Devonian radiolarian faunas from South China - BioOne

Devonian radiolarian faunas from South China 1

2

Yujing Wang1, Jonathan C. Aitchison2, and Hui Luo1

Institute of Geology and Palaeontology, Academia Sinica, Nanjing, China Department of Earth Sciences, University of Hong Kong, Pok Fu Lam Road, Hong Kong, China email: [email protected]

Abstract: Devonian radiolarians from four new sections at Shaijingpo, Lila, Bazhai, and Shiti Reservoir in southern China are described. Faunas from these localities are assigned to eight families, 15 genera and 30 species, with the introduction of one new species Trilonche parapalimbola n. sp. Wang. The stratigraphic significance of the Middle Devonian Eoalbaillella lilaensis fauna, morphotypic variation within the genus Helenifore and the abundance and diversity of the Upper Devonian Holoeciscus foremanae fauna are discussed.

INTRODUCTION

LOCAL STRATIGRAPHY

The distribution of Devonian radiolarians in South China is shown in text-figure 1. They are mainly found in Ali, Huiku, Nanya, Lila, Taierbu, Shaijingpo and Changyuchun districts in western Yunnan Province, Bazhai and Huohong regions in Guizhou Province, Bancheng, Wuxinglin, Yangti, Yulin, Hexiang, Badu districts in Guangxi Province, and Xintian district in Hunan Province. Most faunas occur in oceanic cherts or platform sedimentary lithofacies. Some faunas occur in limestone.

Shaijingpo Section

Pioneering studies of Devonian radiolarians in South China began in the early 1980’s. Sheng Jinzhang and Wang Yujing first described some radiolarians collected from the Middle Devonian Qizhiqiao Formation in Xintian County, Hunan Province (Sheng and Wang 1982). Li (1986) reported some Upper Devonian radiolarians from the Ali area, West Yunnan Province. Li and Wang (1991) described an Upper Devonian (Frasnian) radiolarian fauna from the east and southeast of Guangxi Province. Feng and Liu (1992, 1993) reported a new radiolarian genus Eoalbaillella from west Yunnan Province. Wu et al. (1994), as well as Wang, (1994), reported an Upper Devonian (Famennian) Holoeciscus foremanae fauna from Xiaodong and Bancheng areas in Qingzhou, Guangxi Province. In recent years, Wang et al. (1998) recorded a Frasnian Helenifore laticlavium (=H. robustum) radiolarian fauna from the same region. Wang et al. (2000) reported on radiolarian faunas from cherty rocks in west Yunnan and Luo et al. (2002) report Givetian radiolarians from the Tanhe Formation near Nanning in Guangxi Province. Since 1996, Devonian radiolarian-bearing cherty samples have been collected systematically from the Yangti, Wuxingling and Bancheng areas in Guangxi, the Bazhai and Huohong areas in Guizhou Province, and the Lila, Huiku, Shaijingpo and Changyuchun areas in Yunnan Province. Four Devonian radiolarian faunas, Eoalbaillella lilaensis (Middle Devonian D2), Helenifore laticlavium (Frasnian D31), Helenifore robustum (Frasnian D31), and Holoeciscus foremanae (Famennian D32) faunas have been found. Text-figure 2 shows the local occurrences of these faunas on a lithostratigraphic column.

This section is located near Shaijingpo village, Xianyun County, West Yunnan Province, and is in the immediate vicinity of the road leading to Wujiachun village. A new railroad from Kuanming to Dali now cuts through the section. A stratigraphic section, about 165 m thick, belongs to the Changyuchun Formation (Middle-Upper Devonian). It consists of mainly thinly bedded siliceous rocks; mudstones and shales intruded by various thin diabase dikes and is well exposed in this section. The uppermost and lowermost parts of this formation are covered by Quaternary sediments. The upper part of this formation is about 75m and contains thick gray yellow or thin purple ribbon-bedded cherts. It contains a Helenifore robustum fauna including Palaeoscenidium cladophorum Deflandre, Stigmosphaerastylus variospina (Won), Spongentactinella cf. corynacantha Nazarov and Ormiston. The lower part of this formation is a 90 m thick section of black thinly ribbon-bedded siliceous rocks bearing an Eoalbaillella lilaensis Feng and Liu fauna that includes Polyentactinia miopora Wang, Stigmosphaerastylus cf. dissora (Nazarov), Stigmosphaerastylus sp. A, Spongentactinia shaijingpoensis Wang, Trilonche inusitata (Foreman), Trilonche parapalimbola Wang (n. sp.), Trilonche sp. A, Spongentactinella cf. corynacantha Nazarov and Ormiston and Helioentactinia yunnaensis Wang. Lila Section

This section is located on the south side of No. 38 highway near a Highway Maintenance Base at Lila village, in Lanchan County, West Yunnan Province. Sandstones interbedded with muddy shales and a few thin siliceous rocks are assigned to the Lila Formation which is exposed on this section. These rocks rest conformably on the Lalei Formation, which contains a Lower Devonian Monograptus fauna. Thinly bedded siliceous rocks in the lower part of Lila Formation bear an Eoalbaillella lilaensis fauna including Stigmosphaerastylus spp. Thinly bedded siliceous rocks in the upper part of Lila Formation contain a Helenifore robustum fauna including Trilonche davidi (Hinde), Trilonche minax (Hinde), Stigmosphaerastylus sp. B, and Stigmosphaerastylus cf. dimidiata Nazarov.

micropaleontology, vol. 49, no. 2, pp. 127-145, text-figures 1-2, plates 1-5, 2003

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Yujing Wang, Jonathan C. Aitchison, and Hui Luo: Devonian radiolarian faunas From South China

Bazhai Section

This section is located on a highway at Bazhai village, in southeast Ziyun County, southwest Guizhou Province. The Xiangshudong Formation, a sequence mainly consisting of thinly bedded siliceous rocks and muddy shales, is exposed in this section. This formation is 60m thick and lies conformably on the Middle Devonian Huohong Formation. It conformably underlies the Upper Devonian Daihua Formation. Radiolaria occurring in these thinly bedded siliceous rocks, belong to the Helenifore robustum fauna and include Palaeoscenidium cladophorum Deflandre, Stigmosphaerastylus variospina (Won), S. spp., Trilonche minax (Hinde), T. davidi (Hinde), and Astroentactinia sp. Shiti Reservoir Section

This section is located on the east highway of Shiti reservoir in Bancheng town, Qingzhou, southeast Guangxi Province. The exposed strata are composed mainly of cherts and siliceous mudstones and are named the Shiti Reservoir Formation. This unit was formerly called the Shiti Formation (Wang et al. 1998) but as this name has been pre-empted, we now use the term Shiti Reservoir Formation. The formation lies conformably on the Middle Devonian Xiaodong Group (grey black mudstones) and conformably underlies the Lower Carboniferous Shijia Formation. A Helenifore robustum fauna occurs in thinly bedded siliceous rocks in the lower part of this formation. It includes H. robustum (Boundy-Sanders and Murchey), Ceratoikiscus sp., Spongentactinella cf. corynacantha Nazarov and Ormiston. An abundant Holoeciscus foremanae fauna occurs in the upper thin siliceous rocks, and includes Archocyrtium spp., Astroentactinia sp., Trilonche spp., and Stigmosphaerastylus spp. AGE OF THE EOALBAILLELLA LILAENSIS FAUNA

