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New Lopingian (Late ) rugosochonetid species from Sichuan, South China Monica J. Campi a; Guang R. Shi a a School of Ecology and Environment, Deakin University, Burwood, Australia

Online Publication Date: 01 January 2005

To cite this Article Campi, Monica J. and Shi, Guang R.(2005)'New Lopingian (Late Permian) rugosochonetid species from Sichuan, South China',Alcheringa: An Australasian Journal of Palaeontology,29:2,275 — 285 To link to this Article: DOI: 10.1080/03115510508619306 URL: http://dx.doi.org/10.1080/03115510508619306

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MONICA J. CAMPI and GUANG R. SHI

CAMP|, M.J. & SHI, G.R., 30.9.2005. New Lopingian (Late Permian) rugosochonetid species from Sichuan, South China. Alcheringa 29, 275-285. ISSN 0311 5518.

Two rugosochonetid species, Neochonetes (Huangichonetes) geniculatus sp. nov. and Neochonetes (Zhongyingia) linshuiensis sp. nov., are described from the Lopingian (Late Permian) of the Chuanmu section, Sichuan, South China. Ecological changes from the diverse upper Changhsingian palaeocommunity to the depauperate post- extinction brachiopod community are briefly discussed.

Monica J. Campi [[email protected]] & Guang R. Shi [[email protected]], School of Ecology and Environment, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood 3125, Victoria, Australia; received 13.6.2004, revised 19.11.2004.

Key words: , chonetids, ecology, Lopingian, Permian- boundary.

TWO NEW RUGOSOCHONETID species from The Feixianguan Formation at the Chuanmu the Changhsing and lowermost Feixianguan section is a thick sequence (>1 000 m) of fairly Formations at the Chuanmu section, Sichuan, monotonous thin-bedded marl and micrite. South China are described in this paper. The Brachiopods were only found at the very base of Chuanmu section is located in the Huaying this formation at the Chuanmu section. Early Mountains (Huayingshan) in eastern Sichuan, Triassic sequences are fairly uniform across South about 125 km northeast of Chongqing City (Fig. China (Peng et al. 2001), and were deposited Downloaded By: [Deakin University] At: 04:24 11 May 2009 1). Permian and Triassic sequences are well during a transgressional event and subsequent exposed along a road cutting at this section, highstand following a regression that occurred which is fairly continuous from the Cisuralian towards the end of the Permian (Chert et al. 1998). Liangshan Formation to the Late Triassic. This They are characterised initially by thin bedded work forms part of a larger study on the Permian mudstone and shale, followed by fairly thinly Brachiopoda from the Chuanmu section by MJC. bedded marly to argillaceous micritic facies, and are generally well laminated with little evidence Lithology and biostratigraphy of bioturbation. The new species were collected from the The Changxing Formation is the youngest upper half of the Changxing Formation and the Permian formation in South China, and is lowermost 20 cm of the overlying Feixianguan conformably overlain by Triassic marine sediments at the Chuanmu section. In this section Formation. The abundant and highly diverse and the surrounding area in Sichuan, the brachiopod fauna of the upper half of the Changxing Formation is dominated by cherty Changxing Formation yielded more than 70 limestone with subordinate argillaceous identifiable species (304 individuals) associated limestone and marl, representing a shallow with the two new species. The most common carbonate platform environment (Fig. 2). species was Haydenella kiangsiensis (Kayser, 0311/5518/2005/02275-11 $3.00 © AAP 1883), followed by Spinomarginifera lopingensis 276 MONICA J. CAMPI and G R. SHI ALCHERINGA

R ~Carbonifereua ~~---- LiangshanFormation

~~ okouFormation ~~ LongtanFormation mines ~ -_--~_~

Fig. 1A-B. A, Location map of the Chuanmu section in Sichuan. 1: mountains; 2: rivers, 3: provincial boundaries. B, Distribution of formations at the Chuanmu section. 1: gravel road; 2: river. (Kayser, 1883), Prelissorhynchia pseudoutah 20 cm (Bed 178a) of the overlying Feixianguan (Huang, 1933), Neochonetes (Zhongyingia) Formation (Fig. 2). linshuiensis sp. nov., Spinomarginifera alpha The brachiopod fauna in the Feixianguan Huang, 1932, Acosarina minuta (Abich, 1878), Formation at the Chuanmu section is found only Spinomarginifera chengyaoyenensis Huang, at the very base of this formation in one bed only 1932, Crurithyris speciosa Wang, 1955, (Bed 178a, CC- 11). It consists entirely of Permian Haydenella nasuta Zeng, 1993, Linshuichonetes holdover taxa which did not immediately succumb flatus (Shen & Archbold, 2002), Cathaysia to the end-Permian extinction. The brachiopod orbicularis Xu & Grant, 1994, Neochonetes fauna from Bed 178a (CC-11) contains eight (Huangichonetes) cursothornia (Xu & Grant, species along with an unidentifiable specimen Downloaded By: [Deakin University] At: 04:24 11 May 2009 1994), Neochonetes (Huangichonetes) sub- belonging to the Rugosochonetidae. The species strophomenoides (Huang, 1932), Spirigerella? are Cathaysia orbicularis, Spinomarginifera guizhouensis (Liao, 1980b), Huatangia sulcatifera lopingensis, Crurithyris sp., Linshuichonetes (Liao & Meng, 1986) and Orthotetina frechi flatus, Neochonetes (Huangichonetes) sub- (Huang, 1933), which represented just over 60% strophomenoides, Acosarina minuta, Neo- of the collections; the other 60 species make up chonetes (Huangichonetes) geniculatus sp. nov. the remainder of the species diversity. and Spirigerella? sp., in order of decreasing Neochonetes (Zhongyingia) tinshuiensis sp. abundance. nov. is a relatively common species in the upper As conodont work is yet to be fully part of the Changxing Formation at this section, completed on this section, the Permian-Triassic with twelve specimens collected from seven boundary was primarily located through the use horizons in the upper 110 m of the Changxing of lithostratigraphy, biostratigraphy and Formation (Fig. 2). The second new species, eventostratigraphy (Peng & Tong 1999, Peng et Neochonetes (Huangichonetes) geniculatus sp. al. 2001) mainly through correlation of the clay nov. appears to be much rarer, with only two beds immediately above and below the boundary specimens collected from two horizons, one bed. From correlation with the PTBSS (Permian- within 20 cm (Bed 174) of the top of the Triassic Boundary Stratigraphic Set) of Peng et Changxing Formation and the other in the basal al. (2001) the Permian-Triassic boundary has ALCHERINGA NEW PERMIAN BRACHIOPODS FROM SOUTH CHINA 277

