Devonian (upper Emsian -lower Givetian) radiolarians ...

Devonian (upper Emsian - lower Givetian) radiolarians from the Tanhe Formation, Nanning, Guangxi, southwest China Hui LUO", Jonathan C.

itch is on* and Yu-jing wang'

'Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing 210008, China

email: [email protected] ' ~ e ~ a r t m eof n tEarth Sciences, University of Hong Kong, Pokfulam Road, Hong Kong SAR, China

email: jona@ hku.hk

ABSTRACT: A moderately well-preserved radiolarian fauna consisting of 20 Devonian (upper Emsian-lower Givetian) taxa was recovered from siliceous strata in the Wuxiangling section, Nanning, Guangxi, southwest China. This fauna is dominated by spumellarians, but also includes some distinctive ceratoikiscid forms that have potential for biostratigraphic studies and regional correlation. Six new species, Ceratoikiscum coroniferum n. sp., Stigmosphaerostyluscubicus n. sp., Trilonche nanningensis n. sp., Trilonche remosa n. sp., Trilonche tanheensis n, sp. and Trilonche xinpoensis n. sp., are described.

INTRODUCTION

The Early to Middle Devonian was an important time in radiolarian evolution as major differences exist between Upper Silurian and Upper Devonian radiolarian faunas (Nazarov and Ormiston 1985, 1986, 1993; Stratford and Aitchison 1997). Previous studies from Australia, Japan, Europe and China have yielded important insights into the characteristics of the Lower to Middle Devonian radiolarians and their evolution from Silurian to Late Devonian (Hinde 1899; Kiessling and Tragelehn 1994; Stratford and Aitchison 1997; Aitchison et al. 1999; Kurihara and Sashida 2000; Umeda 1998). These authors also documented the importance of Lower to Middle Devonian radiolarians in paleotectonic reconstructions. Detailed study of Lower to Middle Devonian radiolarian fossils, however, remains in its infancy, especially in China. Lower to Middle Devonian radiolarians have been found recently in many different localities in China, mostly in south China. Sheng and Wang (1982) first reported a Middle Devonian radiolarian fauna from Xintian, Hunan Province. Wu (1985) also found some Middle Devonian radiolarians from the chert belts in Yingde area in northern Guangdong but did not describe them. Feng and Liu (1992) reported some Lower Devonian radiolarians from western Yunnan and named a new radiolarian genus Eoalbaillella. According to Wang et al. (2000) this fauna may be Middle Devonian. Wang et al. (2000) also described some Middle Devonian radiolarians from Xiangyun and Lancang, western Yunnan. In this paper we describe moderately well-preserved Lower to Middle Devonian (upper Emsian-lower Givetian) radiolarians from the Tanhe Formation in the Wuxiangling section, Nanning, Guangxi, southwest China (text-fig. 1). Twenty radiolarian taxa are systematically examined, including six new species. Their documentation will help to further refine the Lower to Middle Devonian radiolarian biostratigraphy of southwest China.

STRATIGRAPHY The Wuxiangling section is located near Xinpo village in the Wuxiangling area (N22"45'53", E108"19'49"), about 4 km south from Nanning, the capital of Guangxi (text-fig. 1). The highway from Nanning to Xinpo passes through this section. Lower to Upper Devonian strata are well exposed in a road-cut along a county road (text-fig. 1) except for small portions covered by soil. The stratigraphic sequence is simple. No definite hiatuses in sedimentation were detected in this section and no complex folds or faults were observed. Fossils are abundant and include planktic tentaculitids, radiolarians and ammonites, as well as some benthic fossils such as brachiopods, corals and crinoids. These fossils, mainly the tentaculitids (Zhong et al. 1992; Kuang et al. 1996), provide good age control for each formation exposed in this section.

Six distinctive formations have been recognized by Zhong et al. (1992) and Kuang et al. (1996) in the Guangxi area. In ascending order they are: Yujiang, Moding, Tanhe, Luofu, Liujiang and Wuzhishan formations. The lowermost Yujiang Formation is considered to be a shallow-water littoral facies deposit, whereas the others accumulated in deep-water environments. In the Wuxiangling section, the lower part of the Tanhe Formation is covered by soil or vegetation, and partly by Tertiary sandstone and conglomerate. The upper part of the formation is well exposed. These upper strata are about 70m thick and consist of thin-bedded dark grey, brown grey cherty rock and cherty mudstone. The top of the Tanhe Formation is conformably overlain by black mudstone of the Luofu Formation. The Tanhe Formation was described by Zhong et al. (1992) as an uppermost Lower Devonian to lower Middle Devonian sequence of yellowish brown, grey brown, and dark thin bedded siliceous rocks, and siliceous shales locally intercalated with dark chert bands that are exposed in the Baise-Nanning region. Strata of the Tanhe Formation exposed in the Wuxiangling section were previously mapped as Dongganlin Formation by the Guangxi Geological Survey Team (see Zhong et al. 1992),

micropaleontology, volume 48, supplement no. 1,2002,pp. 113-127, text-figures 1-2, plates 1-3,2002

113

H . Luo, J . C . Aitchison and Y. Wang: Devonzan radiolariansfi-om the Tanhe Fornzation, Nanning, Guangxi, southwest China

al. 1992). The upper part of the overlying Luofu Formation (text-fig. 2) contains the lower Givetian tentaculitid Nowakia otomari. RADIOLARIAN ASSEMBLAGE

Abundant radiolarians have been recovered from thin-bedded siliceous cherty rocks in the upper (Givetian) part of the Tanhe Formation in the Wuxiangling section (Plate 1 through Plate 3). Radiolarians were found in 4 of 14 samples analyzed (text-fig. 2) and were extracted using standard HF acid-leaching techniques (Pessagno and Newport 1972). The assemblage includes the following taxa: Ceratoikiscum coroniferum n. sp., Ceratoikiscum planistellare Foreman, Ceratoikiscum sp., Helioentactinia sp. cf. H . multa (Won), Magnisphaera sp., Palaeoscenidium cladophorum Deflandre, Spongentactinia sp., Stigmosphaerostylus cubicus n. sp., Stigmosphaerostylus hystricuosa (Aitchison), Stigmosphaerostylus sp., Trilonche dat'idi (Hinde), Trilonche echinata (Hinde), Trilonche cf. T. echinata (Hinde), Trilonche minax (Hinde), Trilonche nanningensis n. sp., Trilonche remosa n. sp., Trilonche tanheensis n. sp., Trilonche vestusta Hinde, Trilonche xinpoensis n. sp., and Trilonche sp.