The genus Eoalbaillella was proposed by Feng and Liu (1992, translation published in 1993). Type specimens were collected from the second layer of siliceous rock on the Lila Section, Lanchang, Yunnan Province. The graptolitic shale that underlies this siliceous rock yields the Lower Devonian graptolites Monograptus uniformis, M. aeguabilis, M. microdon and M. yukonensis. Therefore, Feng and Liu (1992) considered the Eoalbaillella fauna to be Lower Devonian. Later, Feng said (pers comm to Wang) that there is a 20 metre stratigraphic interval between the siliceous rock bearing the Eoalbaillella fauna and the shale bearing graptolites. Recently, we have found a Helenifore robustum fauna in the same section occurring at a higher level than the Eoalbaillella fauna. Aitchison (1990) first reported a radiolarian taxon Holoeciscus sp. coexisting with Palaeoscenidium cladophorum, Entactinia sp. and Entactinosphaera spp. in chert collected from the Djungati Terrane, of the New England Orogen, in eastern Australia. After reassessing the above specimen, Aitchison (1993a) considered this Holoeciscus sp. to be a new genus and species and described it as Paraholoeciscus bingaraensis. He also thought this taxon possibly represented a form transitional between Holoeciscus foremanae and Albaillella paradoxa estimating it to be Upper Devonian Famennian or lowermost Carboniferous. By comparing both their shape and skeletons, we think the genus Paraholoeciscus unquestionably is a junior synonym of Eoalbaillella. Aitchison communicated this to the radiolarian community via the Radfolks Bulletin board in 1995. In 1991, at the Sixth International Radiolaria Conference held in Florence, Italy, our Polish colleague M. Paskowski showed

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SEM photos of a taxon we interpret as Eoalbaillella lilaensis. Unfortunately this material remains unpublished. In 1991, we found that both the Eoalbaillella lilaensis fauna and Helenifore robustum fauna occur in thin siliceous beds in the Shaijingpo section, Xiangyuan County, West Yunnan (Li and Wang 1991). The bed containing the Helenifore robustum fauna occurs at a higher level than that of Eoalbaillella lilaensis fauna. In this area the Lower Devonian Lianhuaqu Formation is limestone with abundant conodonts, corals and tentaculitids. The Eoalbaillella-bearing Changyuchun Formation is composed of siliceous rock and muddy shale with scattered occurrences of Middle Devonian tentaculitids. The contact between the two formations is conformable. In the Huiku Section, Menglian, where Feng and Liu (1992) found the Eoalbaillella fauna, we have found that beds containing the Helenifore robustum fauna occur at higher stratigraphic levels than those with the Eoalbaillella fauna. In sections at Lila, Huiku and Shaijingpo, the Eoalbaillella lilaensis fauna always occurs between the Lower Devonian and Upper Devonian (Frasnian). Therefore, we consider it reasonable to assign a Middle Devonian age to the Eoalbaillella fauna (Wang et al 2000). Radiolarians co-existing with Eoalbaillella lilaensis are Spongentactinia shaijingpoensis, Trilonche inusitata, T. echinata, T. sp. A, Polyentactinia miopora, Stigmosphaerastylus cf. dissora, S. sp. A, Spongentactinella cf. corynacantha and Helioentactinia yunnaensis. Eoalbaillella lilaensis belongs to the family Albaillellidae, whereas the other taxa all belong to the family Entactiniidae and many of them are undescribed. Several of the long-ranging taxa present occur through to the Upper Devonian (Famennian). Eoalbaillella lilaensis fauna has now been found in Shaijingpo, Lila and Huiku in west Yunnan Province, China, the New England orogen of eastern Australia and Poland, and a similar taxon is also known from Alaska (Pat Whalen, pers. comm. to JA) but remains unpublished. MORPHOTYPIC VARIATIONS OF HELENIFORE

The genus Helenifore was introduced by Nazarov and Ormiston (1983) based on specimens collected from Gogo Formation (basal Frasnian, Upper Devonian), in the Canning Basin, West Australia. The type species H. laticlavium is characterized by a wide flat ring of platy fabric, usually with two spines (top spine and base spine) and an opening in one side of thin circular central ring. Aitchison (1993a) described a further species of Helenifore from the Canning Basin. Helenifore gogoense Aitchison is characterized by a narrow and thick platy ring. Ishiga (1988) reported specimens found from tuffaceous siltstone in the Hastings block, New England Fold Belt, east Australia, which he assigned to H. laticlavium. However, these specimens have a narrow and thick elliptical platy ring and are different from H. laticlavium as described from the Canning Basin, which possesses a thin and wide platy ring. We regard them as a new species somewhat similar to H. gogoense (Aitchison 1993a) but different to H. laticlavium. Thus, two morphotypes of Helenifore can be distinguished. One, H. laticlavium Nazarov and Ormiston, (Nazarov and Ormiston 1983; Aitchison 1993b) is characterized by thin and wide platy ring; the other, H. robustum, is characterized by a narrow and thick platy ring (Ishiga 1988; Aitchison 1993b; Sashida et al. 1993; Stratford and Aitchison 1997; Aitchison and Stratford 1997; Wang et al. 1998). Recently, Boundy-Sanders et al. (1999) named a new genus and species Durahelenifore robustum that is the same as this second Helenifore morphotye. Therefore, we adopt their species name

Micropaleontology, vol. 49, no. 2, 2003

TEXT-FIGURE 1 Map showing the distribution of Devonian Radiolarians in south China. 1 - East section of Shiti Reservoir, Guangxi; 2 - West section of Shiti Reservoir, Guangxi; 3 - Wuxiangling section, Nanning, Guangxi; 4 - Yangdi section, Guilin, Guangxi; 5 - Bazhai section, Ziyun, Guizhou; 6 - Shaijingpo section, Xiangyun, Yunnan; 7 - Lila section, Lancang, Yunnan; 8 - Taierbu section, Lancang, Yunnan; 9 - Huiku section, Menglian, Yunnan; 10 - Niuyunling section, Yulin, Guangxi; 11 - Etang section, Hexian, Guangxi; 12 - Nanya section, Menglian, Yunnan; 13 - Ali section, Lancang, Yunnan; 14 - Matangwuo section, Xintian, Hunan

“robustum” but reassign this taxon to the genus Helenifore. H. robustum is an upper Frasnian zonal fossil whereas H. laticlavium represents a lower Frasnian zonal species. Both morphotypes of Helenifore occur in the Bancheng area, Qingzhou, Guangxi, China, and H. robustum also occurs in Shaijingpo, Xiangyun; Lila and Taierbu, Lanchang; Huiku, Menglian in west Yunnan as well as Bazhai, Zhiyun, Guizhou. Generally only a few radiolarians such as Palaeoscenidium cladophorum, Stigmosphaerastylus variospina and Spongentactinella cf. S. corynacantha co-exist with H. robustum. Based on the co-occurrence of conodonts Ancyrodella alata, Ozarkodina immersa, Palmatolepis cf. P. gigas, P. sp., Neoprioniodus sp., Nothygnathellus sp. and Polygnathus sp., which are found in Bancheng area, Qingzhou, Guangxi and Taierbu, Lanchang, west Yunnan, Helenifore robustum-bearing rocks in China can be assigned to the upper Frasnian.

Schmidt-Effing (1988) illustrated a questionable Helenifore? cf. H. laticlavium (Pl. 1, fig. 1; pl. 3, fig. 4) found in the Famennian of the Frankenwald, Germany. Based on their photos we regard those specimens as most likely to be some incomplete fragments of Stigmosphaerastylus sp. ABUNDANCE AND DIVERSITY OF HOLOECISCUS FOREMANAE FAUNA

Since Holoeciscus foremanae was described by Cheng (1986), this taxon has been found in many areas worldwide. It has been treated as a type fossil of a standard radiolarian zone in the Famennian, Upper Devonian. The radiolarian fauna, which Cheng (1986) found co-occurring with Holoeciscus foremanae in phosphate nodules of the Lower Woodford Formation of the Upper Devonian, in Oklahoma, is abundant and diverse. It includes H. brevis, Ceratoikiscum simplum, C. astrum, C. delicatum, C. riedeli, C. extraordinarium, C. avimexpectans,

