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A latest Permian radiolarian fauna from Hushan, South China, and its geological implications

ARTICLE in ALCHERINGA AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY · DECEMBER 2011 Impact Factor: 1.12 · DOI: 10.1080/03115518.2010.536649

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A latest Permian radiolarian fauna from Hushan, South China, and its geological implications 50 5 WEIHONG HE, YANG ZHANG, QIANG ZHANG, KEXIN ZHANG, AIHUA YUAN AND QINGLAI FENG 55 WEIHONG HE,YANG ZHANG,QIANG ZHANG,KEXIN ZHANG,AIHUA YUAN &QINGLAI FENG, XXXX, 2010. A latest Permian radiolarian fauna from Hushan, South China, and its geological implications. Alcheringa 35, 000–000. ISSN 10 0311-5518. A latest Permian (late Changhsingian) radiolarian fauna is recorded from the upper Talung Formation, Hushan, Nanjing, Jiangsu Province, South China. This fauna includes 24 species belonging to 16 genera; new species are Albaillella hushanensis, Copicyntroides stellatus and Trilonche crassus. The presence of the radiolarian fauna and its taxonomic composition reveal that the Eastern Qinling-Dabie deep sea, which was located along the northern margin 60 of the northeastern Yangtze Basin, persisted at least until the end of the Palaeozoic and that the collision between the 15 North China and South China plates had not occurred by the end of the Permian. Weihong He [[email protected]] and Kexin Zhang, Key Laboratory of Biogeology and Environmental Geology of Ministry of Education, China University of Geosciences, Wuhan 430074, PR China; Yang Zhang, Aihua Yuan and Qinglai Feng, Faculty of Earth Sciences, China University of Geosciences, Wuhan 430074, PR China. Qiang Zhang, Key Laboratory of Marginal Sea Geology, South China Sea Institute of Oceanology, Chinese Academy of Sciences, 65 Guangzhou 510301, PR China. Received 15.3.2010, revised 25.6.2010, accepted 25.10.2010. 20 Key words: Radiolaria, Eastern Qinling-Dabie deep sea, late Changhsingian, Hushan, palaeogeography.

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25 CHANGHSINGIAN radiolarian faunas time (Empson-Morin 1984, Catalano et al. from the Palaeotethys have been reported 1991, Kozur 1993, Kuwahara 1999a, Feng widely (Ishiga et al. 1982, Kuwahara & Yao & Ye 2000, He 2006, He et al. 2008). Hence, 1998, Kuwahara 1999a, Yao & Kuwahara radiolarians are useful for palaeobathy- 75 1999, 2000, Sashida et al. 2000a, b, Shang metric and palaeogeographic interpretations 30 et al. 2001, Wang & Shang 2001, Feng et al. of late Palaeozoic terranes. 2007a, Go¨ncu¨oglu et al. 2004). Numerous The timing of convergence and collision works directly or indirectly related to between the North and South China plates radiolarian ecology have also been pub- remains controversial. Debate especially 80 lished (modern radiolarian ecology—see focuses on whether a deep sea (Eastern 35 Gowing & Garrison 1992, Abelmann et al. Qinling-Dabie sea) existed along the north- 1999, Nimmergut & Abelmann 2002, ern margin of the northeastern Yangze Schmidt et al. 2006; fossil radiolarian Basin in the late Palaeozoic. Some research ecology—see Holdsworth 1977, Empson- has suggested that the Eastern Qinling- 85 Morin 1984, Catalano et al. 1991, Afana- Dabie deep sea north to the northeastern 40 sieva & Zamilatskay 1993, Kozur 1993, Yangtze Basin, disappeared and was re- Kuwahara 1999a, Feng & Ye 2000, He placed by an epicontinental sea or even by 2006, He et al. 2008). Most indicate that land at the end of the Silurian (Xu et al. radiolarian associations varied with bathy- 1988, Ren et al. 1991). Other research 90 metry (hence sea-level fluctuations) through suggests that the Eastern Qinling-Dabie 45 deep sea persisted until the Triassic (Zhang ISSN 0311-5518 (print)/ISSN 1752-0754 (online) Ó 2010 Association of Australasian Palaeontologists et al. 1988, Feng et al. 1994, Du et al. 1997). DOI: 10.1080/03115518.2010.536649 The age and location of the Hushan 2 WEIHONG HE et al. ALCHERINGA

95 radiolarian fauna are, therefore, important trough hosting siliceous sediments and a for understanding when this deep sea shallow-water carbonate platform (Fig. 1). disappeared. The Hushan section exposes strata of the Here we re-investigate and describe the siliceous succession of the northern margin 145 radiolarian fauna from the Hushan section of the northeastern Yangtze Basin (Fig. 1). 100 of South China. We incorporate records The Hushan section of this study is from previous studies (Zhang et al. 1992), located ca 20 km east of Nanjing City, and compare the taxonomic representation Jiangsu Province. It can be reached by bus and diversity of this fauna with other in a half hour from Nanjing City (Fig. 2). 150 radiolarian assemblages to clarify whether The Permian–Triassic interval in the Hush- 105 the Eastern Qinling-Dabie deep sea existed an section includes the upper Talung For- at the end of the Permian. mation and the basal part of the Lower Chinglung Formation (Fig. 3). The upper Talung Formation of the 155 Geological setting Hushan section is composed of dark grey 110 Palaeogeographically, the Yangtze Basin thin-bedded cherts and siliceous mudstones was primarily subdivided into a deep-water (with well-developed horizontal bedding)

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Fig. 1. Changhsingian palaeogeography of South China (revised after Feng & Gu 2002). 1, Shallow-water carbonate 185 platform; 2, Terrestrial facies; 3, Deltaic facies; 4, Deep-water siliceous sediment basin (siliceous rock including chert and siliceous mudstone); 5, Submarine rise; 6, Ancient land; 7, Hushan section; 8, Cited sections; (1) Chaohu section, 140 Anhui Province; (2) Meishan section, Changxing, Zhejiang Province; (3) Rencunping section, Sangzhi, Hunan Province. ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 3

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intercalated with medium-bedded grey argil- cephalopod Lopingoceras sp. in bed 89 of laceous limestones, thin-bedded calcareous the Hushan section (Fig. 3). The ammo- 265 mudstones and claystones (Fig. 3). It contains noids Huananoceras sp., Qianjiangoceras 220 abundant radiolarians, ammonoids, cono- sp., Pernodoceras sp., Pseudotirolites sp. donts, small brachiopods, bivalves and a and Pleuronodoceras sp. occur in bed 83 of small number of ostracods and foraminifera. the upper part of the Talung Formation at The lower Talung Formation at the Hushan Hushan. Hunanopecten exilis is regarded as 270 section is covered by Quaternary sediments. the index fossil of the Hunanopecten exilis 225 The basal part of the Lower Chinglung Zone of Late Permian age (Yin 1985). The Formation is composed mainly of yellowish cephalopod Lopingoceras sp. is common calcareous mudstones, thin- to medium- within the Upper Permian of South China bedded argillaceous limestones interbedded (Zhao et al. 1978). The above-mentioned 275 with pale claystones, and contains ammo- ammonoid taxa from the upper Talung 230 noids and bivalves (Fig. 3). Formation are regarded as index fossils of the Pseudotirolites–Pleuronodoceras Zone of late Changhsingian age (Zhao et al. 1978, Geological age Yang et al. 1987). The ammonoid Ophiceras 280 The bivalve Hunanopecten exilis Zhang, sp. was found in beds 94 and 95 of the basal 235 1977 was found in beds 73 and 88, and the part of the Lower Chinglung Formation at 4 WEIHONG HE et al. ALCHERINGA