TEXT-FIGURE 1

Index map showing the location of the Wuxiangling Section in Nanning,

Guangxi, southern China (after Zhong et al. 1992).

called Nabiao Formation by Xian et al. (1980), correlated with the Nabiao Formation by Wu et al. (1987), and assigned to Liujiang Formation (in part) by Yin (1997). Although abundant radiolarians were recovered in this investigation other fossils in the Wuxiangling section are much rarer. The tentaculitids Nowakia (N.) elegans (Barrande 852) Nowakia (N.) cancellata (Richter 1854) Nowakia sp. and Styliolina sp. were reported by Kuang et al. (1996) from the lower part of the Tanhe Formation together with Styliolina sp. and Nowakia sp. from the upper part of the unit. These tentaculitids indicate that the Tanhe Formation (text-fig. 2) possibly ranges from upper Emsian to lower Givetian. The lower part of the Moding Formation (text-fig 2) contains the tentaculitids Nowakia barrandei Boucek and Prantl 1959 Nowakia elegans and Viriatellina pseudogeinitziana Boucek 1964. In addition, the Moding Formation also yields ammonites Erbenoceras sp. and Teicherticeras sp., corals Favosites sp., and Heterophrentis sp., and brachiopods which include Atrypa sp., and Schizophoria sp. This fossil assemblage indicates that the lower part of the Moding Formation is uppermost lower Emsian (Zhong et

cubicus n. sp. only. Sample 96~x1135yields Stigmosphaerostyl~~s Most of the other taxa occur in samples 96~x1128and 96~x1129 within a 30 cm thick interval located about 5.5m below the top of the Tanhe Formation. This radiolarian assemblage, referred herein as the Ceratoikiscum coroniferum-Trilonche remosa assemblage, is characterized by abundant Ceratoikiscum coroniferum n. sp, Trilonche remosa n. sp. (with branching main spines), Stigmosphaerostylus cubicus n. sp. (with cube-like shell) and Trilonche xinpoensis n. sp. (with twisted main spines). Other importantAtaxa include Ceratoikiscum planistellare, Ceratoikiscum sp., Magnisphaera sp., Stigmosphaerostylus hystricuosa and Stigmosphaerostylus sp. In the Wuxiangling section the Ceratoikiscum coroniferum-Trilonche remosa assemblage (text-fig. 2) is the most diverse in terms of the numbers of radiolarian genera and species. Occurrences of Middle Devonian radiolarians are also known from the Qiziqiao Formation of the Middle Devonian at Matangwuo in Xintian district of Hunan province, China. Four genera and seven species from a poorly preserved fauna were described by Sheng and Wang (1982). They are all Spumellaria with spongy . -. shells. This radiolarian fauna has a much lower diversity than that of the Ceratoikiscum coroniferum-Trilonche remosa assemblage from the Tanhe Formation in the Wuxingling section. Although the Ceratoikiscids within the Ceratoikiscum coroniferum-Trilonche remosa assemblage are different from those of the Ceratoikiscum regalinodus assemblage, interpreted to be upper Givetian from the Gamilaroi terrane, New England Orogen in eastern Australia (Stratford and Aitchison 1997; Aitchison et al. 1999), the entactiniids in both assemblages are very similar. SYSTEMATIC PALEONTOLOGY

All taxa described in this paper are illustrated using scanning electron micrographs. Specimens illustrated are assigned a catalog number and deposited in the Department of the Earth Sciences, The University of Hong Kong (abbreviated HKU).

Micropaleontology, vol. 48, no. 1,2002

Legend limestone

chert with

siliceous shale

chert band

chert

mudstone

covered .~ ntewal

TEXT-FIGURE 2

Range chart for radiolarians of Tanhe Formation in the Wuxiangling section, Nanning, Guangxi, southern China.

Subclass RADIOLARIA Miiller 1858 Order POLYCYSTIDA Ehrenberg 1838 Suborder ALBAILLELLARIA Deflandre 1953; emend. Holdsworth 1969 Family CERATOIKISCIDAE Holdsworth 1969 Genus Ceratoikiscum Deflandre 1953 Type species: Ceratoikiscum avimexpectans Deflandre 1953

Remarks: This species is distinguished by its crown-like patagial tissue and by the spherical cavea in the center of test. The spherical cavea bears some resemblance to Ceratoikiscum regalinodus Stratford and Aitchison 1997. This species, however, does not have the crown-like patagial tissue and possesses distinctive prominent nodes at the end of the extra-triangular rods.

Ceratoikiscum coronijerum Luo, Aitchison and Wang n. sp. Plate 1, figures 1-5

Etymology: Latin coronifer crowned, refemng to the crown-like patagial tissue.

Diagnosis: Test with five extra-triangular rods. At least seven pairs of closed caveal ribs all rejoin the plane of the skeletal frame and form a spherical cavea in the center of the test. Patagial vane well-developed. Patagial tissue originates from the b-i joint, forming a crown-like structure extending along the i.v.

Material: Common in samples 96-wx1129 and 96-wx1128 from the Tanhe Formation, Wuxingling, Nanning, Guangxi, SW China. Holotype: HKU-WXL128c-46 (pl. 1, fig. 3). Paratypes: HKU-WXL129a-19 (pl. 1, fig. 1) and HKU-WXL128c-23 (pl. 1, fig. 5).

H. Luo, J . C. Aitchison and Y. Wang.Devonian radiolariansfrom the Tanhe Formation, Nanning, Guangxi, southwest China

Measurements: Based on 14 specimens. a.a.: 87-129pm (holotype: 123pm); a.t.: 38-94pm (holotype: 48ym); a.p.: 96-105ym (holotype: 90pm); b.d.: 81-131ym (holotype: 104ym);b.t.: 45-67pm; i.d.: 79-122ym (holotype: 122pm);i.t.: 44-68ym; i.v.: 81-128pm (holotype: 91pm); diameter o f the spherical cavea 72-147ym (holotype: 73ym). Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation),Nanning, Guangxi so far as is known (this study). Ceratoikiscum planistellare Foreman 1963 Plate 1 , figures 7-9 Ceratoikiscumplanistellare FOREMAN 1963,p. 290, pl. 8, fig. 6;pl. 9, fig. 8.-NAZAROV 1975,~.99,pl. 11,figs.13-16;pl.12,figs.10-14. - AITCHISON 1993, p. 110, pl. 4, figs. 1-5.