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Protoalbaillella pinetopensis, P. deflandrei, Pylentonema typica, P. triangulatum, Quadrapesus dumitricai, Q. unicus, Popofskyellium deflandrei, P. elemense, P. wonae, P. obesum, Cerarchocyrtium singularium, Mostlerium unicum, Huasha holdsworthi, H. densa, Cyrtisphaeractenium delicatum, C. spinosum, Archocyrtium formosum and A. typicum. This fauna includes 11 genera and 27 species belonging to 2 suborders, 2 superfamilies and 6 families. The fauna may also include spumellarians but none were described. Schwartzapfel and Holdsworth (1996) reported further details of faunas from phosphate nodules of the Woodford Formation, Oklahoma, USA. They found abundant well-preserved radiolarians (12 genera and 49 species) occurring together with Holoeciscus foremanae. The fauna they described includes Protoalbaillella pinetopensis, P. deflandrei, P. formosa, Ceratoikiscum ardmorensis, C. astrum, C. avimexpectans, C. bujugum, C. chengi, C. herkommeri, C. labyrintheum, C. mirum, C. planistellatum, C. sandbergi, C. simplum, C. surculum, C. sp. A, C. quinquestellatum, Lapidopiscum transversum, Holoeciscus brevis, H. longus, H. renzae, Popofskyellum pulchrum, P. undulatum, P. hendricksi, P. turpiculum, P. wonae, Cyrtentactinia formosa, C. macrocephala, C. splendida, C. pessagnoi, Totollum deflandrei, T. hendricksi, T. blomei, T. undulatum, T. wonae, T. ziegleri, Kantollum pittsburgense, K. undulatum, Pylentonema mira, P. typica, P. robusta, Quadrapesus araneae, Q. bakeri, Q. conili, Q. (?) jimisonae, Q. sixi, Cyrtisphaeractenium shengi and Staurentactinia nazarovi In the Frankfurt/Main area, Germany, Holoeciscus foremanae, which is found in an Upper Devonian clast within Pleistocene gravel, co-occurs with Archocyrtium cf. delicatum, Astroentactinia biaciculata, Ceratoikiscum sp., Stigmosphaerastylus herculea, Trilonche? palimbola, Palaeoscenidium cladophorum and Polyentactinia? sp. (Braun 1990b). In the Frankenwald, Germany, Holoeciscus foremanae cooccurs with abundant radiolarians including 20 genera and 53 species such as Huasha magnifica, Ceratoikiscum bujugum, C. extraordinarium, C. sp., Holoeciscus brevis, H. elongatus, Archocyrtium cf. angulosum, A. cf. A. delicatum, A. diductum, A. dilapites, A. effingi, A. eupectum, A.(?) formosum, A. aff. A. formosum, A. ludicrum, A. cf. A. riedeli, Pylentonema cf. P. annulatum, P. cf. P. undulatum, P. cf. P. hindei, Astroentactinia biaciculata, A. multispinosa, A. radiata, A. digitosa, A. aff. A. paronae, A. stellata, Stigmosphaerastylus additiva, S. exilispina, S. herculea, S. spongites, S. tortispina, S.(?) pantotolima, S. variospina, S. vulgaris, Trilonche fredericki, T. aitpaiensis, T. symphypora, T. palimbola, T. riedeli, Polyentactinia aranea, P. craticulata, P.(?) perampla, P. sp., P. fenestrata, Tetrentactinia spinulosa, T. teuchestes, Triaenosphaera sicarius, T. hebes, T. sp. A, Stauroplegma sp. A, Staurostylus sp. A. Cerarchocyrtium sp., Cyrtisphaeractenium(?) fluegeli, C. sp., Deflandrellium sp., Robotium sp., Cyrtentactinia cf. C. formosa, Bissylentactinia arrhinia and Palaeoscenidium cladophorum (Kiessling and Tragelehn 1994). In the Oumonhao area of Utubulak (Hofeng grassland), Northern Xinjiang Autonomous Region, China, H. foremanae occurs in silicified limestones of the Gennaren Formation, and co-occurs with Palaeoscenidium cladophorum, Archocyrtium

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ormistoni, Trilonche sp. A, Stigmosphaerastylus sp. A and S. oumonhaoensis (Wang 1997). In Bancheng area, Qinzhou, Guangxi Province, SW China, H. foremanae also co-occurs with abundant radiolarian entactinids, and archocyrtids as well as the conodonts Palmatolepis triangularis, P. delicatula, P. glabra pectinata, P. glabra acuta and P. perlobata schindewolfi (identified by Prof Chenyuan Wang, Nanjing Institute of Palaeontology and Stratigraphy, Academia Sinica in pers com to Yujing Wang). These conodonts indicate that the H. foremanae fauna belongs to the lower to middle Famennian, Upper Devonian. In the western Hastings Block, of the southern New England Orogen, in eastern Australia Holoeciscus foremanae co-occurs with Popofskyellum cf. P. elemense, P. cf. P. dumitricai, Archocyrtium cf. A. delicatum, and Ceratoikiscum planistellare (Ishiga 1988). In the Anaiwan terrane, in the same area, H. foremanae occurs together with Ceratoikiscum sp., Archocyrtium spp., Cyrtentactinia spp. and Palaeoscenidium cladophorum Deflandre (Aitchison and Flood 1990; Aitchison 1993a). Between the late Frasnian and Famennian, a mass extinction affected the biota occurred in many areas of the world. This is the famous F-F event. The event resulted in the disappearance or decline of many invertebrate taxa such as stromatoporids, conodonts, ammonites, tentaculitids, ostracods, brachiopods, corals, and trilobites. About 60% of such taxa became extinct at the end of the Frasnian. However, in our sections from China the above-mentioned Famennian radiolarian fauna has a much higher total abundance and diversity than that of Frasnian from the same areas. This fact seems to indicate that the F-F event only affected shallow water reef-dwelling biota and part of the associated planktonic fauna to disappear. It does not appear to have affected the radiolarian fauna living in deeper water realms. In contrast, during the Famennian the radiolarian fauna developed and diversified. It is possible that the F-F mass extinction may have affected shallow water taxa to a greater extent than those that lived in deeper water. SYSTEMATIC PALEONTOLOGY

Class ACTINOPODA Subclass RADIOLARIA Suborder SPUMELLARIINA Ehrenberg 1875 Family ENTACTINIIDAE Riedel 1967; emend Nazarov and Ormiston 1984 Subfamily ASTROENTACTINIINAE Nazarov 1975 Genus Astroentactinia Nazarov 1975 Type species: Astroentactinia stellata Nazarov 1975 ?Astroentactinia stellata Nazarov 1975

Plate 3, figures 27, 28 Astroentactinia stellata NAZAROV 1975, p. 82-83, pl. 8, fig. 6, pl. 10, figs. 1-3. – NAZAROV and ORMISTON 1983, p. 459, pl. 1, figs. 8, 9. – AITCHISON 1993b, p. 118, pl. 6. – KIESSLING and TRAGELEHEN 1994, p. 235, pl. 4, fig. 7.

Description: Shell is small and spherical with a thin wall and rounded pores. Numerous main external spines, which are generally short with angular oval pores at their bases.


TEXT-FIGURE 2 Lithostratigraphic columns showing the relative positions of local occurrences of four Devonian radiolarian faunas: Eoalbaillella lilaensis (Middle Devonian D2), Helenifore laticlavium (Frasnian D31), Helenifore robustum (Frasnian D31), and Holoeciscus foremanae (Famennian D32) faunas found at Lila, Huiku, Shaijingpo and Changyuchun areas in Yunnan Province.

Material: Abundant in samples from the Gogo Formation, Upper Devonian, (lower Frasnian) Canning Basin, Western Australia. Range and Distribution: Upper Devonian (Frasnian and Famennian), West Australia, South Urals, Russian, West Yunnan and Guizhou , China (Frasnian), Frankenwald, Germany and West Yunnan, Guangxi, China (Famennian). Genus Helioentactinia Nazarov 1975 Type species: Helioentactinia nigra (Hinde 1899)

Helioentactinia yunnanensis Wang

Plate 2, figures 19-24) Helioentactinia yunnanensis WANG et al. 2000, p. 248, pl. 1, figs. 17-20.