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Fig. 3. Radiolarian ranges in the upper Talung Formation of the Hushan section, Nanjing, Jiangsu Province, South 365 China. Data marked by solid dots from this paper and data marked by blank dots revised after Zhang et al. (1992). 320 Hushan (Fig. 3). Ophiceras is an index genus olarian fauna from the upper Talung For- of the Ophiceras–Lytophiceras Zone, char- mation at Hushan is assignable to the late acterizing the earliest Triassic (Guo 1982). Changhsingian (latest Permian; Fig. 3). 370 Therefore, the upper Talung Formation is 325 assigned approximately to the late Changh- singian (latest Permian) at Hushan, and the Materials and methods basal part of the Lower Chinglung Forma- Twenty-two samples (six samples productive) tion at Hushan is assigned to the Early were collected from the greyish-black, thin- 375 Triassic (Fig. 3). Consequently, the radi- bedded, siliceous mudstones or cherts of the ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 5

upper Talung Formation, and five samples a deep sea along the northern margin of the derive from the siliceous mudstones, argillac- northeastern Yangtze Basin at the end of the 425 eous limestones and calcareous mudstones of late Palaeozic. Albaillella triangularis, Neoal- 380 the Lower Chinglung Formation in the baillella sp. cf. N. optima, Ormistonella Hushan section (Fig. 3). All samples were adhaerens, Ormistonella elegans, Shangella etched in a 1–5% solution of HF for 4–8 h at longa, Ishigaum craticula, Ishigaum fusinum, room temperature. The acid residues were Tetratormentum sp. 2, Entactinia itsukaichien- 430 subsequently transferred to other containers sis, Hegleria mammilla, Paroertlispongus 385 filled with water until neutral. Residual fontainei, Paurinella mesotriassica, Copicyn- samples were immersed in new HF solution. troides asteriformis, Tetraspongodiscus staur- After repeating this process more than 40 acanthus, Tetrapaurinella nanjingensis, times, adequate residues were sieved with a Paurinella mesotriassica, Klaengspongus for- 435 mesh diameter of 0.054 mm and dried for mosus and Klaengspongus spinosus, collec- 390 examination. We examined each species tively account for 77% of the species diversity primarily under a binocular microscope, then of the Hushan radiolarian fauna. These selected some specimens from each species species are common in Japan, Thailand, for scanning electron microscopy, and finally West Texas, the southern Alps and South 440 re-examined each species for morphological China, indicating a Palaeotethyan signature 395 consistency. (see the systematic palaeontology of this paper). It seems likely that the radiolarian Results connection between the Hushan area and 445 The radiolarian fauna from the Hushan Palaeotethyan sites occurred via a northern 400 section has been previously reported by link (i.e. along the northern margin of the Zhang et al. (1992), to contain Tetrator- northeastern Yangtze Basin) during the late mentum sp. 1 [identified as Tormentum Changhsingian. This is proposed because delicatum Nazarov & Ormiston, 1985 by late Changhsingian radiolarian faunas from 450 Zhang et al. (1992)], Entactinia itsukaichien- sections to the south of Hushan (such as 405 sis Sashida & Tonishi, 1985, Entactinia sp., Chaohu in Anhui Province, Meishan in Trilonche crassus He & Feng sp. nov. Zhejiang Province and Rencunping in [identified as Entactinosphaera sp. by Zhang Sangzhi, Hunan Province; Fig. 1), are et al. (1992)], Copicyntroides asteriformis dominated by Entactinaria and Spumellaria 455 Nazarov & Ormiston, 1985 and Tetrapaur- forms and are characterized by relatively 410 inella nanjingensis (Zhang, Wu & Liu, 1992) lower diversity, without Albaillellaria forms [identified as Stauroplegma nanjingensis (see He et al. 2005, 2008 for data from Zhang, Wu & Liu, 1992 by Zhang et al. Meishan and Chaohu see He et al. 2005, (1992)]. Here we re-studied this fauna, 2008; data from Rencunping unpublished). 460 recognizing 22 species belonging to 15 It is worth noting that diagenesis and 415 genera (excluding those elements discovered selective dissolution inevitably damage the by Zhang et al. 1992). These taxa are silica skeletons of radiolarians and influence described in the systematic palaeontology the taxonomic composition of assemblages. section. However, we consider that the absence of 465 albaillelids in Chaohu, Meishan and Re- 420 ncunping does not result from diagenesis or Palaeogeography selective dissolution. This is because first, Radiolarian faunas have an important albaillelids can be recognized even if diag- bearing on resolution of whether there was enesis is relatively intense. For example, 470 6 WEIHONG HE et al. ALCHERINGA

many albaillellarian forms have been recog- late Changhsingian of South China indicate nized from the Talung Formation of Shang- that radiolarian diversity and taxonomic si section, Guangyuan, Sichuan Province representation (especially the percentage of 520 (Yao & Kuwahara 1999) even though Latentifistularia and Albaillellaria forms) 475 diagenesis and dissolution in Shangsi were are useful bathymetric indices (He et al. more intense than in other studied sections 2008). The diversity (number of species and of South China [see plates 2 and 3 of Yao & genera) and the percentage of Latentifistu- Kuwahara (1999)]. Second, the damage (by laria and Albaillellaria forms (the ratio of 525 diagenesis and dissolution) to biogenetic Latentifistularia and Albaillellaria species to 480 silica should be less intensive in Chaohu and the total number of species) in radiolarian Rencunping than in Hushan, since less faunas appear to be higher in a deep-water cherts and more clay minerals constitute setting (He et al. 2008). Albaillellaria and the Talung Formation in Chaohu and Latentifistularia forms account for 50% of 530 Rencunping, and the damage (by diagenesis the fauna at species level in the upper 485 and dissolution) to biogenetic silica is Changhsingian of the Hushan section (24 relatively weaker in rocks with more clay species in 16 genera). This is higher than the minerals than in cherts (Blome & Reed specific and generic diversity and percentage 1993). Third, the frames of albaillellarian of Albaillellaria and Latentifistularia forms 535 forms are generally stronger than those of from Chaohu [see Fig. 4; 15 species in 12 490 entactinarian and spumellarian forms, be- genera; Latentifistularia forms accounting cause the frames of albaillellarian species for 30–40% at Chaohu and Albaillellaria are sheet-like and less easily dissolved, forms being absent; He et al. 2008]. The whereas the frames of two other kinds diversity at Meishan section is four species 540 consist of fine beams or spicules and are in three genera, and the representation of 495 more readily degraded. The abundance of Latentifistularia forms accounts for 40% of albaillellarian forms might indicate a dee- the assemblage (He 2006). At Rencunping, per-water setting compared with the faunas these values are seven species in six genera dominated by Entactinaria and Spumullar- and 21% Latentifistularia (unpublished 545 ia, which characterize waters ca 200 m deep data). The specific and generic diversity 500 (Kozur 1993). However, a small proportion and the percentage of Albaillellaria and of albaillellarian forms have been recorded Latentifistularia forms at Hushan, there- in Lower Permian shallow-water carbonate fore, are also higher than at Meishan and platform deposits in the Ural Mountains Rencunping (Fig. 4). This pattern suggests 550 (albeit only 32 albaillellarian specimens that bathymetry deepened from south to 505 found among over 1000 radiolarian speci- north across the northern part of the mens; see Nazarov & Ormiston 1985). The Yangtze Basin, reaching in excess of 200 m absence of Albaillellaria forms in the by Hushan. This hypothesis is supported by sections south of Hushan (Chaohu, the difference in lithology and biotic asso- 555 Meishan and Rencunping) during the late ciations from south to north (from Re- 510 Changhsingian suggests water depths were ncunping, Meishan, Chaohu to Hushan). possibly too shallow for deep-water albail- The upper Changhsingian is characterized lellarians. The representation of Albaillel- by grey medium-bedded carbonates at laria forms, therefore, favours a northerly Meishan (Yin et al. 2001) and by abundant 560 deepwater Palaeotethyan connection to grey cherty limestones at Rencunping 515 Hushan. (Zhang et al. 2009), whereas the upper Integrated sedimentological, biotic asso- Changhsingian at Hushan is dominated by ciation and radiolarian faunal data from the dark grey thin-bedded cherts and siliceous ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 7