Description: Test with five sturdy extra-triangular rods, generally straight. Central skeletal frame not clear. A well-developed wide lamellar patagium connects the extra-triangular rods. Cavea formed o f more than two pairs o f simple ribs, not long enough to join the patagium beyond the intersector. Remarks: Although all specimens are not well preserved, the main characteristics o f this species, a flat star-shaped outline with wide lamellar patagium, can be observed. Material: Four specimens from the Tanhe Formation, Wuxingling, Nanning, Guangxi, SW China. Measurements: Based on four specimens; length o f extratriangular rods 30-70pm. Range and occurrence: Lower to Middle Devonian from the Tanhe Formation, Wuxingling, Nanning, Guangxi. Also known from the Upper Devonian (Frasnian)o f western Australia and the southern Urals as well as the Upper Devonian (Famennian) o f North America. Ceratoikiscum sp.

Plate 1, figure 6 Description: Test with six straight extra-triangularrods. A wide patagial vane developed, irregularly perforate and occasionally slightly spongy. Cavea formed o f two pairs o f short caveal ribs, generally not long enough to join the patagium. Measurements: Based on 4 specimens. a.a.: 112-127ym; a.t.: 46-64ym; a.p.: 106-117ym; b.d.: 82-89pm; b.t.: 51-56pm; b.v.: 105ym; i.d.: 97-130ym; i.t.: 50-60pm; i.v.: 71-103ym. Material: Rare specimens from the Tanhe Formation, Wuxingling, Nanning, Guangxi, SW China. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi so far as is known (this study). Suborder SPUMELLARIA Ehrenberg 1875 Family ENTACTINIIDAE Riedel 1967; emend. Nazarov and Onniston 1984 Genus Spongentactinia Nazarov 1975 Type species: Spongentactiniafungosa Nazarov 1975 Spongentactiniu sp.

Plate 3, figures 19, 20

Description: Two spherical shells with six (?) three-bladed spines. Outer shell large, spongy and very thick with inner shell small, latticed and thin. Five radial beams connect the two shells. No by-spines observed. Measurements: Based on measurement o f two specimens. Diameter o f the outer shell 182-230ym, diameter o f the inner shell 42ym, the thickness o f the outer shell 39ym, width o f the base o f the main spines 28-30pm. Material: Rare specimens from the Tanhe Formation, Wuxinglin Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Genus Stigmosphaerostylus Riist 1892; emend. Foreman 1963 Type Species: Stigmosphaerostylusnotabilis Rust 1892 Stigmosphaerostylus RUST 1892, p. 142. - AITCHISON and STRATFORD 1997,p. 380. AITCHISON et al. 1999,p. 148. Ellipsostinzga HINDE 1899, p. 5 1 . Entactinia FOREMAN 1963, p. 271. Diagnosis: Species with a single well-developed latticed or spongy shell (sometimes with external spongy material) and an internal six-rayed double spicule more delicate than the main spines (fromForeman 1963, p. 271). Remarks: Based on the reassessment o f the Hinde (1899)fauna, Aitchison and Stratford (1997) regarded the widely used name Entactinia introduced by Foreman as a junior synonym o f Stigmosphaerostylus (Rust 1892). The more precise diagnosis o f Foreman (1963) based on descriptions o f matrix-free specimens was adopted (Aitchison and Stratford 1997) as an appropriate emendment for this genus. Range and occurrence: Upper Paleozoic, worldwide. Stigmosphaerostyhs cubicus Luo, Aitchison and Wang n. sp. Plate 1 , figures 15-17

Diagnosis: Cube-like spongy shell with 6 stout three-bladed main spines, variable in size and rapidly tapering in distal part. One spine commonly more robust than others displaying minor to moderate torsion. Wall o f shell rather thick. Remarks: This species is distinguished from all other species o f Stigmosphaerostylusby its cube-like spongy shell. No by-spines observed. As the inner structure could not be examined further study o f this species is necessary. Etymology: Latin cubicus-a-um, refemng to the cube-like shell. Measurements: Based on measurement o f nine specimens. Diameter o f shell 132-175pm (holotype: 146pm, average 144pm), length o f the longest spines 124-182pm (holotype: 153pm, average 155 pm), width o f the base o f the main spines 35-60ym (holotype: 35-44pm), thickness o f the shell wall up to 30 pm. Material: Specimens abundant in the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Holotype: HKUWXL134-24 ( p l . 1 , fig. 16). Paratypes: HKU-WXL134b-62 (pl. 1, fig. 15) and HKU-WXL134b-64 (pl. 1 , fig. 17). Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation),Nanning, Guangxi, SW China so far as is known.


Stigmosphaerostylus hystricuosa (Aitchison) 1993 Plate 2, figures 16-18

Staurodruppa HINDE 1899 p. 5 1. Entactinosphaera FOREMAN 1963 p. 274.

Entactinia hystricuosa AITCHISON 1993, p. 113, pl. 5, figs. 1,2. Apophysisphaera hystricuosa WON 1997b, p. 375, pl. 2, figs. 1-6.

Diagnosis: Two or more well developed, spherical or sub-

Diagnosis: See Aitchison (1993, p. 112). Description: Test with one small spherical lattice shell with six

three-bladed main spines. Main spines massive, very long and deeply bladed from base to tip. Several tiers of short spinules evident on the main spines; each tier with three spinules, one developed on each blade of the main spine. Some short thorn-like by-spines observed. Pores of shell polygonal-oval, relatively large in size, about 8-12 per hemisphere. Internal spicule not clear.

spherical shells, and an internal, six-rayed double spicule. When secondary spines are present, the six spines developed from the rays of the internal spicule are generally considerably more robust and are easily recognizable as main spines. Up to six main spines may be present (from Aitchison and Stratford 1997, p. 374). Trilonche davidi (Hinde 1899); emend. Nazarov 1975 Plate 2, figures 7 , 8

tion, Nanning, Guangxi.

Staurolonche davidi HINDE 1899, p. 46, pl. 8, fig. 13.

Staurolonche tenella HINDE 1899, p. 47, pl. 8, fig. 14.

Staurolonche laterna HINDE 1899, p. 47, pl. 8, fig. 15.

Staurodruppa nucula HINDE 1899, p. 52, pl. 9, fig. 7.

Staurolonchidium obliquurn HINDE 1899, p. 47, pl. 8, fig. 17.

Entactinosphaera grandis NAZAROV 1975, p. 65, pl. 5, figs. 11, 12;pl.

7, figs. 1-4. Trilonche davidi, (HINDE) AITCHISON and STRATFORD 1997, p. 380, pl. 1, fig. 8; pl. 2, fig. 4. - AITCHISON et al. 1999,p. 144, pl. 3, figs. K and L; pl. 4, figs. C, D, F, G, K, L; pl. 5, figs. M and N; pl. 6, figs. C and S.