Description: Two concentric spherical lattice shells with more than 20 short robust three-bladed main spines. Comparison: This species has an external appearance closely similar to that of Spongentactinella corynacantha Nazarov and Ormiston, but differ in having two lattice shells not one;

Plate 1, figures 20, 21

Range and Distribution: Middle Devonian (Givetian), West Yunnan, China.


Genus Spongentactinella Nazarov 1975 Type species: Spongentactinella veles (Foreman 1963)

Helioentactinia aster Aitchison

Spongentactinella corynacantha Nazarov and Ormiston


Plate 3, figures 23-26

Helioentactinia aster AITCHISON 1993b, p. 118, pl. 7, figs. 16.

Spongentactinella corynacantha NAZAROV and ORMISTON 1983, p. 460-461, pl. 1, figs. 1-2. Spongentactinella cf. corynacantha WANG et al. 2000, p. 248, pl. 1, figs. 21-23; pl.2, figs. 14-15.

Helioentactinia sp. A

Range and Distribution: Upper Devonian (Frasnian and Famennian), West Australia (Frasnian) and West Yunnan, Guangxi, China (Famennian).

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Range and Distribution: Upper Devonian (Frasnian), West Australia and West Yunnan, Guizhou, Guangxi, China. Spongentactinella veles (Foreman)

Plate 5, figure 6) Tetrentactinia veles FOREMAN 1963, p. 283-284; pl. 7, fig. 1. Spongentactinella veles (Foreman) NAZAROV 1975, p. 92, pl. 12, fig. 9. - NAZAROV AND ORMISTON 1983, p. 460, pl. 1, fig. 10.

Range and Distribution: Upper Devonian (Frasnian and Famennian), South Urals, Russia, West Australia (Frasnian), Ohio, USA, West Yunnan, China (Famennian). Subfamily ENTACTINIINAE Riedel 1967; emend. Nazarov 1975 Genus Stigmosphaerostylus Rüst 1892; emend. Foreman 1963

Germany (Famennian), Western Australia and NSW, Australia, Southern Urals, Russian, West Yunnan, China (Givetian and Frasnian). Stigmosphaerostylus variospina (Won)

Plate 4, figures 1-7 Palaeoxyphostylus variospina WON 1983, p. 156-157, pl. 8, figs. 1-4, 6-22. - NOBLE 1992, pl. 1, fig. 8. Entactinia variospina GOURMELON 1986, p.183-184, pl. 4, fig. 1. GOURMELON 1987, p. 49-50, pl. 3, figs. 6-11. - BRAUN 1989, p. 368, pl. 2, figs. 3-4; pl. 4, fig. 5. - BRAUN 1990a, pl. 1, fig. 4. BRAUN 1990b, p. 109-110, pl. 7, figs. 4-6. - KIESSLING and TRAGELEHN 1994, p. 236-237, pl. 4, figs. 23-24. – WANG et al. 1998, p. 365, pl. 1, figs. 6-7. Entactinia cf. variospina (Won) LI and WANG 1991, p. 399, pl. 1, figs. 16-17. Stigmosphaerostylus variospina WANG et al. 2000, p. 245, pl. 2, figs. 10-13; pl. 3, figs. 19-20.

Type Species: Stigmosphaerostylus notabilis (Rüst 1892) Stigmosphaerostylus RÜST 1892, p. 142 - AITCHISON and STRATFORD 1997, p. 380. Ellipsostigma HINDE 1899, p. 51. Entactinia FOREMAN 1963, p. 271.

Remarks: Foreman (1963) introduced a new genus Entactinia described from matrix-free specimens. This genus is a junior synonym of Stigmosphaerostylus Rüst 1892. The diagnosis of Foreman (1963) is more precise and it was adopted as an appropriate emendment by Aitchison and Stratford (1997).

Range and Distribution: Upper Devonian Famennian) to Lower Carboniferous, Worldwide.

(Frasnian-

Stigmosphaerostylum sp. A

Plate 2, figures 17, 18 Range and Distribution: Middle Devonian (Givetian), West Yunnan, China. Stigmosphaerostylus sp. B

Plate 5, figure 13

Range and occurrence: Present in Upper Paleozoic radiolarian-bearing strata worldwide.

Range and Distribution: Upper Devonian (Famennian), West Yunnan and Guangxi , China.

Stigmosphaerostylus herculea (Foreman)

Stigmosphaerostylus sp. C

Plate 5, figure 14 Entactinia herculea FOREMAN 1963, p. 271, pl. 1, figs. 3a-d. SCHMIDT-EFFING p. 34, pl. 1, fig. 9. - BRAUN 1990b, p. 11, pl. 2, figs. 1-4. - BRAUN et al. 1992, p. 169-170, figs. 2-8. - LI and WANG 1991, p. 398-399, pl. 1, fig. 12. - KIESSLING and TRAGELEHN 1994, p. 236, pl. 4, figs. 12, 21-22. - WANG 1997, p. 151-152, pl. 1, fig.4.

Range and Distribution: Upper Devonian (Famennian), Ohio, U.S.A., Frankenwald and Main gravels, Germany, Vosges, France, and West Yunnan, Guangxi, China. Stigmosphaerostylus pusilla (Hinde 1899); emend. Foreman 1963

Plate 2, figures 11-16 Staurosphaera pusilla HINDE 1899, p. 46, pl. 8, fig. 12. Entactinia? additiva FOREMAN 1963, p. 273, pl. 1, fig. 10; pl. 3, fig. 9. Stigmosphaerostylus pusilla (Hinde) - AITCHISON and STRATFORD 1997, p. 381.

Diagnosis: The diagnosis presented for the junior synonym Entactinia additiva by Foreman (1963, p. 273) is taken as an emended diagnosis for this species. One small spherical lattice shell with seven three-bladed spines, variable in width and length. The pores are sub-circular to sub-angular, variable in size and shape, about nine to twelve per half a circumference. Remarks: Specimens figured by Hinde (1899) are nearly identical to material figured by Foreman (1963) thus Entactinia additiva is considered the junior synonym. Other specimens later assigned by Nazarov to Entactinia additiva do not belong to this species (see discussion in Aitchison 1993b). Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian and Famennian) Ohio, U.S.A., Frankenwald,

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Plate 5, figures 20, 21 Range and Distribution: Upper Devonian (Famennian), Guangxi, China. Genus Trilonche Hinde 1899; emend. Foreman 1963; emend. Aitchison and Stratford 1997

Type Species: Trilonche vetusta Hinde Trilonche HINDE 1899, p. 48. Staurodruppa HINDE 1899, p. 51. Entactinosphaera FOREMAN 1963, p. 274. Trilonche AITCHISON and STRATFORD 1997 p. 373-374.

Diagnosis: Two or more, well developed, spherical or subspherical 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 differentiated and recognizable as main spines. Up to six main spines may be present. Remarks: The diagnosis of Hinde (1899) restricted the genus to specimens with three radial spines. In 1963 Foreman published a less restrictive, but more general diagnosis of a synonymous genus Entactinosphaera that encompassed material described as Trilonche by Hinde. Aitchison and Stratford (1997) placed Entactinosphaera in synonymy and adopted the definition of (Foreman 1963) as an appropriate emendation for the diagnosis of the senior synonym Trilonche. Although up to six (possibly more) main spines may be present it is common for not all spines to be developed to a similar degree. Where, in addition to the three bladed main spines, smaller “by-spines” are present these subordinate spines typically have a rod- or needle-like form.


Range and Distribution: Present in Upper Paleozoic radiolarian-bearing strata worldwide.

Range and Distribution: Middle (Givetian) and Upper (Frasnian-Famennian) Devonian, Worldwide.