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620 Fig. 4. Comparison of the late Changhsingian radiolarian faunas from the studied sections of South China. The 575 shaded area of the pie graphs indicates the percentage of Latentifistularia þ Albaillellaria forms within the radiolarian fauna of the different sections. The number of species and genera of these combined groups is indicated within the shaded area. 625 mudstones. Generally, the limestone and Order ALBAILLELLARIA Deflandre, 580 cherty limestone accumulated in shallower 1953 water than the siliceous (opaline) mudstone Family ALBAILLELLIDAE Deflandre, (He et al. 2008). The biotic association of 1953 the upper Changhsingian at Chaohu and 630 Meishan sections contains abundant small Albaillella Deflandre, 1952 585 benthic brachiopods or abundant benthic foraminifera (He et al. 2008, Song et al. Type species. Albaillella paradoxa Defla- 2009), whereas the upper Changhsingian at ndre, 1952 Hushan mainly contains abundant planktic 635 radiolarians, nektic ammonoids, conodonts, Albaillella hushanensis He & Feng sp. nov. 590 and a small number of brachiopods and (Fig. 5A–G) bivalves. This deepwater corridor constitu- tes the so-called Eastern Qinling-Dabie deep Etymology. Named for the type locality at sea, which apparently persisted until at Hushan. 640 least the end of the Permian (Fig. 1). 595 Furthermore, based on the discovery of Material. Holotype, HS-66-96. This species Lower Triassic radiolarians in the Tongbai is common in the Hushan section; 17 Mountains of Eastern Qinling (Du et al. specimens were examined by SEM and 1997), the collision between the North seven illustrated. 645 China and South China plates probably 600 occurred in the Middle to Late Triassic (Du Occurrence. Upper Changhsingian; Hushan, et al. 1997). South China.

Diagnosis. A species of Albaillella with 9–11 650 Systematic palaeontology horizontal bands and straight ventral and 605 All specimens described in this paper are dorsal margins. deposited in the Micropaleontology La- boratory, Faculty of Earth Sciences, China Measurements. Height of shell excluding University of Geosciences, Wuhan, People’s H-frame, 145–200 mm; width of shell ex- 655 Republic of China. The classification of cluding H-frame, 80–115 mm; ratio of 610 Radiolaria adopted herein follows De We- width to height, 0.42–0.58. Based on five ver et al. (2001). specimens. 8 WEIHONG HE et al. ALCHERINGA

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Description. Shell body straight, cone- 1999 Albaillella triangularis Ishiga, Kito & 800 shaped, with 9–11 horizontal bands and Imoto; Kuwahara, p. 92, pl. 3, figs 6–11. 755 straight ventral and dorsal margins. Bands [1999b] gradually becoming narrower (in width) 2000 Albaillella triangularis Ishiga, Kito & and shorter (in height) from aperture to Imoto; Sashida et al., pp. 795–797, figs apical cone. Apical cone straight. Ventral 7.13–7.16. [2000a] 805 wing straight, long, and protruding hor- 2000 Albaillella triangularis Ishiga, Kito & 760 izontally or slightly downwards from the Imoto; Sashida et al., p. 250, pl. 1, figs 6– third horizontal band counted from the 10, 13. [2000b] aperture. H-frame not completely pre- 2001 Albaillella triangularis Ishiga, Kito & served. Imoto; Shang et al., pl. 2, figs 2, 3. 810 2001 Albaillella triangularis Ishiga, Kito & 765 Discussion. The new species differs from Imoto; Wang & Shang, p. 113, pl. 1, figs Albaillella triangularis Ishiga, Kito & 17–23. Imoto, 1982 from the Upper Permian of 2007 Albaillella triangularis Ishiga, southwestern Japan in having more hor- Kito & Imoto; Jin et al., p. 10, figs 815 izontal bands and straight ventral and 4.1–4.6. 770 dorsal margins. The new species is similar to Albaillella triangularis of Caridroit & Material. One specimen was examined by De Wever, 1986 from the lower Changh- SEM and illustrated. singian of southwestern Japan in the 820 absence of a curved ventral margin, but Occurrence. Common in the Upper Permian 775 differs in the former having more hor- of Japan, north Thailand, Russian Far East, izontal bands and a straight dorsal mar- North America, Malaysia, the Philippines, gin, and the latter having a higher ratio of Turkey and South China. shell width to height (0.59–0.90). The new 825 species can be readily distinguished from Measurements. Height of shell excluding H- 780 other Albaillella species in its higher cone- frame, 130 mm; width of shell excluding H- shaped shell with more horizontal bands, frame, 70 mm; ratio of width to height, 0.54; and with straight ventral and dorsal length of ventral wing, 115 mm. Based on margins. one specimen. 830

785 Albaillella triangularis Ishiga et al., 1982 Discussion. This specimen is consistent with (Fig. 5H) Albaillella triangularis from the Upper 1982 Albaillella triangularis Ishiga, Kito & Permian of Nabejiri-yama, southwestern Imoto, p. 17, pl. 2, figs 8–10. Japan in its triangular outline, curved 835 1986 Albaillella triangularis Ishiga, Kito & ventral and dorsal margins, curved apical 790 Imoto; Caridroit & De Wever, pp. 58– cone, and wider horizontal bands in the 59, fig. 3. lower part of the shell.