Measurements: Based on measurement of seven specimens. Di-

Description: Two concentric spherical lattice shells with 6

Remarks: Imperfectly preserved specimens are observed which are externally similar to specimens of Stigmosphaerostylus hystricuosa (Aitchison), figured as Entactinia hystricuosa by

Aitchison (1993, pl. 5, figs. 1, 2). Material: Common in the Tanhe Formation, Wuxingling Sec-

Range and occurrence: Lower to Middle Devonian from

three-bladed main spines. Main spines robust all about same size and shape. One main spine with slight torsion. Outer shell relatively thick. Pores of shell rounded-oval and of fairly uniform size. No by-spines observed. Structure of internal spicule not clear.

Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Also known from the Devonian of eastern NSW, Australia, and Canning Basin, Western Australia.

Remarks: Based on Aitchison and Stratford (1997), the detailed diagnosis of the synonymised taxon Entactinosphaera grandis

ameter of shell 48-71pm (average 59pm), length of the main spines up to 100-216pm, width of the base of the main spines 16-35pm.

?Stigmosphaerostylus sp. Plate 2, figure 19,20 Description: One (?) large spherical to ellipsoidal lattice shell

with six (7) three-bladed main spines. One main spine significantly large, long, robust and strongly three-bladed, and may exhibit slight torsion; others apparently smaller and shorter. No by-spines observed. Numerous pores of shell relatively coarse, polygonal to rounded. Internal structure not visible. Remarks: This species is superficially like Trilonche echinata

(Hinde 1899) emend. Aitchison and Stratford (1997), but differs from it by having a much larger shell with coarser pores and no by-spines. As the inner structure is not clear, the genus assignment of this morphotype remains questionable. Material: Abundant in the Tanhe Formation, Wuxinglin Sec-

tion, Nanning. Measurements: Based on measurement of five specimens. Di-

ameter of outer shell 189-213pm, length of the largest main spines 199-259pm, basal width of the largest main spine 33-37pm, diameter of the pores 9-29pm. Range and occurrence: Lower to Middle Devonian from

Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Genus Trilonche Hinde 1899; emend. Foreman 1963; emend. Aitchison and Stratford 1997 Type species: Trilonche vetusta Hinde 1899 Triloriche HINDE 1899p. 48. AITCHISON and STRATFORD 1997, p. 373. AITCHISON et al. 1999, p. 144.

(Nazarov 1975, p. 65) is accepted as an emendation of the diagnosis of this species. Measurements: Based on measurement of two specimens. Di-

ameter of the outer shell 125-131pm , of the inner shell 42pm; length of the main spines up to 114-120pm, width of the base of the main spines 32-42pm. diameter of pores 4-6pm. Material: Two specimens from the Tanhe Formation, Wuxing-

ling Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from

Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Also known from the Devonian of eastern Australia, North America and Russia. Trilonche echinata (Hinde 1899); emend. Aitchison and Stratford 1997 Plate 1, figures 13, 14 Heliosoma echinatum HINDE 1899, p. 50; pl. 9, fig. 1, 2. Heliosomaparonae, HINDE 1899, p. 50; pl. 9, fig. 3. Entactinosphaera palimbola, FOREMAN 1963, p. 277; pl. 2, fig. 7a-e; pl. 3, fig. 3a-d. -LI and WANG 1991, p. 400, pl. 1, figs. 11-14. Entactinosphaera echinata? FOREMAN 1963, p. 279; pl. 3, fig. 10;pl. 4, fig. 12a-b. Entactinosphaera assidera, NAZAROV 1975, p. 64; pl. V, figs. 6,7; pl. VI, figs. 6-8. - LI and WANG 1991, p. 399, pl. 1, figs. 17. Entactinosphaera echinata, NAZAROV 1975,p. 60;pl. 111, figs. 1-3;pl. IV, figs. 1-4. ~ntactinos~haera cf. E. assidera, AITCHISON 1993, p. 115;pl. 6, figs. 2, 6. Trilonche echinata, AITCHISON and STRATFORD 1997, p. 377-378, pl. 1, figs. 5,6,10.- AITCHISON et al. 1999 p.145, pl. 1, figs. G-K, N; pl. 2, figs. D, K, L, Q; pl. 3, figs. 0 and X; pl. 4, fig. S;pl. 5, figs. B-D, G, I; pl. 6, figs. A, I, L-N, P.

H . Luo, J. C. Aitchison and Y. Wang: Devonian radiolarians from the Tanhe Formation, Nanning, Guangsi, southwest Clrina

Diagnosis: Two concentric spherical lattice shells with approximately six three-bladed main spines of variable width and length. By-spines are common but are variable and typically thorn-like. One main spine is typically substantially longer and more robust than the others. An internal six-rayed spicule is present within the inner shell (from Aitchison and Stratford 1997 p. 377). Description: Two concentric spherical lattice shells with six three-bladed main spines. One long, robust main spine, strongly three-bladed, sometimes with slight torsion; other spines smaller and shorter. Rod-like by-spines often well developed. Numerous pores of outer shell small, rounded and of approximately uniform size. Remarks: For a detailed discussion of this species see Aitchison and Stratford (1997) and Aitchison et al. (1999). Measurements: Based on measurement of 10 specimens. Diameter of outer shell 103-136pm, diameter of the inner shell up to 45pm, length of the largest main spines 117-206pm, width of the base of the largest main spine 29-41pm, length of by-spines up to 20-32pm, diameter of pores 5-9pm. Material: Abundant specimens from the Tanhe Formation, Wuxinglin Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi, SW China (this study). Also known from Upper Devonian (Frasnian) of Hexian County and Yulin City, Guangxi, China as well as Australia, North America and Russia. Trilonche sp. cf. T. echinata (Hinde 1899) Plate 2, figures 9, 10

Description: Two spherical lattice shells with five (six?) three-bladed main spines. One main spine extremely large, robust, typically much longer than other four (five?) spines, tapering abruptly to a sharp tip. Four other main spines about equal in size, present on a plane nearly perpendicular to the largest main spine and disposed at approximately 90" to each other. Outer shell very thick but the inner shell thinner. No by-spines observed. Pores of shells relatively small, fairly uniform in size. Internal structure not clear. Remarks: This species is distinguished from the typical T. echinata (Hinde 1899; emend. Aitchison and Stratford 1997) by its largest main spine which is heavier and somewhat shorter, its thicker outer shell and the lack of by-spines. The described specimens are similar to Trilonche hindea (Hinde 1899) with four smaller main spines present on a plane nearly perpendicular to the largest main spine and disposed at approximately 90" to each other. Only one larger main spine is observed, however, and the nature of the opposing spine is unclear due to the state of preservation. Thus, the species assignment is tentative. Measurements: Based on measurement of 3 specimens. Diameter of outer shell 113-141pm, diameter of inner shell 44pm; length of the largest main spine 88-115pm, width of spine base 42-54pm: outer shell thickness 25pm; inner shell thickness 8pm; pore diameter 5-8pm. Material: 3 specimens from the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi.

Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Trilonche minax (Hinde 1899)emend. Aitchison and Stratford 1997 Plate 1, figures 10-12

Xiphosphaera minax HINDE 1899, p. 45; pl. 8, fig. 8.

Stylosphaera obtusa, HINDE 1899, p. 45; pl. 8, fig. 9.

StaurodruppapraelongaHINDE 1899, p. 51; pl. 9, fig. 6.

Staurodruppa?prolataFOREMAN 1963, p. 281; pl. 8, figs. la-b; pl. 9,

fig. 12. Entactinosphaera egindyensis NAZAROV 1975, p. 61; pl. 5, figs. 2,4; pl.6,figs. 1,5.-LIand WANG 1991,p. 399,pl. l,figs.3,4. Trilonche minax (Hinde) AITCHISON and STRATFORD 1997, pl. 1, figs. 3,4; pl. 2, figs. 3,5. - AITCHISON et al. 1999, p. 145, pl. 2, figs. H, I; pl. 5, figs. J, K. Description: Two spherical concentric lattice shells with 6 main spines. Outer shell with small pores but inner medullary shell relatively coarsely porous. Two polar main spines extremely, long, robust and strongly three-bladed. Four smaller main spines short and present on plane perpendicular to two polar main spines. No by-spines observed. Pores hexagonal-oval. Remarks: For a detailed discussion of this species see Aitchison and Stratford (1997) and Aitchison et al. (1999). Measurements: Based on measurement of 3 specimens. Diameter of the outer shell 76-123pm, length of polar main spines 76-175pm, width of the base of the polar main spines 29-48pm, length of short main spines 32-59pm. Material: 3 specimens from the Tanhe Formation, Wuxinglin Section; Guangxi, SW China. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi, SW China (this study). Also known from Upper Devonian (Frasnian) of Hexian County, Guangxi, China as well as Australia, North America and Russia. Trilonche nanningensis Luo, Aitchison and Wang n. sp. Plate 2, figures 11-15

Entactiniidae gen. et sp. indet, sp. C, STRATFORD and AITCHISON 1997, pi. 7, figs. A, B. Entactiniid gen. et sp. indet, AITCHISON et al. 1999, pi. 4, fig. P. Diagnosis: Two concentric spherical lattice shells with 5 (6?) three-bladed main spines. All main spines robust and about same size and shape. Along the three-bladed edge, each main spine with a distinctive flattened rounded projection between 1/2 to 2/3 distance from base of spine. Spines taper more abruptly beyond projection. Both outer and inner shells relatively thin. Few by-spines observed. Pores of outer and inner shells small, rounded and of fairly uniform size. Remarks: This new species has an external appearance similar to taxa described from the Middle Devonian Ceratoikiscum regalinodus assemblage from Gamilaroi terrane in eastern Australia, figured as Entactiniidae gen. et sp. indet, sp. C (pl. 7, figs. 1, 2) by Stratford and Aitchison (1997), and material figured as Entactiniid gen. et. sp. indet. (pl. 4, fig. P) by Aitchison et al. (1999). The common characteristic is that all main spines have a rounded projection, whereas their internal structures are not visible. The specimens examined in this study, however, exhibit two latticed shells indicating that they belong to the genus Trilonche Hinde. The detailed internal structure is not clear.


Etymology: From Nanning City, the capital of Guangxi, in which this species is found. Measurements: Based on measurement of five specimens. Diameter of outer shell 117-132pm (holotype: 132pm), diameter of inner shell 41pm, length of the main spines up to 66-11 1pm (holotype: 85-106pm), width of the base of the main spines 30-41pm (holotype: 30-32pm), diameter of pores 3-7pm, thickness of the outer shell 6pm. Material: Five specimens from the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Holotype: HKUWXL128c-10 (pl. 2, fig. 13); Paratypes: HKU-WXL129-23 (pl. 2, fig. 11) and HKU-WXL129a-65 (pl. 2, fig. 14). Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Middle Devonian of the Gamilaroi terrane in eastern Australia. Trilonche remosa Luo, Aitchison and Wang n. sp. Plate 3, figures 11-1 8

Enractinosphaer-asp. C. ISHIGA 1988, pl. 5, fig. 8. Entactiniid gen. et sp. indet. A, AITCHISON et al. 1999p. 152, pl. 3, fig. I; pl. 4, figs. J and N. Diagnosis: Spherical lattice shells with six three-bladed main spines. All main spines robust and about same size and shape disposed at approximately 90" to each other; proximal half of spine from base straight but develops torsion of between 1 to 1.25 rotations, coiling sinistrally when looking towards tip; spines taper rapidly near tip. By-spine not developed. Pores of outer and inner shells small, rounded and fairly uniform size. Remarks: This species differs from Trilonche xinpoensis n. sp. in having only six main spines which are disposed at approximately 90" to each other, by the torsion of spine which occurs only in the distal part rather than along the entire length of the spine, and by the less developed by-spines. Of described species it is most like Entactiniid gen. et sp. indet, A (pl. 4, fig. J) by Aitchison et al. (1999) of the Middle Devonian Ceratoikiscum regalinodus assemblage from Gamilaroi terrane in eastern Australia. The detailed internal structure is not clear. Etymology: From the Tanhe Formation, in which this species is found.

Diagnosis: Two concentric spherical lattice shells with six three-bladed main spines. Main spines massive, subequal in width and length, and deeply grooved; most main spines with a single tier of short rod-like spinules on distal portion disposed at 90" to each other on a plane perpendicular to the main spines; at least one main spine, without short spinules, tapers to a sharp point. Outer shell relatively thin with numerous small pores. Inner shells commonly poorly preserved due to recrystallization. Some short rod-like by-spines observed. Pores of the outer shell small polygonal-oval.

Measurements: Based on measurement of five specimens. Diameter of outer shell 93-120pm (holotype: 96pm, average 106pm), length of the main spines up to 64-107pm (holotype: 94-107pm), width of the base of the main spines 19-28pm (holotype: 24-25pm).

Remarks: This species is distinguished from all other species of Trilonche by four short spinules on distal section of the main spine. This form of main spine bears some resemblance to that of Stigmosphaerostylus gogoense (Aitchison 1993). S. gogoense, however, only has one latticed shell rather than two. Although Won (1997, p. 374) claimed that she had some Devonian specimens with two latticed shells and outer apophyses (= the external spinules), the same characteristics of T. remosa, no formal descriptions of this kind of taxon have been published. Details of the internal spicule are not clear.

Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Probable Middle to Upper Devonian in New England Orogen, eastern Australia.

Etymology: Latin ramosus-a-um, branching, referring to the branching main spines. Measurements: Based on measurement of 20 specimens. Diameter of outer shell 96-156pm (holotype: 113pm, average: 118pm), diameter of inner shell 36-40pm, maximum length of the main spines 92-169pm (holotype: 118pm), width of the base of the main spines 27-41pm (holotype: 35-36pm), thickness of the outer shell 7-lOpm, diameter of pores 2-8pm. Material: Specimens are common in the Tanhe Formation, Wuxinglin, Nanning, Guangxi, SW China. Holotype: HKUWXL128c-32 (pl. 3, fig. 14). Paratypes: HKU-WXL128a-52 (pl. 3, fig. 11) and HKU-WXL129-09 (pl. 3, fig. 16). Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi, southwest China so far as is known. Trilonche tanheensis Luo, Aitchison and Wang n. sp. Plate 3, figures 6-9

Entactinia sp. AITCHISON 1988, p. 54, fig. 3S

Material: Five specimens from the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Holotype: HKUWXL128c-14 (pl. 3, fig. 8); Paratype: HKU-WXL129c-23 (pl. 3, fig. 6) and HKU-WXL128c-94 (pl. 3, fig.7)

Trilonche vetusta Hinde 1899;emend. Aitchison and Stratford 1997 Plate 1, figures 18, 19

Trilonche vetusra HINDE 1899,p. 48, pl. 8, figs. 18, 19. - AITCHISON and STRATFORD 1997,pl. 1,figs. l,2; pl. 2, figs. 1,2. - AITCHISON et al. 1999, p. 145, pl. 2, figs. J, R; pl. 3, fig. E; pl. 5, figs. F, H; pl. 6, figs. K, 0, Q. Trilonche pirtmani HINDE 1899, p. 48, pl. 8, figs. 20,21. Theodiscus hastatus HINDE 1899, p. 53, pl. 9, fig. 9. ?Triactiscus lanceola HINDE 1899, p. 53, pl. 9, fig. 11. Entactinosphaer-aaitpaiensis NAZAROV 1973,p. 697, pl. 1, figs 4-5. NAZAROV 1975, p. 63, pl. 5, figs. 1, 3, 5; pl. 6, figs. 2-4. - LI and WANG 1991, p. 399, pl. 1, figs. 8-10. Description: Two spherical concentric lattice shells with six main spines. Two strongly three-bladed polar main spines present on one axis; these spines extremely large, longer and more robust than other four main spines. Four smaller main spines short and may or may not be present on a plane perpendicular to the two polar main spines. Few by-spines observed. Pores of outer shell are hexagonal-oval. Remarks: For a detailed discussion of this species see Aitchison and Stratford (1997) and Aitchison et al. (1999). Measurements: Based on measurement of eleven specimens. Diameter of the outer shell 99-165pm, length of polar main

H. Luo, J. C. Aitchison and Y. Wang: Devonian radiolarians from the Tanhe Formation, Nanning, Guangxi, southwest China

spines 110-254pm, width of the base of the two polar main spines 41-75pm, length of short main spines 23-36pm. Material: Specimens are common in the Tanhe Formation, Wuxinglin Section, Nanning, Guangxi, SW China. Range and occurrence: Lower to Upper Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi, Guangxi, SW China (this study). Also known from the Upper Devonian (Frasnian) of Hexian County, Guangxi, China as well as Australia and Russia. Trilonche xinpoensis Luo, Aitchison and Wang n. sp. Plate 3, figures 1-5, 10

Diagnosis: Two concentric spherical lattice shells with 7-8 three-bladed main spines. All main spines subequal and of moderate length. Most exhibit sinistral or dextral torsion of between 1to 1.25 rotations from base to tip. Radial beams continuous with main spines, three-bladed with no torsion between shells. Numerous needle-like by-spines well developed on surface of outer shell. Outer shell and inner shell relatively thin. Pores of outer and inner shell small, rounded and fairly uniform in size Remarks: This species differs from Trilonche tanheensis n. sp. in having 7-8 main spines, by the torsion of spine which affects the entire length of the spine, and by the development of by-spines. Details of structure of internal spicule not clear. Etymology: From Xinpo village, where this species is found. Measurements: Based on measurement of eight specimens. Diameter of outer shell 111-163pm (holotype: 159pm, average 146pm), diameter of inner shell 51-55pm (holotype: 55pm); length of the main spines up to 64-106pm (holotype: 102-106pm), width of the base of the main spines 23-35 pm (holotype: 24-31pm). Diameter of the radial beams 12-18 pm; thickness of outer shell 9-17pm (holotype: 17pm), thickness of inner shell 6-7pm (holotype: 6pm); pore diameter 3-9pm (holotype: 3-4pm).

Material: Abundant in the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Holotype: HKU-WXL128c-51 (pl. 3, fig. 10); Paratypes: HKU-WXL128a-58 (pl. 3, fig. 4) and HKU-WXL129a-57 (pl. 3, fig. 5). Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi, SW China (this study). ?Trilonche sp.

Plate 2, figures 1-3

Description: Spherical lattice shell with 6 slender three-bladed main spines. Main spines heavy, very long and only slightly variable in width from base to tip. Numerous by-spines well developed on shell surface, generally needle-like and circular in cross section, and rarely three-bladed at base. Pores of shell small, rounded-oval and uniform in size. Details of the structure of the internal spicule not clear. Remarks: Specimens examined are externally similar to the morphotype described by Foreman (1963) as Entactinosphera echinata? (Hinde) (p. 279, pl. 3, fig. 10, pl. 4, fig. 12a-b., This form is characterized by the greater length of the slender main spines and numerous by-spines. Because the internal structure is not visible, the generic assignment here remains questionable. Measurements: Based on measurement of 7 specimens. Diameter of outer shell 113-162pm, length of the main spines up to 221-265pm, basal width 24-30pm; length of the long by-spines up to 48-60pm, diameter 12-20pm; pore diameter 6-10pm. Material: Abundant in the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). Genus Helioentactinia Nazarov 1975 Type species: Helioentactinia polyacanthina Foreman 1963 Helioentactinia NAZAROV 1975,p. 88

PLATE 1

Radiolarians from the Tanhe Formation, Wuxiangling section, Nanning, Guangxi.

All figures are scanning electron micrographs. All scale bars=lOOym.

1-5 Ceratoikiscum coroniferum n. sp. 1, paratype, HKU-WXL129a 19.2, HKU-WXL128a-34. 3, holotype, H K U - ~ ~ ~ 1 2 8 c - 44,6HKU.