Trilonche cf. T. cancellicula (Foreman)

Trilonche elegans Hinde 1899; emend. Foreman 1963

Plate 5, figure 25


cf. Entactinosphaera cancellicula FOREMAN 1963, p. 278, pl. 3, fig. b; pl. 4, figs. 1a-f.

Trilonche elegans HINDE 1899, pl. 8, fig. 22. - AITCHISON and STRATFORD 1997, p. 378, figs. 2-7, 3-6. Staurodruppa nana HINDE 1899, pl. 9, fig. 8. Entactinosphaera symphypora FOREMAN 1963, p. 277; pl. 2, figs. 5a-c; pl. 3, fig. 4. - KIESSLING and TRAGELEHN 1994, p. 238, pl. 5, fig. 7.

Range and Distribution: Upper Devonian (Famennian), Ohio, U.S.A., Guangxi, China. Trilonche davidi (Hinde 1899); emend. Nazarov 1975

Plate 1, figures 14, 15; Plate 3, figure 21 Staurolonche davidi HINDE 1899, p. 46; pl. 8, fig. 13. Staurolonche tenella HINDE 1899, pl. 8, fig. 14. Staurolonche laterna HINDE 1899, pl. 8, fig. 15. Staurodruppa nucula HINDE 1899, pl. 9, fig. 7. Staurolonchidium obliquum HINDE 1899, pl. 8, fig. 17. Entactinosphaera grandis NAZAROV 1975, p. 65; pl. 5, figs. 11, 12; pl. 7, figs. 1-4. Trilonche davidi AITCHISON and STRATFORD 1997, p. 380, figs. 2-8, 3-4. - WANG et al. 2000, p. 246-247, pl. 2, figs. 16-17.

Diagnosis: The detailed diagnosis of the synonymized taxon Entactinosphaera grandis (Nazarov 1975, p. 65) was accepted (Aitchison and Stratford 1997) as an emendation of the diagnosis of this species. Remarks: The test has two concentric spherical shells, robust main spines and rare, long, by-spines. The main spines are massive, approximately equal in size, and three-bladed. They have the same width along their entire length. The outer shell is thick, with rounded or rounded-oval pores. By-spines are rare, quite thin, rod-like and terminate bluntly. Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian), Eastern Australia,, Southern Urals, Russia, West Yunnan, China (Givetian and Frasnian). Trilonche echinata (Hinde 1899); emend. Aitchison and Stratford 1997

Plate 1, figures 16-19; Plate 5, figures 26-30) Heliosoma echinatum HINDE 1899, p. 50; pl. 9, figs. 1, 2. Heliosoma paronae HINDE 1899, p. 50; pl. 9, fig. 3. Entactinosphaera palimbola FOREMAN 1963, p. 277; pl. 2, figs. 7a-e; pl. 3, figs. 3a-d. - SCHMIDT-EFFING 1988, p. 34, pl. 1, figs. 5-6. BRAUN 1990b, p. 12, pl. 2, figs. 5-6. - KIESSLING and TRAGELEHN 1994, p. 237, pl. 5, figs. 2, 4-5. Entactinosphaera echinata? FOREMAN 1963, p. 279, pl. 3, fig. 10; pl. 4, figs. 12a-b. Entactinosphaera assidera NAZAROV 1975, p. 64, pl. V, figs. 6, 7; pl. VI, figs. 6-8. Entactinosphaera echinata NAZAROV 1975, p. 60, pl. III, figs. 1-3; pl. IV, figs. 1-4. Entactinosphaera cf. E. assidera LI and WANG 1991, p. 399, pl. 1, fig. 17. - AITCHISON 1993b, p. 115; pl. 6., figs. 2, 6. Trilonche echinata AITCHISON and STRATFORD 1997, p. 377-378, figs. 2-5, 2-6, 2-10. - WANG et al. 2000, p. 246, pl. 1, fig. 24; pl. 3, figs. 9-11, 26.

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. Remarks: See Aitchison and Stratford (1997) for a detailed discussion of this taxon.

Remarks: Aitchison and Stratford (1997) synonymized Entactinosphaera symphypora Foreman with the senior synonym Trilonche elegans but adopted the more detailed diagnosis of Foreman (1963, p. 277), who worked with matrix free material, as an appropriate emendation. Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian and Famennian), E-Australia (Frasnian), Ohio, USA, Frankenwald, Germany (Famennian), West Yunnan, China (Givetian). Trilonche minax (Hinde 1899); emend. Aitchison and Stratford 1997

Plate 2, figures 6-10; Plate 5, figures 15-19 Xiphosphaera minax HINDE 1899, p. 45, pl. 8, fig. 8. Stylosphaera obtusa HINDE 1899, p. 45, pl. 8, fig. 9. Staurodruppa praelonga HINDE 1899, p. 51, pl. 9, fig. 6. Staurodruppa? prolata FOREMAN 1963, p. 281, pl. 8, figs. 1a-b; pl. 9, fig. 12. Entactinosphaera egindyensis NAZAROV 1975, p. 61; pl. 5, figs. 2, 4; pl. 6, figs. 1-5 - LI and WANG 1991, pl. figs. 3-4. Trilonche minax AITCHISON and STRATFORD 1997, p. 377, figs. 2-3, 2-4; figs. 3-3, 3-5. - WANG et al., 2000, p. 247, pl. 2, fig. 24.

Diagnosis: Spherical radiolarian with two concentric shells. Relatively thick cortical shell with small pores. Inner medullary shell small and coarsely porous. Two very large, long, robust, strongly grooved, polar main spines. The width of the base of the two main spines is approximately one radius of the outer shell or greater. Four other small, commonly rodded, main spines may be present on a plane perpendicular to the two major heavy three bladed spines. Remarks: See Aitchison and Stratford (1997) for a detailed discussion of this taxon. Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian and Famennian), E-Australia, S-Urals, Russia (Frasnian), Guangxi and West Yunnan, China (Frasnian and Famennian) and Ohio, U.S.A. (Famennian). Trilonche parapalimbola Wang n. sp.

Plate 2, figures 1-5; Plate 5, figures 22-23 Entactinosphaera palimbola LI and WANG 1991, p. 400, pl. 1, figs. 11-14.

Diagnosis: Two concentric spherical lattice shells with six three-bladed main spines. Two main spines are robust but one of them is extremely large and massive. Four other main spines are smaller and about equal in size. No by-spines or by-spines not developed. Comparison: This species is distinguished from Trilonche echinata by the lack of by-spines or by undeveloped by-spines. Holotype: 96SHT179, pl. 5, fig. 22.

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Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian and Famennian), Guangxi, and West Yunnan, China (Givetian to Famennian). Trilonche tetractinia (Foreman)

figs. 2-12, 14. - KIESSLING and TRAGELEHN 1994, p. 237, pl. 4, fig. 25. not Stylosphaera vetusta HINDE 1899, p. 46; pl. 8, fig. 10. not Entactinosphaera vetusta NAZAROV 1975, p. 64; pl. 5, figs. 8-10; pl. 6, figs. 9-11.

Plate 5, figure 24 Entactinosphaera tetractinia FOREMAN 1963, p. 276, pl. 2, figs. 6a-b; pl. 6, figs. 2a-b.

Range and Distribution: Upper Devonian (Famennian), Ohio, U.S.A., Guangxi, China. Trilonche vetusta Hinde 1899; emend. Aitchison and Stratford 1997

Diagnosis: Thin cortical test with small lattice pores and secondary spines. Inner shell has small circular to polygonal pores. Three main spines that are moderately stout, typically sub-equal in length although the third may be somewhat smaller than the other two. Main spines are strongly three-bladed and connect through to the inner shell and are disposed at approximately 120° to each other.