840 3 Fig. 5. Scanning electron micrographs of the late Changxingian radiolarians from the Hushan section, Nanjing, 795 Jiangsu Province, South China. All scale bars ¼ 50 mm. Specimen designations, e.g. for HS-66-24—HS: Hushan section; 66: bed number; 24: specimen number. A–G, Albaillella hushanensis He & Feng sp. nov.; A, HS-66-24; B, HS- 66-27; C, HS-66-54; D, HS-66-57; E, holotype, HS-66-96; F, HS-66-63; G, HS-66-35. H, Albaillella triangularis Ishiga et al., 1982, HS-66-53. I–K, Albaillella sp.; I, HS-66-81; J, HS-66-89; K, HS-66-16. L, Neoalbaillella sp. 2, HS-66-18. M, N, Neoalbaillella sp. cf. N. optima Ishiga et al., 1982; M, HS-66-64; N, HS-66-34. O, P, Neoalbaillella sp. 1; O, HS- 845 66-17; P, HS-66-13. 10 WEIHONG HE et al. ALCHERINGA

Albaillella sp. (Fig. 5I–K) Neoalbaillella sp. 1 (Fig. 5O–P) 895 Material. This species is uncommon in the Material examined. This species is rare at 850 Hushan section. Three specimens were Hushan. Two specimens were examined by examined by SEM and illustrated. SEM and illustrated.

Occurrence. Upper Changhsingian; Hushan, Occurrence. Upper Changhsingian; Hushan, 900 South China. South China. 855 Measurements. Height of shell excluding H- Discussion. The cylindrical shell with rows of frame, 180–250 mm; width of shell excluding poresstronglysuggeststhisformisaspecies H-frame, 80–135 mm; ratio of width to of Neoalbaillella, but the poor preservation 905 height, 0.44–0.64; length of ventral wing, prevents a more resolved attribution. 860 105–170 mm. Based on three specimens. Neoalbaillella sp. 2 (Fig. 5L) Discussion. This material is similar to Material examined. One specimen was ex- Albaillella triangularis from the Upper amined by SEM and illustrated. 910 Permian of Nabejiri-yama, southwestern 865 Japan, in its triangular outline and curved Occurrence. Upper Changhsingian; Hushan, ventral and dorsal margins, but poor pre- South China. servation prevents confident attribution to a formal species. Discussion. A cylindrical shell with two 915 wings favours assignment to Neoalbaillella, 870 Neoalbaillella Takemura & Nakaseko, 1981 but the poor preservation prevents attribu- tion to an established species. Type species. Neoalbaillella ornithoformis Takemura & Nakaseko, 1981. Order LATENTIFISTULARIA Caridroit 920 et al., 1999 875 Neoalbaillella sp. cf. N. optima Ishiga et al., 1982 (Fig. 5M–N) Family ORMISTONELLIDAE De Wever & Caridroit, 1984 emend. De Wever et al., Material. Two specimens were examined by 2001 925 SEM and illustrated. 880 Ormistonella De Wever & Caridroit, 1984 Occurrence. Upper Changhsingian; Hushan, South China. Discussion. Ormistonella is similar to Nazar- ovella De Wever & Caridroit, 1984 in its 930 Discussion. This material is similar to four arms arrayed on a tetrahedral struc- 885 Neoalbaillella optima Ishiga, Kito & Im- ture, but the latter has a larger central part oto, 1982 from the Upper Permian of and four unequal arms, one being closed Nabejiri-yama, southwestern Japan, in its and small. cylindrical shell with an apical cone 935 slightly curved ventrally and its ladder- Type species. Ormistonella robusta De We- 890 shaped wings. However, the present speci- ver & Caridroit, 1984. mens are difficult to compare precisely with Neoalbaillella optima because of poor Ormistonella adhaerens Feng in Feng et al., preservation. 2006a (Fig. 6A–C) 940 ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 11

1997 Latentifistridae gen. et sp. indetermi- from Ormistonella adhaerens, as the latter nate A; Kuwahara et al., pl. 3, figs 10, has a row of pores located in each groove 11. of the arms, and has two dentate sides of 990 1998 Nazarovella scalae Caridroit & De each arm. 945 Wever; Wang et al., pl. 4, fig. 9. 2001 Nazarovella gracilis De Wever & Family LATENTIFISTULIDAE Nazarov Caridroit; Shang et al., pl. 4, fig. 6. & Ormiston, 1983 2006 Ormistonella adhaerens Feng in 995 Feng et al., p. 845, figs 10.11–10.16. Shangella Feng in Feng et al., 2006a 950 [2006a] Discussion. Shangella is similar to Material. This species is uncommon in the Quadricaulis Caridroit & De Wever, 1986 Hushan section. Three specimens were in its perforate central shell with hollow, 1000 examined by SEM and illustrated. tubular, perforate arms, but the latter has 955 Occurrence. Upper Changhsingian; South an inflated tetrahedral central shell with China. four tetrahedrally arranged cylindrical arms. Discussion. This species is distinguished from 1005 other forms of Ormistonella in having gutter- Type species. Shangella longa Feng in Feng 960 shaped arms with dentate sides and a row of et al., 2006a pores located in each groove of the arms. Shangella longa Feng in Feng et al., 2006a Ormistonella elegans (Feng in Feng & Liu, (Fig. 6F–K) 1010 1993) (Fig. 6D–E) 965 2006 Shangella longa Feng in Feng et al., 1993 Wangia elegans Feng in Feng & Liu, p. pp. 829–831, figs 6.6, 6.7, 6.9–6.15. 544, pl. 2, figs 16, 17. [2006a] 2006 Ormistonella elegans (Feng in Feng & 1015 Liu); Feng et al., p. 845, figs 10.17–10.20. Material. This species is common in the 970 [2006a] Hushan section. Ten specimens were exam- ined by SEM and six illustrated. Material. This species is uncommon in the Hushan section. Three specimens were Occurrence. Upper Changhsingian; South 1020 examined by SEM and two illustrated. China. 975 Occurrence. Upper Permian; South China. Discussion. A large, triangular central shell and three long arms are evident, despite the Diagnosis. A species of Ormistonella with end of each arm not being preserved in our 1025 four three-bladed arms, each arm consisting material. This species differs from Shangella 980 of three wide grooves and three round and regularis Feng in Feng et al., 2006a from the coarse ridges; central part of shell small. Talung Formation, Dongpan section, southern Guangxi, South China, in the Discussion. This species differs from latter having a smaller central shell and 1030 Ormistonella robusta De Wever & Cari- three shorter arms. 985 droit, 1984 from the Maizuru Group (Upper Permian), Japan, in the latter Family CAULETELLIDAE Caridroit having four U-shaped arms. It differs et al., 1999 12 WEIHONG HE et al. ALCHERINGA

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1080 ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 13

Ishigaum De Wever & Caridroit, 1984 Discussion. This species differs from Ishi- 1130 gaum trifustis De Wever & Caridroit, 1984 Type species. Ishigaum trifustis De Wever & from the Upper Permian of Japan in having Caridroit, 1984. a fine spongy structure covering the distal end of each arm. 1180 Ishigaum craticula Shang et al., 2001 (Fig. 1135 6L) Family PSEUDOLITHELIIDAE Kozur & Mostler, 1989 2001 Ishigaum craticula Shang et al., p. 233, pl. 1, figs 12–15. Tetratormentum Nazarov & Ormiston, 1985 1185 2006 Ishigaum craticula Shang et al.; Feng 1140 et al., p. 835, fig. 8.24. [2006a] Diagnosis. A pyramidal shell with four- rayed internal spicules enclosed in a rela- Material examined. One specimen was ex- tively thick spongy framework. amined by SEM and illustrated. 1190 Type species. Tetratormentum narthecium 1145 Occurrence. Upper Changhsingian; South Nazarov & Ormiston, 1985 China. Tetratormentum sp. 2 (Fig. 6P)

