). Revue de l'Institut Français de Pétrole 23 (4), 439–469. Jardiné, S., Combaz, A., Magloire, L., Peniguel, G., Vachey, G., 1972. Acritarches du Silurien terminal et du Dévonien du Sahara algérien. VII Congrès International de Stratigraphie et de Géologie du Carbonifère, Krefeld (1971). Compte Rendu 1, 295–311. Jardiné, S., Combaz, A., Magloire, L., Peniguel, G., Vachey, G., 1974. Distribution stratigraphique des Acritarches dans le Paléozoique du Sahara algérien. Review of Palaeobotany and Palynology 18 (1–2), 99–129. Legault, J.A., 1973. Chitinozoa and Acritarcha of the Hamilton Formation (Middle Devonian) southwestern Ontario. Geological Survey of Canada Bulletin 221, 1–103. Le Hérissé, A., Deunff, J., 1988. Acritarches et Prasinophycés (Givétien supérieurFrasnien moyen) de Ferques (Boulonnais-France). In: Brice, D. (Ed.), Le Dévonien de Ferques, Bas-Boulonnais (N. France). Biostratigraphy du Paléozoïque, vol. 7, pp. 104–152. Le Hérissé, A., Servais, T., Wicander, R., 2000. Devonian acritarchs and related forms. Courier Forschungsinstitut Senckenberg 220, 195–205. Li, J., Servais, T., Brocke, R., 2002. Chinese Paleozoic acritarch research: review and perspectives. Review of Palaeobotany and Palynology 118 (1–4), 181–193. Li, J., Wicander, R., Yan, K., Zhu, H., 2006. An Upper Ordovician acritarch and prasinophyte assemblage from Dawangou, Xinjiang, northwestern China: biostratigraphic and paleogeographic implications. Review of Palaeobotany and Palynology 139, 97–128. Li, Z.X., Powell, C.M., 2001. An outline of the palaeogeographic evolution of the Australasian region since the beginning of the Neoproterozoic. Earth-Science Reviews 53 (3–4), 237–277. Liao, W., 1990. The biogeographic affinities of east Asian corals. In: McKerrow, W.S., Scotese, C.R. (Eds.), Palaeozoic Palaeogeography and Biogeography. Geological Society Memoir, vol. 12, pp. 175–179. Loboziak, S., Streel, M., 1980. Miospores in Givetian to Lower Frasnian sediments dated by conodonts from the Boulonnais, France. Review of Palaeobotany and Palynology 29 (3–4), 285–299. Loboziak, S., Streel, M., 1989. Middle–Upper Devonian miospores from the Ghadamis basin (Tunisia-Libya): systematics and stratigraphy. Review of Palaeobotany and Palynology 58 (2–4), 173–196. Lu, L., 1980. Devonian miospores from the Longhuashan section in Zhanyi of Yunnan and their stratigraphic significance. Memoirs of Nanjing Institute of Geology and Palaeontology, Academia Sinica 14, 1–62. In Chinese with English abstract. Lu, L., 1988. Middle Devonian microflora from Haickou Formation at Shijiapo in Zhanyi of Yunnan, China. Memoirs of the Nanjing Institute of Geology and Palaeontology, Academia Sinica 24, 109–234. In Chinese with English abstract. Lu, L., Wicander, R., 1988. Upper Devonian acritarchs and spores from the Hongguleleng Formation, Hefeng District in Xinjiang, China. Revista Española de Micropaleontología 20 (1), 109–148. Magloire, L., 1967. Etude stratigraphique, par la palynologie, des depots argilo-gréseux du Silurien et du Dévonien inférieur dans la region du Grand Erg Occidental (Sahara algérien). International Symposium on the Devonian System, Calgary. Alberta 2, 473–491. Marshall, J.E.A., 1995. A simple cost-effective infra-red microscope for palynology. Journal of Micropalaeontology 14, 106. Marshall, J.E.A., Allen, K.C., 1982. Devonian miospore assemblages from Fair Isle, Shetland. Palaeontology 25 (2), 277–312. Martin, F., 1981. Acritarches du Famennien inférieur à Villiers-sur-Lesse (Belgique). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre 52 (2), 1–49. Martin, F., 1982. Acritarches et Chitinozoaires de la partie supérieure du Frasnien dans un affleurement au nord immediate de Frasnes (Belgique). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre 54 (2), 1–17. McGregor, D.C., 1973. Lower and Middle Devonian spores of eastern Gaspé, Canada. I. Systematics. Palaeontographica Abteilung B 142, 1–77. McGregor, D.C., 1977. Lower and Middle Devonian spores of eastern Gaspé, Canada. II. Biostratigraphic significance. Palaeontographica Abteilung B 113, 111–142.