Plate 1, figures 25-30 Trilonche vetusta HINDE 1899, p. 48; pl. 8, figs. 18, 19. - AITCHISON and STRATFORD 1997 p. 374-376, figs. 2-1, 2-2; figs. 3-1, 3-2. WANG et al. 2000, p. 247, pl. 3, figs. 25-26. Trilonche pittmani 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. Entactinosphaera aitpaiensis NAZAROV 1973, p. 697; pl. 1, figs. 4-5. NAZAROV 1975, p. 63, pl. 5, figs. 3, 5; pl. 6, figs. 2-4. - LI and WANG 1991, p.399, pl. 1, figs. 8-10. - BRAUN et al. 1992, p. 171,

Remarks: See Aitchison and Stratford (1997) for a detailed discussion of this taxon. Range and Distribution: Middle (Givetian) and Upper Devonian (Frasnian and Famennian), E-Australia, S-Urals, Russia, Guangxi and West Yunnan (Frasnian), Vosges, France and Frankenwald, Germany (Famennian), West Yunnan, China (Givetian).

Explanation of plates All figures are scanning electron micrographs and scale bars are 100µm. Specimens illustrated are deposited in Nanjing Institute of Geology and Palaeontology, Academia Sinica. Their locations collected, sample numbers, and ages are as follows: 1. Shaijingpo, Xianyun, Yunnan: 91SJP12, 91SJP 13 (Givetian); 91SJP 18, 91SJP 28 (Frasnian). 2. Lila, Lanchan, Yunnan: 97LL4 (upper Frasnian). 3. Taierbu, Yunnan: 91LL’88 (upper Frasnian). 4. Nanya, Yunnan: 97Ny2 (Famennian). 5. Bazhai, Ziyun, Guizhou: 98Ba12 (upper Frasnian). 6. Shiti Reservoir, Qingzhou, Guangxi: 96SHT159 (lower Frasnian); 96SHT174 (upper Frasnian); 96SHT179, 96SHT181, 96SHT182, 96SHT185, 96SHT219 (Famennian). 7. Shijia Reservoir, Qingzhou, Guangxi: 91J12 (Famennian).

PLATE 1 Middle Devonian D2 1-6 Eoalbaillella lilaensis Feng and Liu. 1, 2, 91SJP13; 3-6, 91SJP12. 7-13 Spongentactinia shaijingpoensis Wang. 7, 11, 91SJP13; 8-10, 12, 13, 91SJP12. 14,15 Trilonche davidi (Hinde). 14, 15, 91SJP13. 16-19 Trilonche echinata (Hinde). 16, 19, 91SJP13; 17, 18, 91SJP12.

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20-21 Helioentactinia sp. A. 20, 21, 91SJP12. 22,23 Trilonche elegans (Hinde). 22, 91SJP13; 23, 91SJP12. 24 Triaenosphaera sp. A, 91SJP12. 25-30 Trilonche vetusta Hinde. 25-30, 91SJP12.

Yujing Wang, Jonathan C. Aitchison, and Hui Luo

micropaleontology, vol. 49, no. 2, 2003

Plate 1

135


Trilonche sp. A

Plate 3, figures 19, 20 Range and Distribution: Upper Devonian (Frasnian), West Yunnan and Guizhou, China.

Montagne Noire, France (Tournaisian), Frankenwald, Germany, West Yunnan and Guangxi, China (FrasnianFamennian). Polyentactinia miopora Wang

Plate 2, figures. 25-30

Trilonche sp. B

Plate 3, figure 22

Polyentactinia miopora WANG et al. 2000, p. 248, pl. 1, figs. 13-15.

Range and Distribution: Upper Devonian (Frasnian), Guizhou, China.

Diagnosis: One sub-spherical lattice shell with six strong, approximately equal in size, three-bladed main spines. The pores are less and large, polygonal to rounded. By-spines not developed.

Genus Spongentactinia Nazarov 1975 Type species: Spongentactinia fungosa Nazarov 1975

Comparison: This species differs from Polyentactinia leptosphaera Foreman by the smaller shell and less pores.

Spongentactinia shaijingpoensis Wang

Plate 1, figure 7-13 Spongentactinia shaijingpoensis WANG et al. 2000, p. 247, pl. 1, figs. 8-12.

Range and Distribution: Middle Devonian (Givetian). West Yunnan, China.

Description: Two concentric spherical shells with six massive three-bladed main spines. The outer shell is spongy whereas the inner shell is latticed. The main spines are deeply bladed. Five of them possess a distinctive projection along the each bladed edge at 2/3 part of the way from the base to tip.

Family HAPLENTACTINIIDAE Nazarov 1980 Subfamily HAPLENTACTINIINAE Nazarov 1980 Genus Haplentactinia Foreman 1963 Type species: Haplentactinia rhinophyusa Foreman 1963

Comparison: This species has an external appearance and main spines closely similar to that of Stigmosphaerostylus profundiculcus Aitchison (1993b). It is distinguished from Stigmosphaerostylus profundiculcus by possessing two shells and by its spongy outer shell.

Haplentactinia rhinophyusa Foreman

Plate 3, figure 18 Haplentactinia rhinophyusa FOREMAN 1963, p. 270, pl. 1, fig. 2; pl. 3, fig. 7. - AITCHISON 1993b, p. 120, 121, pl. 5, figs. 12, 16. Haplentactinia cf. rhinophyusa WANG et al. 2000, p. 249, pl. 3, fig. 29


Range and Distribution: Upper Devonian (lower Frasnian), Guangxi, China.

Family POLYENTACTINIIDAE Nazarov 1975 Genus Polyentactinia Foreman 1963 Type species: Polyentactinia craticulata Foreman 1963

Genus Triaenosphaera Deflandre 1973 Type species: Triaenosphaera sicarius Deflandre

Polyentactinia aranea Gourmelon

Plate 5, figure 7

Triaenosphaera cf. T. sicarius Deflandre

Plate 5, figures 8-12 Polyentactinia aranea GOURMELON 1986, p. 185-186, pl. 4, fig. 5. GOURMELON 1987, p. 77-78, pl. 10, figs. 7-9. - BRAUN 1990b, p. 130-131, pl. 14, figs. 4-5, 10; pl. 15, figs. 13-15. - KIESSLING and TRAGELEHN 1994, p. 238, pl. 5, fig. 10.

Range and Distribution: Upper Devonian (Frasnian-Famennian) to Lower Carboniferous (Tournaisian), Pyrenees and

cf. Triaenosphaera sicarius DEFLANDRE 1960, pl. 1, fig. 10. DEFLANDRE 1973, p. 1150, pl. 2, figs. 3-4. - HOLDSWORTH et al. 1978, p. 781, figs.2a-c. - GOURMELON 1986, p. 186-187, pl. 2, fig. 1. - SCHMIDT-EFFING 1988, p. 35, pl. 3, fig. 5. - BRAUN 1990, p. 114-115, pl. 11, figs. 8-9. - KIESSLING and TRAGELEHN 1994, p. 239, pl. 5, figs. 21-22. - WANG et al. 2000, p. 249, pl. 2, fig. 31; pl. 3, fig. 27.

PLATE 2 Middle Devonian D2 1-5 Trilonche parapalimbola n. sp. Wang. 1, 2, 3, 5, 91SJP12; 4, 91SJP13. 6-10 Trilonche minax (Hinde). 6, 8, 91SJP12; 7, 9, 91SJP13; 10, 91SJP28. 11-16 Stigmosphaerostylus pusilla (Hinde). 11-14, 91SJP13; 15, 16, 91SJP12.

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17,18 Stigmosphaerostylus sp. A, 91SJP12. 19-24 Helioentactinia yunnanensis Wang. 19, 21-22, 24, 91SJP12; 20, 91SJP13; 23, 97Ny2. 25-30 Polyentactinia miopora Wang. 25-27, 30, 91SJP12; 28, 29, 91SJP13.



Plate 2

137


Range and Distribution: Upper Devonian (Famennian)-Lower Carboniferous (Visean), Montagne, Noire and Herault, France (Visean), Alaska, USA, Frankenwald, Germany, West Yunnan, China (Famennian). Triaenosphaera sp. A

Helenifore laticlavium NAZAROV and ORMISTON 1983, p. 464-466, pl. 2, figs. 8, 11, 13, 14. - AITCHISON 1993b, p. 112; pl. 2, figs. 11, 12.