Discussion. The present species is similar to Material. One specimen was examined by 1195 Ishigaum tristylum Feng in Feng et al., SEM and illustrated. 1150 2006a from the Talung Formation, Dong- pan section, southern Guangxi, South Chi- Occurrence. Upper Permian; West Texas na, in having long arms with rows of pores, and South China. but differs in the former having an elongate 1200 and spongy distal part and the latter having Measurements. Height of pyramid except 1155 a subspherical and spongy distal end. terminal spines, 180 mm; basal width of pyramid, 220 mm; length of terminal spine, Ishigaum fusinum Feng in Feng et al., 2006a 65 mm; basal width of terminal spine, (Fig. 6M–O) 60 mm. Based on one specimen. 1205

1160 2006 Ishigaum fusinum Feng in Feng et al., Discussion. This species differs from Tetra- p. 837, figs 8.12–8.17. [2006a] tormentum sp. 1 [identified as Tormentum delicatum Nazarov & Ormiston, 1985 by Material examined. This species is common Zhang et al. (1992)], from the upper 1210 but not well preserved in the Hushan Talung Formation of the Hushan section 1165 section. Three specimens were examined by in its shorter and conical terminal spines. SEM and illustrated. It differs from Tetratormentum narthecium Nazarov & Ormiston, 1985 from the Occurrence. Upper Changhsingian; South Upper Carboniferous to the Lower Per- 1215 China. mian of the southern Urals and Tetra- 1170 3 Fig. 6. Scanning electron micrographs of the late Changxingian radiolarians from the Hushan section, Nanjing, Jiangsu Province, South China. All scale bars ¼ 100 mm. A–C, Ormistonella adhaerens Feng in Feng et al., 2006a; A, HS-66-11; B, HS-66-49; C, HS-66-79. D, E, Ormistonella elegans (Feng in Feng & Liu, 1993); D, HS-66-008; E, HS- 1220 66-39. F–K, Shangella longa Feng in Feng et al., 2006a; F, HS-62-11; G, HS-62-56; H, HS-66-009; I, HS-66-29; J, HS- 62-18; K, HS-66-10. L, Ishigaum craticula Shang et al., 2001, HS-66-98. M–O, Ishigaum fusinum Feng in Feng et al., 1175 2006a; M, HS-66-50; N, HS-66-59; O, HS-66-47. P, Tetratormentum sp. 2, HS-66-33. 14 WEIHONG HE et al. ALCHERINGA

tormentum acutum Sashida & Tonishi, Occurrence. Common in the Upper Permian 1270 1988 from the Upper Permian of central of eastern and northern Thailand and South 1225 Japan in its more rounded pyramidal shell China. and four thicker and conical terminal spines covered by a spongy structure. Measurements. Diameter of test along short The rarity and preservational state of the axis, 120–140 mm; diameter of test along 1275 material prevent us establishing a new long axis, 180–210 mm. Based on two speci- 1230 formal species. mens.

Order SPUMELLARIA Ehrenberg 1875 Discussion.Thismaterialisconsistentwith emend. De Wever et al., 2001 the description of Paroertlispongus fontainei 1280 Family OERTLISPONGIDAE Kozur & (Sashida)ofSashidaet al., 2000b from the 1235 Mostler in Dumitrica et al., 1980 Upper Permian of eastern Thailand in having an elliptical outline, and two polar spines of Paroertlispongus Kozur & Mostler, 1981 variable length. This species is similar to Paroertlispongus chinensis (Feng, 1992) from 1285 Diagnosis. An elliptical, spongy shell with the Triassic Muyinhe Formation, Yunnan 1240 two spines of variable length. Province, southwestern China in shell outline and presence of two polar spines, but differs Type species. Paroertlispongus multispinosus in having spines with variable length. Kozur & Mostler, 1981. 1290 Family PYRAMISPONGIIDAE Kozur & 1245 Paroertlispongus fontainei (Sashida in Sa- Mostler, 1978 emend. Dumitrica in De shida et al., 2000b) (Fig. 7G–H) Wever et al., 2001

2000 Pseudospongoprunum? chiangdaoensis Paurinella Kozur & Mostler, 1981 1295 Sashida in Sashida et al., p. 805, figs 1250 7.21, 23. [2000a] Type species. Paurinella curvata Kozur & 2000 Pseudospongoprunum? fontainei Sashi- Mostler, 1981. da in Sashida et al., pp. 255–256, pl. 2, figs 15–18. [2000b] Paurinella mesotriassica Kozur & Mostler, 1300 2002 Pseudospongoprunum? chiangdaoensis 1981 (Fig. 7I) 1255 Sashida in Sashida et al.; Sashida & Salyapongse, p. 695, fig. 3.24. 1981 Paurinella mesotriassica Kozur & 2002 Copiellintra fontainei (Sashida); Mostler, p. 50, pl. 44, fig. 1. Feng & Gu, pp. 803–804, figs 5.11, 1994 Paurinella mesotriassica Kozur & 1305 5.12. Mostler; Kozur & Mostler, p. 73, pl. 1260 2006 Paroertlispongus fontainei (Sashida); 15, fig. 7, pl. 16, fig. 4. Feng et al., p. 27, pl. 3, figs 4–7. 2002 Paroertlispongus? sp. Feng & Gu, p. [2006b] 805, figs 5.13, 5.14. 2008 Paroertlispongus fontainei (Sashida); 2006 Paurinella mesotriassica Kozur & 1310 He et al., p. 210, figs 5A–B. Mostler; Feng et al., p. 29, pl. 3, figs 8– 1265 11. [2006b] Material. This species is common but not well preserved in the Hushan section. Six Material. This species is rare in the Hushan specimens were examined by SEM and two section. One specimen was examined by 1315 illustrated. SEM and illustrated. ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 15

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1320

1370

1325

1375

1330

1380

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1345

1395

1350

1400

1355

1405

1360

1410 16 WEIHONG HE et al. ALCHERINGA

Occurrence. Upper Changhsingian of South 2006 Copicyntroides sp. Feng et al., p. 39, pl. China and Middle Triassic of the southern 7, fig. 20 [2006b]. Alps. 1460 Etymology. This species is named for the 1415 Measurements. Diameter of test, 115 mm; star-like outline of the shell. length of preserved long spine, 4100 mm; length of preserved short spine, 90 mm. Material. Holotype, HS-66-30. This species Based on one specimen. is common in the Hushan section. Twelve 1465 specimens were examined by SEM and eight 1420 Discussion. This specimen conforms to illustrated. Paurinella mesotriassica Kozur & Mostler, 1981 from the Middle Triassic of the Occurrence. Upper Changhsingian; Hushan, southern Alps in the form and arrangement South China. 1470 of rod-like spines of variable length. This 1425 species differs from Paurinella aequispinosa Diagnosis. A species of Copicyntroides with Kozur & Mostler, 1981 from the Triassic of seven massive, equatorial, three-bladed the southern Alps, by the latter having three spines covered by spongy structures. spines of equal length. 1475 Measurements. Diameter of test, 180– 1430 Family RELINDELLIDAE Kozur & Mos- 230 mm; length of spines, 70–130 mm; basal tler in Dumitrica et al., 1980 width of spines, 35–90 mm; ratio of spine basal width to spine length, 0.50–0.93 Copicyntroides Nazarov & Ormiston, 1985 (average 0.70). Based on five specimens. 1480