WXL128a-13.5, paratype, HKU-WXL128c-23.

1@12 Trilonche minax (Hinde). 10, HKU-WXL134-04. 11, HKU-WXL134b-42. 12, HKU-WXL134b-43. 13-14 Trilonche echinata (Hinde). 13,

HKU-WXL128a-59. 14, HKU-WXL128c-77.

6 Ceratoikiscum sp. HKU-WXL134b-01.

7-9 Ceratoikiscum planistellare Foreman. 7, HKU-WXL128a-27.8, HKU-WXL128c-37.9, HKU-WXL128a-25.

1517 Stigmosphaerostylus cubicus n.sp. 15, paratype, HKU-WXL134b-62. 16, holotype, HKUWXL134-24. 17, paratype, HKU-WXL134b-64. 18-19 Trilonche vetusta Hinde. 18, HKU-WXL134b-44. 19, HKU-WXL134b-39.

H . Luo, J . C . Aitchison and Y. Wang

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W . Luo, J. C. Aitc,hisonand Y. Wang: Devonian radiolarians from the Tanhe Formation, Nanning, Guangxi, southwe.\t China

Radiobisphaera WON 1997a. p. 344 Diagnosis: Two spherical latticed shells with numerous external spines forming from ends of either multi-ray spicule or polyhedron located eccentrically or in the centre of internal shell (from Nazarov 1975, p. 88). Helioentactinia sp. cf. H. multa (Won 1997a) Plate 2, figure 6 Description: Two concentric spherical lattice shells with one very strong main spine and numerous thorn-like by-spines extended from radial beams. Outer shell quite thin with small rounded-oval pores. Inner shell with coarser pores than outer shell possessing many radial beams that arise from its surface. Internal structure not clear. Remarks: Specimens examined have an external appearance similar to Helioentactinia multa (Won) (Won 1997a, p. 352, pl. 3, figs 17-19) from the Upper Devonian Gogo Formation in Western Australia, but differ in having a smaller outer shell and short by-spines. Measurements: Based on measurement of 3 specimens. Diameter of outer shell 99-135pm, diameter of inner shell 42-46pm, length of the main spine up to 96pm, width of the base of the main spines 24-35pm, thickness of the outer shell 67-lOpm, thickness of the inner shell 7pm. Material: 3 specimens from the Tanhe Formation, Wuxiangling Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study).

Suborder INCERTAE SEDIS ~ ~PALAEOSCENIDIIDAE ~ i l ~~ i ~ (1967); d ~ l emend. Holdsworth (1977); emend. Goodbody (1982); emend. Goodbody (1986) Subfamily PALAEOSCENIDIINAE Riedel (1967); emend. Goodbody (1986) Type genus: Palaeoscenidium Deflandre 1953 Genus Palaeoscenidium Deflandre (1953); emend. Goodbody

(1986)

Type species: Palaeoscenidium cladophorum Deflandre (1953)

Diagnosis: For diagnosis see Goodbody (1986, p. 150). Palaeoscenidium cladophorum Deflandre 1953 Plate 2, figures 23-25 Palaeosrenidiunz cladophorunz DEFLANDRE 1953, p. 408, text-fig. 308.-DEFLANDRE 1960,p.214,pl. 1,fig. 21. -FOREMAN 1963,p. 302, pl. 8, fig. 10; pl. 9, fig. 6. - HOLDSWORTH 1973, p. 128, pl. 1, fig. 19. - NAZAROV 1975,p. 96, pl. 13, figs. 4-5; pl. 14, figs. 5-6. NAZAROV et al. 1982 p. 172,figs. 5D-F. -NAZAROV and ORMISTON 1983,p. 465, pl. 2, figs. 6,7. - ISHIGAet al. 1987, pp. 299-300, pl. 1, figs. 1-10. - LI and WANG 1991, p. 401, pl. 2, figs. 18-19. AITCHISON 1993, p. 121, pl. 1, figs. 15-17, 19, pl. 2, 17, 20. KIESSLING and TRAGELEHN 1994, pp. 240-241, pl. 6, figs. 5-7, 9-10. -STRATFORD and AITCHISON 1997,pl. 2-0. -WANG 1997, p. 157, pl. 1, fig. 6. - SASHIDA et al. 1998, pl. 19, fig. 8. 11-14. AITCHISON et al. 1999, p. 153, pl. 5, fig, 0;pl. 6, figs. F, J. Description: Four subequal apical spines and four rod-like straight or curved tapering basal spines arising at approximately 45" from short median bar; one spine may be considerably longer than other spines. Some short spinules developed at right angles to basal spines and disposed irregularly along them, below a small tent-like lamellar shell.

PLATE 2


All figures are scanning electron micrographs. All scale bars=lOOym.

1-3 ?Trilonche sp. 1, HKU-WXL134-43. 2, HKU-

WXL134b-67.3, HKU-WXL134b-68.

4-5 Trilonche echinata (Hinde). 4, HKU-WXL134b-36. 5, HKU-WXL 128a-63. 6 Helioentactinia sp. cf. H. multa (Won). HKU-

WXL128c-16.

7-8 Trilonche davidi (Hinde). 7, HKU-WXL129c-29. 8, HKU-WXL134-47. 9-10 Trilonche sp. cf. T. echinata (Hinde). 9, HKUWXL128-06. 10, HKU-WXL128c-13.

11-15 Trilonche nanningensis n. sp. 11, paratype, HKU-WXL129-23. 12, HKU-WXL128c-03.13, holotype, HKU-WXL128c-10. 14, paratype, HKU-WXL129a-65, 15, HKU-WXL 129~-04. 1618 Stigmosphaerostylus hystricuosa (Aitchison). 16, HKU-WXL128a-45. 17, HKU-WXL128a-44. 18, HKU-WXL128c-43. 1920 Stigmosphaerostylus sp. 19, HKU-WXL134b-30. 20, HKU-WXL134-40. 21-22 Magnisphaera sp. 21, HKU-WXL128c-85.22, HKU-WXL 129a-45.