H. Zhu et al. / Review of Palaeobotany and Palynology 152 (2008) 141–157 McGregor, D.C., 1979a. Spores in Devonian stratigraphical correlation in the Devonian System. In: House, M.R., Scrutton, C.T., Bassett, M.G. (Eds.), The Devonian System. Special Papers in Palaeontology, vol. 23, pp. 163–184. McGregor, D.C., 1979b. Devonian miospores of North America. Palynology 3, 31–52. McGregor, D.C., Owens, B., 1966. Illustrations of Canadian fossils: Devonian spores of eastern and northern Canada. Geological Survey of Canada Paper 66–30, 1–66. McGregor, D.C., Camfield, M., 1976. Upper Silurian (?) to Middle Devonian spores of the Moose River Basin, Ontario. Geological Survey of Canada Bulletin 263, 1–63. McGregor, D.C., Camfield, M., 1982. Middle Devonian miospores from the Cape De Bray, Weatherall, and Hecla Bay Formations of northeastern Melville Island, Canadian Arctic. Geological Survey of Canada Bulletin 348, 1–105. McGregor, D.C., Playford, G., 1993. Canadian and Australian Devonian spores: zonation and correlation. Geological Survey of Canada Bulletin 438, 1–84 Imprinted 1992. McNestry, A., 1988. The palynostratigraphy of two uppermost Devonian–Lower Carboniferous borehole sections in South Wales. Review of Palaeobotany and Palynology 56 (1–2), 69–87. Molyneux, S.G., Le Hérissé, A., Wicander, R., 1996. Chapter 16. Paleozoic phytoplankton. In: Jansonius, J., McGregor, D.C. (Eds.), Palynology: Principles and Applications. American Association of Stratigraphic Palynologists Foundations, vol. 2, pp. 493–529. Moreau-Benoit, A., 1974. Recherches de palynologie et de planctologie sur le Dévonien et quelques formations siluriennes dans le Sud-Est du Massif Armoricain. Mémoires de la Société géologique et minéralogique de Bretagne 18, 1–251. Moreau-Benoit, A., 1984. Acritarches et Chitinozoaires du Dévonien moyen et supérieur de Libye Occidentale. Review of Palaeobotany and Palynology 43 (1–3), 187–216. Naumova, S.N., 1953. Spore-pollen assemblages of the Upper Devonian of the Russian Platform and their stratigraphic significance. Transactions of Institute of Geological Sciences, Academy of Sciences of the USSR 143 (60), 1–204. In Russian. Ottone, E.G., 1996. Devonian palynomorphs from the Los Monos Formation, Tarija Basin, Argentina. Palynology 20, 105–155. Owens, B., 1971. Miospores from the Middle and early Upper Devonian rocks of the western Queen Elizabeth Islands, Arctic Archipelago. Geological Survey of Canada Paper 70–38, 1–157. Playford, G., 1977. Lower to Middle Devonian acritarchs of the Moose River Basin, Ontario. Geological Survey of Canada Bulletin 279, 1–87. Playford, G., 1981. Late Devonian acritarchs from the Gneudna Formation in the western Carnarvon Basin, Western Australia. Geobios 14 (2), 145–171. Playford, G., Dring, R.S., 1981. Late Devonian acritarchs from the Carnarvon Basin, Western Australia. Special Papers in Palaeontology 27, 1–78. Playford, G., McGregor, D.C., 1993. Miospores and organic-walled microphytoplankton of Devonian–Carboniferous boundary beds (Bakken Formation), southern Saskatchewan: A systematic and stratigraphic appraisal. Geological Survey of Canada Bulletin 445, 1–107. Pöthe de Baldis, E.D., 1974. El microplancton de Devónico Medio de Paraguay. Revista Española de Micropaleontología 6 (3), 367–369. Pöthe de Baldis, E.D., 1978. Paleomicroplancton adicional del Devonico inferior de Uruguay. Revista Española de Micropaleontología 9 (2), 235–250. Richardson, J.B., 1960. Spores from the Middle Old Red Sandstone of Cromarty, Scotland. Palaeontology 3 (1), 45–63. Richardson, J.B., 1962. Spores with bifurcate processes from the Middle Old Red Sandstone of Scotland. Palaeontology 5 (2), 171–194. Richardson, J.B., 1965. Middle Old Red Sandstone spore assemblages from the Orcadian Basin, north-east Scotland. Palaeontology 7 (4), 559–605. Richardson, J.B., McGregor, D.C., 1986. Silurian and Devonian spore zones of the Old Red Sandstone Continent and adjacent regions. Geological Survey of Canada Bulletin 364, 1–79. Riegel, W., 1973. Sporenformen aus den Heisdorf-, Lauch- und Nohn-Schichten (Emsium und Eifelium) der Eifel, Rheinland. Palaeontographica Abteilung B 142, 78–104. Riegel, W., 1974. Phytoplankton from the Upper Emsian and Eifelian of the Rhineland, Germany — a preliminary report. Review of Palaeobotany and Palynology 18 (1–2), 29–39.