1435 Diagnosis. Shell discoidal, consisting of one Description. Star-like in outline; test (ex- spongy outer shell and nine or ten con- cluding spines) discoid, flat on both sides; centric inner tests, with 6–9 equatorial test is covered by surficial spongy structures, three-bladed spines. and consists of numerous equally and 1485 densely spaced concentric disks. The seven 1440 Discussion. Copicyntroides is similar to equatorial three-bladed spines are unequal Klaengspongus Sashida in Sashida et al., in length and covered by spongy structures. 2000b in having a discoidal shell, but the latter has an elevated central part and Discussion. This species is similar to Copi- 1490 prominently depressed flanks on both sides, cyntroides? sp. A of Sashida et al. (2000b) 1445 whereas the former has two flat sides. from the Changhsingian of eastern Thailand in its discoidal shell with sturdy, three- Type species. Copicyntroides asteriformis bladed and equatorial spines, but differs in Nazarov & Ormiston, 1985. that the former has seven stronger spines 1495 and thick spongy structures covering the 1450 Copicyntroides stellatus He & Feng sp. nov. proximal part of each spine and the latter (Fig. 8A–H) has eight thinner spines. The new species is 3 Fig. 7. Scanning electron micrographs of the late Changxingian radiolarians from the Hushan section, Nanjing, 1500 Jiangsu Province, South China. All scale bars ¼ 50 mm. A–C, Klaengspongus formosus Feng in Feng et al., 2007b; A, 1455 HS-75-002; B, HS-75-006; C, HS-88-009. D–F, Klaengspongus spinosus Sashida in Sashida et al., 2000b; D, HS-87-13; E, HS-87-007; F, HS-87-12. G, H, Paroertlispongus fontainei (Sashida in Sashida et al., 2000b); G, HS-87-008, ss indicates scar of broken spine; H, HS-87-10. I, Paurinella mesotriassica Kozur & Mostler, 1981, HS-62-23. J–M, Entactinia itsukaichiensis Sashida & Tonishi, 1985; J, HS-66-48; K, HS-66-51; L, HS-66-68; M, HS-66-80. ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 17

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1560

1515

1565

1520

1570

1525

1575

1530

1580

1535

1585

1540

1590

1545

1595

1550 18 WEIHONG HE et al. ALCHERINGA

similar in outline to Copicyntroides parvulus na, in having a spongy, flat outer spherical 1600 Feng in Feng et al., 2006b from the Talung test, four coplanar three-bladed external Formation of southern Guangxi, South spines, and in the ratio of the length of China, but differs by the latter having 9–12 spine to diameter of test. This species differs longer spines that are not covered by from Tetraspongodiscus tetragonius Feng in 1650 spongy structures. The new species differs Feng et al., 2006b from the Talung 1605 from Copicyntroides sp. of He et al. (2005) Formation, southern Guangxi, South Chi- from the Meishan D section since the latter na, by the latter having a quadrangular shell has thinner conical main spines. and more massive main spines. It differs from Tetraspongodiscus inaequispinosus 1655 Tetraspongodiscus Kozur & Mostler, 1979 (Kozur & Mostler, 1981) since the latter 1610 has a higher ratio of spine length to test Type species. Tetraspongodiscus longispino- diameter. sus Kozur & Mostler, 1979. Family PALAEOLITHOCYCLIIDAE Ko- 1660 Tetraspongodiscus stauracanthus Feng in zur & Mostler, 1989 1615 Feng et al., 2006b (Fig. 8I–K) Klaengspongus Sashida in Sashida et al., 2005 Lepingosphaera stauracanthus Feng in 2000b emend. Feng & Gu, 2002 Feng et al.; He et al., pp. 215–217, figs 1665 5.1–5.5. Type species. Klaengspongus spinosus Sashi- 1620 2006 Tetraspongodiscus stauracanthus Feng da in Sashida et al., 2000b. in Feng et al., pp. 39–40, pl. 9, figs 5–16. [2006b] Klaengspongus formosus Feng in Feng et al., 2008 Tetraspongodiscus stauracanthus Feng 2007b (Figs 7A–C, 8L) 1670 in Feng et al.; He et al., p. 208, fig. 4E–J. 1625 2007 Klaengspongus formosus Feng in Feng Material. Four specimens were examined by et al., pl. 2, figs 6–11. [2007b] SEM and three illustrated. 2008 Klaengspongus formosus Feng in Feng et al.; He et al., p. 209, fig. 4O–S. 1675 Occurrence. Common in the upper Changh- 1630 singian of South China and Japan. Material. This species is common in the Hushan section. Nine specimens were ex- Measurements. Diameter of test, 90– amined by SEM and four illustrated. 175 mm; length of well-preserved spines, 1680 45–160 mm. Based on two specimens. Occurrence. Common in the Changhsingian 1635 of South China. Discussion. The studied specimens are con- sistent with Tetraspongodiscus stauracanthus Diagnosis. Outline discoidal, test consisting Feng in Feng et al., 2006b from the Talung of a central phacoid sphere and a flat 1685 Formation, southern Guangxi, South Chi- peripheral ring. 1640 3 Fig. 8. Scanning electron micrographs of the late Changxingian radiolarians from the Hushan section, Nanjing, Jiangsu Province, South China. All scale bars ¼ 50 mm. A–H, Copicyntroides stellatus He & Feng sp. nov.; A, HS-66- 1690 002; B, HS-66-97; C, HS-66-40; D, HS-66-31; E, HS-66-52; F, HS-66-46; G, HS-66-60; H, holotype, HS-66-30. I–K, Tetraspongodiscus stauracanthus Feng in Feng et al., 2006b; I, HS-66-001; J, HS-66-005; K, HS-66-99. L, 1645 Klaengspongus formosus Feng in Feng et al., 2007b, HS-87-005. ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 19