Diagnosis: See (Nazarov and Ormiston 1983)

Plate 1, figure 24

Range and Distribution: Upper Devonian (lower Frasnian), west Australia, Guangxi, China.


Helenifore robustum (Boundy-Sanders and Murchey)

Suborder ALBAILLELLARIA Deflandre 1953; emend. Holdsworth 1969 Superfamily ALBAILLELLACEA Cheng 1986 Family CERATOIKISCIDAE Holdsworth 1969 Genus Ceratoikiscum Deflandre 1953 Type species: Ceratoikiscum avimexpectans Deflandre 1953

Helenifore laticlavium ISHIGA 1988, p. 82, pl. 1, figs. 1-4. - ISHIGA et al. 1988, fig. 2g. - STRATFORD and AITCHISON 1997, pl. 1, figs. C, D. - SASHIDA et al. 1993, fig. 4: 24-26. - WANG et al. 1998, pl. 1, figs. 4, 5. - AITCHISON et al. 1999, p. 158, pl. 1, fig. A; pl. 6, figs. G, H. Durahelenifore robustum BOUNDY-SANDERS and MURCHEY 1999, p. 66, pl. 1, figs. A-G.

Ceratoikiscum sp. A

Discussion: Test consists of a thicker, narrow ring with two main (basal and apical) spines. The ring has a circular-elliptical central opening and is strongly elongate. The surface of the ring is smooth without any small denticulate spinules. Apical main spine (i.d.) is short, sturdy and tapering. Basal main spine (i.v.) is rod-like, stout and moderately long. A short thorn-like (b.a.) spine is developed. No b.p. spine is present.

Plate 3, figure 29 Range and Distribution: Upper Devonian (Frasnian), Guangxi, China. Ceratoikiscum planistellare Foreman

Plate 3, figure 30


Ceratoikiscum planistellare FOREMAN 1963, p. 290, pl. 8, fig. 6; pl. 9, fig. 8. - NAZAROV 1975, p. 99, pl. 11, figs.13-16; pl. 12, figs. 10-14. AITCHISON 1993b, p. 110, pl. 4, figs.1-5. - NAZAROV and ORMISTON 1983, p. 461-462, pl. 2, figs. 3, 5. - 1988, p. 79-81, pl. 3, figs. 6-10. - SCHWARTZAPFEL and HOLDWORTH 1996, p. 115-116, pl. 7, figs. 1, 2, 6, 7, 14, 18, 19. - WANG et al. 2000, p. 243, pl. 2, fig. 19.

Remarks: This species differs from Helenifore laticlavium Nazarov and Ormiston (1983) in that it has a thicker, narrow ellipsoidal ring without any denticulate spinules.

Range and Distribution: Upper Devonian (Frasnian), Guangxi, China.

Family HOLOECISCIDAE Cheng 1986 Genus Holoeciscus Foreman 1963 Type species: Holoeciscus anceps Foreman 1963

Genus Helenifore Nazarov and Ormiston 1983 Type species: Helenifore laticlavium Nazarov and Ormiston 1983

Range and Distribution: Upper Devonian (upper Frasnian), worldwide.

Holoeciscus formanae Cheng

Plate 4, figures 8-12 Holoeciscus foremanae CHENG 1986, p. 91-92, pl. 5, figs. 4-6, 9-10, 14, 18. - AITCHISON 1988, figs. 4A, B. - ISHIGA 1988, p. 83-84, pl. 3, figs. 1-5. - ISHIGA et al. 1988, fig. 2c. - AITCHISON 1990, p. 179, pl.

Helenifore laticlavium Nazarov and Ormiston 1983


PLATE 3

Upper Devonian, Frasnian D31 1-5 Helenifore laticlavium Nazarov and Ormiston, 96SHT159.

22 Trilonche sp. B, 98Ba12.

6-13 Helenifore robustum (Boundy-Sanders and Murchey). 6, 8, 9, 96SHT174; 7, 97LL4; 10-12, 91LL’88; 13, 91SJP28.

23-26 Spongentactinella corynacantha Nazarov and Ormiston. 23, 24, 98Ba12; 25, 96SHT159; 26, 91SJP28.

14-17 Palaeoscenidium cladophorum Deflandre. 14, 91SJP28; 15, 16, 97LL4; 17, 97Ny2.

27,28 Astroentactinia stellata Nazarov. 27, 98Ba12; 28, 96SHT185.

18 Haplentactinia rhinophyusa Foreman, 96SHT159. 19,20 Trilonche sp. A. 19, 98Ba12; 20, 97LL4;

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21 Trilonche davidi (Hinde), 91SJP28.

29 Ceratoikiscum sp. A, 96SHT159. 30 Ceratoikiscum planistellare Foreman, 96SHT159.



Plate 3

139


6.5. - AITCHISON 1990, figs. 4a-b - BRAUN 1990b, p. 9-10, pl. 1, figs. 1-3. - WANG 1991, pl. 2, fig. 8. – AITCHISON and FLOOD 1992, figs. 3-2. - AITCHISON 1993a, pl. 1, fig. 4. - KIESSLING and TRAGELEHN 1994, p. 230, pl. 1, figs. 8-9, 11-12. - SPILLER and METCALFE 1995, fig. 3.1. - SCHWARTZAPFEL and HOLDSWORTH 1996, p. 125-126, pl. 20, figs. 1-4, 7-9. - WANG 1997, p. 158, pl. 1 fig. 15. - WANG et al. 1998, p. 365, pl. 1, figs. 1-3. WANG et al. 2000, p. 242-243, pl. 3, figs. 1-5.

Range and Distribution: Upper Devonian (Famennian), worldwide. Holoeciscus elongatus Kiessling and Tragelehn

Plate 4, figures 13, 14 Holoeciscus elongatus KIESSLING and TRAGELEHN 1994, p. 230, pl. 1, figs. 14-17. Holoeciscus longus SPILLER and METCALFE 1995, figs. 3.2-3.4. SCHWARTZAPFEL and HOLDSWORTH 1996; p. 126-127, pl. 12, figs. 3-6, 18-20.

Range and Distribution: Upper Devonian (Famennian), Oklahoma, North America, Frankenwald, Germany, Guangxi, China.

lower and middle Lower Devonian. However, Wang et al. (2000) consider that it is more reasonable to assign Eoalbaillella lilaensis fauna to the Middle Devonian than the Lower Devonian. Holotype: See Feng and Liu 1992, pl. 1, fig. 1. Range and Distribution: Middle Devonian (Givetian), West Yunnan, China, New England Orogen, Eastern Australia and Poland Suborder NASSELLARIINA Ehrenberg 1875 Superfamily CYRTOIDEA Haeckel 1862 Family POPOFSKYELLIDAE Deflandre 1964; emend. Cheng 1986 Subfamily POPOFSKYELLINAE Cheng 1986 Genus Popofskyellum Deflandre 1964; emend Cheng 1986 Type species: Popofskyellum palchrum Deflandre 1964 Popofskyellum cf. P. elemense Cheng

Plate 5, figures 1-5 cf. Popofskyellum elemense CHENG 1986, p. 156, pl. 3, figs. 1, 3, 14, 17, 23.