2325 Palaeoscenidium cladophorum Deflandre. 23, HKU-WXL128a-42.24, HKU-WXL134b-24.25, HKU-WXL134b-13.


micropaleonrology,vol. 48, supplement no. 1,2002

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123

H. Luo, J. C .Aitchison and Y. Wang: Devonian radiolariansji-on1 the Tanhe Formation, Nanning. Guangsi, soutlzu~estChina

Remarks: P. cladophorum has fairly wide intraspecific variation in the same structural framework (Nazarov and Ormiston 1983; Ishiga et al. 1987; Sashida et al. 1998 and Aitchison et al. 1999). Specimens are commonly fragmentary and poorly preserved. Two forms within this species can be discriminated from our material: type one with long and thick spicules (pl. 8, fig. 21 and pl. 17, figs. 1,4) and type two with shorter and thinner spicules (pl. 3, figs. 3-5 and pl. 8, figs. 22-25). Measurements: Based on measurement of 12 specimens. Apical and basal spines are often broken and the maximum length is given. Length of basal spines 76-230pm, diameter 11-35pm; length of PA spines 11-70pm, diameter 10-15pm; height of tent-like shell up to 60pm; length of spinules up to 29pm. Material: Common in the Tanhe Formation, Wuxinglin Section, Guangxi. Range and occurrence: Lower-Upper Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi SW China (this study). Also known from the Middle Devonian to Lower Carboniferous of Europe, North America, USSR, Middle Asia, Malaysia, South China, Japan and Australia. Family INCERTAE SEDIS Genus Magnisphaera Won 1997a Tjpe species: Magnisphaera gigantea Won 1997a Magnisphera sp.

Plate 2, figures 2 1, 22

Description: Two spherical to subspherical lattice shells. Outer

shell large, relatively coarse, and loosely latticed. Pores vary in

shape and size, rounded to polygonal. Some mesh bars protrude

outward like by-spines. Inner shell relatively small and irregu-

larly latticed. Inner spicules and outer spines not clear.

Remarks: Most specimens examined are incomplete. But the

two latticed shells, an unusually large shell size and an ex-

tremely loose outer shell wall show the diagnostic features of

the genus Magnisphaera Won (1997a p. 366-367).

Measurements: Based on 6 specimens. Diameter of outer shell 170-236pm (average 195pm), diameter of the inner shell 50pm, diameter of mesh bar 7-8pm. Material: Six specimens from the Tanhe Formation, Wuxingling Section, Nanning, Guangxi. Range and occurrence: Lower to Middle Devonian from Wuxiangling (Tanhe Formation), Nanning, Guangxi (this study). ACKNOWLEDGMENTS This project has been supported by funding from The University of Hong Kong Committee on Research and Conference Grants in a research grant awarded to Dr. Jonathan Aitchison, a research student support fund of The University of Hong Kong and a Research Fund of Laboratory of Paleaeobiology and Stratigraphy, Chinese Academy of Sciences (No. 013 111) awarded to Hui Luo. We thank Dr. Andreas Braun for his helpful review and comments on the manuscript. REFERENCES AITCHISON, J. C., 1988. Radiolaria from the southern part of the New England Orogen. In: Kleeman J. D., Ed., New England Orogen Tectonics and Metallogenesis, 49-60. University of New England, Armidale. , 1993. Late Devonian (Frasnian) Radiolaria of the Canning Basin, Western Australia. Palaeontographica Abteilung A, 228: 105-128.

AITCHISON, J. C., and STRATFORD, J. M. C., 1997. Middle Devonian (Givetian) Radiolaria from eastern New South Wales, Australia: areassessment of the Hinde (1899) fauna. Neues Jahrbuch fiir Geologie und Palaontologie Abhandlungen, 203: 369-390. AITCHISON, J. C., DAVIS A. M., STRATFORD J. M. and SPILLER F. C. P., 1999. Lower and Middle Devonian radiolarian biozonation of the Gamilaroi terrane New England Orogen, eastern Australia. Micropaleontology, 45: 138-162. DEFLANDRE, G., 1953. Radiolaries fossiles. In: Grass P. P., Ed., Trait6 de Zoologie. 1 pt. 2: 389-436. Paris, Masson et Cie.

PLATE 3


All figures are scanning electron micrographs. All scale bars=lOOpm.

1-5,10 Trilonche xinpoensis n. sp. 1, HKU-WXL128a-58. 2, HKU-WXL128a-57.3, HKU-WXL128d-18.4, paratype, HKU-WXL128a-58. 5, paratype, HKUWXL129a-57. 10, holotype, HKU-WXL128c-51. 6 9 Trilonche tanheensis n. sp. 6, paratype, HKUWXL129c-23.7, paratype, HKU-WXL128c-94.8, holotype, HKU-WXL128c-14.9, HKUWXL129c-22,

11-18 Trilonche remosa n. sp. 11, paratype, HKUWXL128a-52. 12, HKU-WXL128a-48. 13, HKUWXL128c-15. 14, holotype, HKU-WXL128c-32. 15, HKU-WXL128c-11. 16, paratype, HKUWXL129-09. 17, HKU-WXL128c-44. 18, HKUWXL129-54. 1920 Spongentactinia sp. 19, HKU-WXL129a-66. 20, HKU-WXL129-72.


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, 1960. A propos du developpement des recherches sur les radiolares fossiles. Revue de Micropaltontologie, 2: 212-21 8.

EHRENBERG C. G., 1838, ~ b e die r Bildung der Kreidefelsen und des Kreidemergel durch unsichtbare Organismen. Abhandlungen Konigliche Akademie der Wissenschaften zu Berlin, Jahre 1838: 59-147. ---, 1875. Fortsetzung der mikrogeologischen Studien als Gesammt-Uebersicht der mikroskopischen Palaontologie gleichartig analysirter Gebirgsarten der Erde, mit specieller Riiksicht auf den Polycystinen-Merge1 von Barbados. Konigliche Akademie der Wissenschaften zu Berlin, Abhandlungen, Jahre 1875:l-225.

FENG Q. L. and LIU, B. P., 1992: ANew Radiolarian genus of Early Devonian from western Yunnan. Chinese Sciences (B). 5: 549-553. FOREMAN, H. P., 1963. Upper Devonian Radiolaria from the Huron member of the Ohio shale. Micropaleontology, 9: 267-304. GOODBODY, Q. H., 1982. Silurian Radiolaria from the Cape Phillips Formation, Canadian Arctic Archipelago. Proceedings Third North American Paleontological Convention, August, 1982, 1: 21 1-216. , 1986. Wenlock Palaeoscenidiidae and Entactiniidae (Radiolaria) from the Cape Phillips Formation of the Canadian Arctic Archipelago. Micropaleontology, 32: 129-157.

HINDE, G. J., 1899. On the Radiolaria in the Devonian Rocks of New South Wales. Quarterly Journal of the Geological Society of London, 55: 38-64. HOLDSWORTH, B. K., 1969. Namurian Radiolaria of the genus Ceratoikiscurn from Staffordshire and Derbyshire, England. Micropaleontology, 15: 221-229. ,1973. The Radiolaria of the Baltalimani Formation, Lower Carboniferous, Istanbul: Paleozoic of Istanbul. Ege universitesi. Fen fakultesi, Kitaplar Serisi, Jeol., 40: 117-134.

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