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1745

1700

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1705

1755

1710

1760

1715

1765

1720

1770

1725

1775

1730

1780

1735

1785 20 WEIHONG HE et al. ALCHERINGA

Measurements. Diameter of test, 175– Material. Five specimens were examined by 250 mm; diameter of centric part, 80– SEM and three illustrated. 1835 125 mm. Based on four specimens. 1790 Occurrence. Common in the Discussion. The present material is consis- Changhsingian of South China and east- tent with Klaengspongus formosus Feng in ern Thailand. Feng et al., 2007b from the Talung Forma- 1840 tion, southern Guangxi, South China, in its Measurements. Diameter of test excluding 1795 discoidal outline, a large phacoid central outer spines, 200–275 mm; diameter of sphere, flat peripheral ring and absence of centric part, 150–200 mm. Based on three an outer spine, but material from the type specimens. locality is generally larger (324–427 mm). 1845 The species is similar to Klaengspongus Discussion. This material is consistent with 1800 spinosus Sashida in Sashida et al., 2000b Klaengspongus spinosus Sashida in Sashida from the Changhsingian of eastern Thailand et al., 2000b from the Changhsingian of in lateral view, but the latter differs by eastern Thailand in its discoidal outline, having a larger central phacoid sphere and phacoid central sphere, flat peripheral ring 1850 gear-shaped outer spines. and prominent outer spines. 1805 Klaengspongus spinosus Sashida in Sashida Order ENTACTINARIA Kozur & Mos- et al., 2000b (Fig. 7D–F) tler, 1982 1855 2000 Orbiculiforma? sp. Feng et al., pp. 312– Family ENTACTINIIDAE Riedel, 1967 1810 313, figs 2.1–2.4. 2000 Orbiculiformidae gen. et sp. indeter- Entactinia Foreman, 1963 minate. Sashida et al., p. 805, figs 7.25– 7.27. [2000a] Type species. Entactinia herculea Foreman, 1860 2000 Klaengspongus spinosus Sashida in 1963. 1815 Sashida et al., p. 256, pl. 3, figs 7, 8. [2000b] Entactinia itsukaichiensis Sashida & To- 2001 Copicyntroides sp. Shang et al., pl. 2, nishi, 1985 (Figs 7J–M, 9A–B) fig. 19. 1865 2001 Spongosphaeradiscus shaiwaensis 1985 Entactinia itsukaichiensis Sashida & 1820 Wang in Wang & Shang, pl. 1, figs 24– Tonishi, p. 9, pl. 1,figs 1–6. 33. 1992 Stauracontium sp. Zhang et al., pl. 3, 2002 Klaengspongus spinosus Sashida in figs 6, 9. Sashida et al.; Feng & Gu, p. 805, figs 1992 Copicyntra akikawaensis Sashida & 1870 6.1–6.14, 8.1–8.7, 9.1–9.4. Tonishi; Zhang et al., pl. 3, fig. 7. 1825 2008 Klaengspongus spinosus Sashida in 1998 Entactinia itsukaichiensis Sashida & Sashida et al.; He et al., pp. 209–210, Tonishi; Kuwahara & Yao, p. 36, pl. 2, fig. 4T–V. fig. 59. 1875 3 1830 Fig. 9. Scanning electron micrographs of the late Changxingian radiolarians from the Hushan section, Nanjing, Jiangsu Province, South China. All scale bars ¼ 50 mm. A, B, Entactinia itsukaichiensis Sashida & Tonishi, 1985; A, HS-66-28; B, HS-66-44. C–I, Trilonche crassus He & Feng sp. nov.; C, holotype, HS-86-002; D, HS-86-003; E, HS-86- 004; F, HS-86-005; G, HS-86-006; H, HS-86-007; I, paratype, HS-66-41. J, K, Hegleria mammilla (Sheng & Wang, 1985); J, HS-87-003; K, HS-87-002. L, Paracopicyntra sp., HS-66-006. 1880 ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 21

2000 Entactinia itsukaichiensis Sashida & spines and other smaller needle-like sub- Tonishi; Sashida et al., p. 251, pl. 1, figs ordinate spines according to the emended 24–26 [2000b]. definition of Aitchison & Stratford (1997). 1930 2005 Entactinia itsukaichiensis Sashida & 1885 Tonishi; He et al., p. 210, figs 4.7–4.8. Type species. Trilonche vetusta Hinde, 1899 2007 Entactinia itsukaichiensis Sashida & Tonishi; Feng et al., p. 20, pl. 1,figs 1–2. Trilonche crassus He & Feng sp. nov. (Fig. [2007c] 9C–I) 1935 2008 Entactinia itsukaichiensis Sashida & 1890 Tonishi; He et al., pp. 202–203, figs 3A– 1992 Entactinosphaera sp. Zhang et al., pl. L. 3, fig. 12.

Material. This species is common in the Etymology. This species is named for the 1940 Hushan section. Twenty specimens were thick outer shell. 1895 examined by SEM, and six illustrated. Material. Holotype, HS-86-002; paratype, Occurrence. Common in the Upper Permian HS-66-41. This species is common in the of central Japan, USA, Italy, Thailand and Hushan section. Seven specimens were 1945 South China. examined by SEM and illustrated. 1900 Diagnosis. A species of Entactinia with a Occurrence. Upper Changhsingian; Hushan, small lattice shell, six three-bladed main South China. spines and numerous needle-like by-spines 1950 on the shell surface. Diagnosis. A species of Trilonche with a 1905 thick outer shell and numerous spine-like Measurements. Diameter of test, 80– nodes arising from junctions of pore frames 125 mm. Based on six specimens. of outer shell. 1955 Discussion. This species is similar to En- Measurements. Diameter of test, 180– 1910 tactinia modesta Sashida & Tonishi, 1985 200 mm; basal width of main spines, 25– from the Upper Permian of Itsukaichi, 40 mm, thickness of cortical shell, 25– central Japan in shell outline, but differs in 30 mm. Based on seven specimens. having a smaller shell. It is similar to 1960 Entactinia parapycnoclada Nazarov & Or- Description. Test medium-sized, spherical 1915 miston, 1985 from the Bone Springs Lime- and latticed, with 140–160 subcircular pores stone and Lamar Limestone of West Texas per hemisphere. The main spines are mas- in the shape and size of the shell and in the sive, three-bladed, with round ridges and shape of the main spines, but the latter deep grooves. The main spines slightly 1965 differs by having torsional main spines and descrease in width towards their distal parts. 1920 by-spines of non-uniform length and dia- Numerous spine-like nodes arise from junc- meter. tions of the pore frames and thicken the cortical shell. One latticed medullary shell Trilonche Hinde, 1899 emend. Foreman, connects to the cortical shell by six three- 1970 1963 emend. Aitchison & Stratford, 1997 bladed beams. 1925 Diagnosis. Trilonche has two or more Discussion. This species resembles Wuyia spherical shells, up to six three-bladed main dongpanica Feng in Feng et al., 2007c 22 WEIHONG HE et al. ALCHERINGA

1975 from the Talung Formation of southern 2006 Hegleria mammilla (Sheng & Wang); Guangxi, South China, in its numerous Wonganan & Caridroit, p. 6, pl. 1, fig. spine-like nodes arising from the pore 29. frames, but the former has sturdy main 2007 Hegleria mammilla (Sheng & Wang); 2025 spines and two concentric shells, whereas Feng et al., pp. 12–15, pl. 5, figs 14–16. 1980 the latter has three concentric shells. The [2007c] new species is similar to Trilonche textilis 2008 Hegleria mammilla (Sheng & Wang); Feng in Feng et al., 2007c from the He et al., pp. 205–206, figs 3T–U, 4A– Talung Formation of southern Guangxi, C. 2030 South China in having numerous spine- 1985 like nodes arising from the pore frames, Material. Five specimens were examined by but the former has massive, three-bladed SEM and two illustrated. main spines, and more and smaller pores on the outer shell. The new species is Occurrence. Common in the Middle and 2035 distinguished from other species of Tri- Upper Permian of Thailand, North Amer- 1990 lonche by its thick cortical shell and spine- ica, Sicily and South China. like nodes on the outer shell. Measurements. Diameter of test, 250– Hegleria Nazarov & Ormiston, 1985 260 mm; number of mammae, 70–84. Based 2040 on two specimens. 1995 Type species. Hegleria mammifera Nazarov & Ormiston, 1985. Discussion. The shape and size of the external sphere, number of conical mammae Hegleria mammilla (Sheng & Wang, 1985) on the surface, and each mamma having 2045 (Fig. 9J–K) numerous small pores framed by thick bars, 2000 suggest that these specimens are referable to 1985 Phaenicosphaera mammilla Sheng & Hegleria mammilla (Sheng & Wang, 1985) Wang, pp. 179–180, pl. 3,figs 1–8. first described from the Kufeng Formation, 1985 Hegleria mammifera Nazarov & Or- Longtan, Nanjing, Jiangsu Province, South 2050 miston, p. 22, pl. 6,figs 3–5. China. 2005 1987 Phaenicosphaera mammilla Sheng & Wang; Kozur & Krahl, pp. 365–366, fig. Paracopicyntra Feng in Feng et al., 2006b 7a. 1990 Hegleria mammifera Nazarov & Or- Type species. Copicyntra ziyunensis Feng & 2055 miston; Noble & Renne, p. 384, pl. 1, Gu, 2002 2010 figs 9, 10. 1992 Hegleria mammilla (Sheng & Wang); Paracopicyntra sp. (Fig. 9L) Blome & Reed, p. 369, pl. 11, figs 10, 12, 13. Material. This species is rare in the Hushan 2060 1994 Hegleria mammilla (Sheng & section. One incomplete specimen was 2015 Wang); Wang & Li, pp. 209–210, pl. 1, examined by SEM and illustrated. figs 22, 23. 2000 Hegleria mammilla (Sheng & Wang); Occurrence. Upper Changhsingian; South Sashida et al., p. 252, pl. 3, figs 1–4, 6. China. 2065 [2000b] 2020 2002 Hegleria mammilla (Sheng & Wang); Measurements. Diameter of test, 150 mm. Feng & Gu, p. 801, figs 4.1–4.3. Based on one specimen. ALCHERINGA PERMIAN RADIOLARIANS FROM CHINA 23