Genus Eoalbaillella Feng and Liu 1992

Type species: Eoalbaillella lilaensis Feng and Liu 1992 Diagnosis: “The test bilaterally symmetrical, composed of a lamellar shell and a triangular framework. The lamellar shell, supported by the triangular framework, imperforate, subcylindrical or conical, having numerous marginal spines around apertural margin. A-rod much shorter than i, b-rod forming an elongate triangular framework, a-rod and upper parts of I, b-rod covered by the shell; the lower parts of i, b-rod, without shell, like the shape of the letter X”(Feng and Liu 1993, p. 244)

Range and Distribution: Upper Devonian (Famennian), Oklahoma and Arkansas, North America, West Yunnan and Guangxi, China. Family ARCHOCYRTIIDAE Kozur and Mostler 1981; emend, Cheng 1986 Genus Archocyrtium Deflandre 1972; emend. Cheng 1986 Type species: Archocyrtium riedeli Deflandre 1972 Archocyrtium cf. A. effingi Kiessling and Tragelehn

Eoalbaillella lilaensis, Feng and Liu 1992


Plate 1, figures 1-6 Holoeciscus sp. AITCHISON 1990, p. 369, figs. 3N, O. Eoalbaillella lilaensis FENG and LIU 1992, p. 550-551, pl. 1, figs. 1-16. Paraholoeciscus bingaraensis AITCHISON 1993, p. 363, pl. 1, fig. 5.

cf. Archocyrtium effingi KIESSLING and TRAGELEHN 1994, p. 231, pl. 2, figs. 14-15.

Range and Distribution: Upper Devonian (Famennian), Frankenwald, Germany, West Yunnan, China.

Diagnosis: same as genus diagnosis (See Feng and Liu 1993, p. 244)

Archocyrtium ormistoni Cheng

Discussion: Feng and Liu first reported the Eoalbaillella lilaensis in Lila, Ximeng county, southwestern Yunnan as

Archocyrtium ormistoni CHENG 1986, p. 125-126, pl. 5, figs. 1-2, 4, 11, 21; pl. 6, figs. 2, 19. - WANG 1997, p. 157, pl. 1, figs. 7-8.


PLATE 4

Upper Devonian, Frasnian D31 and Famennian D32 1-7 Stigmosphaerostylus variospina (Won). 1-3, 5, 98Ba12; 4, 6, 7, 91SJP18. 8-12 Holoeciscus formanae Cheng. 8-9, 12, 96SHT185; 10, 97Ny2; 11, 96SHT179. 13,14 Holoeciscus elongatus Kiessling and Tragelehn, 96SHT181. 15-18 Archocyrtium ormistoni Cheng. 15, 96SHT182; 16, 96SHT181; 17, 96SHT219; 18, 91J12.

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19,27-31 Archocyrtium venustum Cheng. 19, 96SHT181; 27, 28, 96SHT219; 29-31, 96SHT182 20,25 Archocyrtium wonae Cheng, 96SHT185. 21-24 Archocyrtium cf. A. effingi Kiessling and Tragelehn. 21, 23, 24, 96SHT182; 22, 96SHT185. 26 Archocyrtium sp. A, 96SHT185. 32,33 Helioentactinia aster Aitchison. 32, 96SHT179; 33, 96SHT185.



Plate 4

141


Archocyrtium cf. ormistoni WANG et al. 2000, p. 244, pl. 3, figs. 13-14.

Range and Distribution: Upper Devonian (Famennian), Oklahoma, North America, Xinjiang, Yunnan and Guangxi, China. Archocyrtium venustum Cheng

Plate 4, figures 19, 27-31 Archocyrtium venustum CHENG 1986, p. 128, pl. 1, figs. 1-2, 6-7, 9-10; pl. 5, figs. 3, 6, 18, 26. 5

Range and Distribution: Upper Devonian (Famennian), Oklahoma, North America, Guangxi, China. Archocyrtium wonae Cheng

Plate 4, figures 20, 25 Archocyrtium wonae CHENG 1986, p. 128-129, pl. 5, fig. 8; pl. 6, fig. 4. – KIESSLING and TRAGELEHN 1994, p. 232, pl. 2, fig. 4 - WANG et al. 2000, p. 244, pl. 3, fig. 12.

Range and Distribution: Upper Devonian (Famennian), Oklahoma, North America, Frankenwald, Germany, Xinjiang, Yunnan and Guangxi, China. Archocyrtium sp. A

Plate 4, figure 26 Range and Distribution: Upper Devonian (Famennian), Guangxi, China. Incertae sedis

Family PALAEOSCENIDIIDAE Riedel 1967; emend. Holdsworth 1977; Goodbody 1982; Furutani 1983; Goodbody 1986 Genus Palaeoscenidium Deflandre 1953; emend. Goodbody 1986 Type species: Palaeoscenidium cladophorum Deflandre 1953 Palaeoscenidium cladophorum Deflandre group

Plate 4, figures 14-17 Palaeoscenidium cladophorum DEFLANDRE 1953, p. 408, 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. - SCHMIDT-EFFING 1988, p. 35, pl. 1, figs. 11-12. - ISHIGA et al. 1987, p. 299; pl. 1, figs. 1-10. ISHIGA 1988, p. 87-89, pl. 2, figs. 4, 6-8. - BRAUN 1990b, p. 17-18, pl. 1, figs. 4-5. - AITCHISON 1990, figs. 2E, 3M, I-J. - WANG 1991, pl. 2, fig. 15. - LI and WANG 1991, p. 401, pl. 2, figs. 18-19. AITCHISON and FLOOD 1992, figs. 3-7. - AITCHISON 1993b, p. 121; pl. 1, figs. 15-17, 19; pl. 2, figs. 17, 20. - KIESSLING and TRAGELEHN 1994, p. 240-241, pl. 6, figs. 5-7, 9-10. STRATFORD and AITCHISON 1997, pl. 2O. - WANG 1997, p. 157, pl. 1, fig. 6. - WANG et al. 2000, p. 249-250, pl. 2, figs. 7-9.

Range and Distribution: This is a cosmopolitan species group reported from Australia, Europe, Asia minor, North America, the former Soviet Union and Japan. Middle Devonian to Lower Carboniferous as far as is known. The biostratigraphic potential of Palaeosceniids remains to be established. ACKNOWLEDGMENTS

The authors wish to thank Ian Metcalfe for his helpful review of the manuscript. The University of Hong Kong is thanked for provision of SEM facilities used in the production of this paper. Work described in this paper was supported by grants to J. Aitchison from the Research Grants Council of the Hong Kong Special Administrative Region, China. REFERENCES AITCHISON, J.C., 1988. Late Paleozoic radiolarian ages from the Gwydir terrane, New England orogen, eastern Australia. Geology 16: 793-795. ———, 1990. Significance of Devonian-Carboniferous radiolarians from accretionary terranes of the New England Orogen, eastern Australia. Marine Micropaleontology, 15(3-4): 365-378. ———, 1993a. Albaillellaria from the New England orogen, Eastern NSW, Australia. Marine Micropaleontology, 21(4): 353-367. ———, 1993b. Late Devonian (Frasnian) Radiolaria of the Canning Basin, Western Australia. Palaeontographica, Abteilung A, 228: 105-128.

PLATE 5

Upper Devonian, Famennian D32

1-5 Popofskyellum cf. elemense Cheng. 1-4, 91J12; 5, 97Ny2. 6 Spongentactinella veles (Foreman), 97Ny2. 7 Triaenosphaera cf. sicarius Deflandre, 97Ny2 8-12 Polyentactinia aranea Gourmelon. 8, 96SHT179; 9-11, 96SHT181; 12, 91J12. 13 Stigmosphaerostylus sp. B, 96SHT179 14 Stigmosphaerostylus herculea (Foreman), 96SHT219

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15-19 Trilonche minax (Hinde). 15, 16, 19, 96SHT219; 17, 97Ny2; 18, 96SHT181. 20,21 Stigmosphaerostylus sp. C, 96SHT219 22,23 Trilonche parapalimbola n. sp. Wang. 22 (holotype), 96SHT179; 23, 96SHT185. 24 Trilonche tetractinia (Foreman), 96SHT181. 25 Trilonche cf. cancellicula (Foreman), 96SHT179. 26-30 Trilonche echinata (Hinde). 26, 27, 29, 96SHT219; 28, 91SJP28; 30, 91J12.



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Manuscript received August 21, 2002 Manuscript accepted March 25, 2003

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