157
Riegel, W., 1982. Palynological aspects of the Lower/Middle Devonian transition in the Eifel region. Courier Forschungsinstitut Senckenberg 55, 279–292. Scotese, C.R., McKerrow, W.S., 1990. Revised world maps and introduction. In: McKerrow, W.S., Scotese, C.R. (Eds.), Palaeozoic Palaeogeography and Biogeography. Geological Society Memoir, vol. 12, pp. 175–179. Stockmans, F., Willière, Y., 1969. Acritarches du Famennien inférieur. Académie royale des sciences, des letters et des beaux artes de Belgique, Classe des sciences, Mémoires 38 (6), 1–63. Streel, M., 1964. Une association de spores du Givétien inférieur de la Vesdre, à Goé (Belgique). Annales de la Société Géologique de Belgique 87 (7), 1–30. Streel, M., 1967. Associations de spores du Dévonien inférieur belge et leur signification stratigraphique. Annales de la Société Géologique de Belgique 90 (1–3), 11–53. Streel, M., Higgs, K., Loboziak, S., Riegel, W., Steemans, P., 1987. Spore stratigraphy and correlation with faunas and floras in the type marine Devonian of the Ardene– Rhenish regions. Review of Palaeobotany and Palynology 50 (3), 211–229. Turnau, E., 1996. Miospore stratigraphy of Middle Devonian deposits from Western Pomerania. Review of Palaeobotany and Palynology 93, 107–125. Turnau, E., Racki, G., 1999. Givetian palynostratigraphy and palynofacies: new data from the Bodzentyn Syncline (Holy Cross Mountains, central Poland). Review of Palaeobotany and Palynology 106, 237–271. Upshaw, C.F., 1964. Palynological zonation of the Upper Cretaceous Frontier Formation near Dubois, Wyoming. In: Cross, A.T. (Ed.), Palynology in Oil Exploration. Society of Economic and Paleontology and Mineralogy Special Publication, vol. 11, pp. 153–168. Wicander, E.R., 1974. Upper Devonian–Lower Mississippian acritarchs and prasinophycean algae from Ohio, U.S.A. Palaeontographica Abteilung B 148, 9–43. Wicander, R., 1984. Middle Devonian acritarch biostratigraphy of North America. Journal of Micropaleontology 3 (2), 19–24. Wicander, R., Wood, G.D., 1981. Systematics and biostratigraphy of the organic-walled microphytoplankton from the Middle Devonian (Givetian) Silica Formation, Ohio, U.S.A. American Association of Stratigraphic Palynologists Contribution Series 8, 1–137. Wicander, R., Wright, R.P., 1983. Organic-walled microphytoplankton abundance and stratigraphic distribution from the Middle Devonian Columbus and Delaware Limestones of the Hamilton Quarry, Marion County, Ohio. The Ohio Journal of Science 83 (1), 2–13. Wicander, R., Playford, G., 1985. Acritarchs and spores from the Upper Devonian Lime Creek Formation, Iowa, U.S.A. Micropaleontology 31 (2), 97–138. Wicander, R., Wood, G.D., 1997. The use of microphytoplankton and chitinozoans for interpreting transgressive/regressive cycles in the Rapid Member of the Cedar Valley Formation (Middle Devonian), Iowa. Review of Palaeobotany and Palynology 98 (1–2), 125–152. Wood, G.D., 1986. Some Middle Devonian (Givetian) chitinozoans and acritarchs from New York State: Preliminary results. In: Miller, M.A. (Ed.), A Field Excursion to Trenton Group (Middle and Upper Ordovician) and Hamilton Group (Middle Devonian) Localities in New York, and a Survey of Their Chitinozoans. American Association of Stratigraphic Palynologists Field Trip Guidebook, pp. 101–114. Wood, G.D., Clendening, J.A., 1985. Organic-walled microphytoplankton and chitinozoans from the Middle Devonian (Givetian) Boyle Dolomite of Kentucky, U.S.A. Palynology 9, 133–145. Zhu, H., 1999. Late Devonian spore assemblages from the Qizilafu Formation of Sache County, southern Xinjiang and their palynofacies analysis. Acta Palaeontologica Sinica 38 (1), 56–85. In Chinese with English summary. Zhu, H., 2000. Upper Devonian spores from the Tarim Basin, NW China. Acta Palaeontologica Sinica 39 (2), 159–176. In Chinese with English summary. Zhu, H., Lu, L., 2002. A Middle Devonian spore assemblage from Zhenyi, Yunnan, South China. Abstracts from Palaeozoic Palynology in the Third Millennium; New Directions in Acritarch, Chitinozoan and Miospore Research. CIMP 2002. Lille, Sept. 2002, 75. Abstract.