Discussion. The studied specimen conforms CARIDROIT, M., DE WEVER,P.&DUMITRICA, P., 1999. 2070 to Paracopicyntra Feng in Feng et al., 2006b Un nouvel ordre, une nouvelle famille et un nouveau genre de radiolaires du Paleozoique: in having more than four concentric shells Latentifistularia, Cauletellidae et Cauletella. and in the distance between two adjacent Compte Rendu de la Acade´mie des Sciences concentric shells. Francaise, Sciences de la terre et des planets 329, 2120 603–608. CATALANO, R., STEFANO, P.DI.&KOZUR, H., 1991. 2075 Permian circumpacific deep-water fauna from the Acknowledgements western Tethys (Sicily, Italy)—new evidence for the The authors express sincere thanks to Wu Jun position of the Permian Tethys. Palaeogeography, Palaeoclimatology, Palaeoecology 87, 75–108. and Zhang Suxin for photography, and to Ms 2125 DE WEVER,P.&CARIDROIT, M., 1984. Description de Liu Weihong from the Nanjing Institute of quelques nouveaux Latentifistulidae (Radiolaires 2080 Geology and Mineral Resources, and Prof. polycystines) paleozoiques du Japon. Revue de Wu Shunbao for offering much help with Micropale´ontologie 27, 98–106. DE WEVER, P., DUMITRICA, P., CAULET, J.P., NIGRINI, fieldwork. We wish to express sincere thanks C. & CARIDROIT, M., 2001. Radiolarians in the to four anonymous reviewers and the editor Sedimentary Record, Gordon and Breach Science 2130 of Alcheringa for their reviews of the manu- Publishers, Singapore, 533 pp. 2085 script and for offering good suggestions that DEFLANDRE, G., 1952. Albaillella nov. gen., du Carbo- nife` re infe´rieur, type d’une lignee aberrante eteinte. improved the paper. This research has been Comptes Rendus Acade´mie des Sciences, Paris 223, supported by NSFC (Grant Nos. 40872008, 872–874. 40921062, 40502001), the Ministry of Educa- DEFLANDRE, G., 1953. Radiolaires fossils. In Traite de 2135 tion of China (NCET-10-0712, B08030) and Zoologie,P.P.GRASSE, Masson et Cie, ed., Paris, 389–486. 2090 the Fundamental Funds for the Central DU,Y.S.,FENG, Q.L., YIN, H.F., ZHANG,Z.H.&ZENG, Universities. X.Y., 1997. The Late Hercynian–Early Indosinian Eastern Qinling-Dabie Sea. Scientia Geologica Sinica 32, 129–135 (in Chinese with English abstract). 2140 References DUMITRICA, P., KOZUR,H.&MOSTLER, H., 1980. ABELMANN, A., BRATHAUER, U., GERSONDE, R., SIEGER, Contribution to the radiolarian fauna of the Geologisch- 2095 R. & ZIELINSKI, U., 1999. Radiolarian-based Middle Triassic of the Southern Alps. transfer function for the estimation of sea-surface Pala¨ontologische Mitteilungen Innsbruck 10, 1–46. temperatures in the Southern ocean (Atlantic EHRENBERG, C.G., 1875. Fortestzung der mikrogeolo- gischen studien als gesammtuebersicht der mikros- sector). Paleoceanography 14, 410–421. 2145 AFANASIEVA, M.S. & ZAMILATSKAY, T.Y., 1993. The kopischen palaeontologie gleichartig analysirter paleobiogeography of the northeast Pricaspian gebirgsarten der erde, mit specieller rucksicht auf 2100 Basin and pre-Uralian Depression in Artinskian den polycystinen-mergel von Barbados. Abhandlun- time based on Radiolaria and Foraminifera. gen der Koeniglische Akademie der Wissenschaften, Micropaleontology, special issue 6, 61–65. zur Berlin (Jahrgang) 1875, 1–226. MPSON ORIN AITCHISON, J.C. & STRATFORD, J.M.C., 1997. Middle E -M , K.M., 1984. Depth and latitude Devonian (Givetian) Radiolaria from eastern New distribution of radiolaria in Campanian (Late 2150 South Wales, Australia: a reassessment of the Cretaceous) tropical and subtropical oceans. Mi- cropaleontology 30, 87–115. 2105 Hinde (1899) fauna. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie Abhandlungen 203, 369–390. FENG, Q.L., 1992. Permian and Triassic radiolarian Earth BLOME, C.D. & REED, K.M., 1992. Permian and Early biostratigraphy in South and Southwest China. (?) Triassic radiolarian faunas from the Grindstone Science—Journal of China University of Geosciences terrane, central Oregon. Journal of Paleontology 66, 3, 51–62 (in Chinese with English abstract). 2155 351–383. FENG, Q.L. & GU, S.Z., 2002. Uppermost Changhsin- BLOME, C.D. & REED, K.M., 1993. Acid processing of gian (Permian) radiolarian fauna from southern 2110 pre-Tertiary radiolarian cherts and its impact on Guizhou, southwestern China. Journal of Paleon- faunal content and biozonal correlation. Geology tology 76, 797–809. 21, 177–189. FENG, Q.L. & LIU, B.P., 1993. Late Permian and Early– Middle Triassic radiolarians from southwestern CARIDROIT,M.&DE WEVER, P., 1986. Some Late 2160 Permian radiolarians from pelitic rocks of the Yunnan. Earth Science—Journal of China Unviver- Tatsuno Formation (Hyogo Prefecture), Southwest sity of Geosciences 18, 540–552 (in Chinese with 2115 Japan. Marine Micropaleontology 11, 55–90. English abstract). 24 WEIHONG HE et al. ALCHERINGA

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