In U n n I I 139 181 I U I I U I I $ 0 nil n ! n u n I I U I I~1 180 ~ -- 181 138 iI~ ~ I ' ~.,~. ~-i]: ¢ 178b i,.i.~i.~l~::.~ ,:-'~ ;"~ ;"-'~i:i _.., A U- [ 17~" ~JO ~ 'II]I'll .... IIl"'.i] 21II :In :II ~ i

t37 174 i~_ 17z

173 175 136 172 ,° (~ 174 171

135 170

134 169 d) _ 168

133 167 165 164 132 163 162 131 161 3 rn general 160 ~ scale 159 o 158 130 0 C0 157 o ¢o 1. lJ~ l in =. E E ~55 L~. 154 2_ 2. ~ 129 153 152 3 gN ¢.~ 151 0 128 150

149 4. 148

147 146 127 Downloaded By: [Deakin University] At: 04:24 11 May 2009 145 144

143 142 126 7.

141 125 8. 124

140 123 9.

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Fig. 2. Stratigraphic column of the Changhsing (top 110 m only) and Feixianguan Formations at the Chuanmu section. 1, limestone; 2, argillaceous limestone; 3, cherty limestone; 4, cherty bioclastie limestone; 5, mudstone; 6, calcareous mudstone; 7, claystone; 8, Neochonetes (Zhongyingia) linshuiensis sp. nov.; 9, Neoehonetes (Huangichonetes) geniculatus sp. nov.; 10, other brachiopods. The arrow marked 'A' at top right indicates the Permian-Triassic biostratigraphic boundary, which occurs between Beds 178a and b, about 20 cm above the top of the Changxing Formation. The arrow marked 'B' indicates the lithostratigraphic boundary between the Changxing and Feixianguan Formations. 278 MONICA J. CAMPI and G. R. SHI ALCHERINGA

been placed by us between Beds 178a and 178b, China (pers. obs.). The S-H-NPalaeocommunity which appear to be the equivalents of Beds 27a/ was associated with an abundant invertebrate fauna b and 27c/d of Meishan D section respectively dominated by crinoids, with bivalves, corals, (Peng et al. 2001, Yin et al. 1996). Placement of ammonoids and trilobites present in smaller the boundary at this point locates the numbers (Fig. 3A). brachiopods of Bed 178a in the very last bed of The depositional environment for this the Permian, although they are above the community is interpreted as a back reef or eventostratigraphic boundary marked by the clay lagoonal environment, below wave base, rather beds (Beds 25 and 26 at the Meishan section), than in a reef or bioherm deposit. Reefs and and the main extinction horizon, which is in Bed bioherm build-ups were quite common in the 176 at the Chuanmu section [this extinction event Changhsingian in South China, and were usually was discussed in detail in Campi (2003)]. This characterised by abundant corals and calcareous also means that the basal 20 cm of the Feixianguan algae, along with reef-building brachiopod genera Formation is Permian, as the lithostratigraphic boundary between the Feixianguan and Changxing Formations was placed by us at the top of Bed 174, in accordance with Yin et al. (2001). Peng et al. (2001) reported the conodont Hindeodusparvus from Bed 178 (their Bed 5) at the Chuanmu section; however, the exact identity of the specimen is somewhat questionable according to Peng Yuanqiao (pers. comm. 2003).

Ecology These new species occur within the Spinomarginifera - Haydenella - Neochonetes (S - H- N) Palaeocommunity recognised in the upper part (top 40 m) of the Changxing Formation at the Chuanmu section (Campi 2003). Downloaded By: [Deakin University] At: 04:24 11 May 2009 Neochonetes (Zhongyingia) linshuiensis sp. nov. is a prominent member of this community, whereas N. (Huangichonetes) geniculatus sp. nov. played a very minor role, as it first appeared at the very top (top 20 cm) of the Changxing Formation. The S- H- N Palaeocommunity was / very diverse (> 70 species) and was dominated by small, spiny productids and chonetids, in addition to genera such as Prelissorhynchia and Acosarina. It was generally found in micrite to Fig. 3. Palaeocommunity reconstructions. A, typical cherty and bioclastic limestones and is interpreted members of the Spinomarginifera - Haydenella - Neochonetes Palaeocommunity from the upper part of as a relatively quiet environmental setting with a the Changxing Formation; a, crinoids; b, ammonoid; c, soft, muddy lime substrate. There is abundant Prelissorhynchia; d, Acosarina; e, Sehuchertella; f, pyrite in some beds at the Chuanmu section, chonetids; g, Spirigerella?; h, trilobite; i, Crurithyris; j, which possibly indicates periodic presence of productids (e.g. Spinomarginifera and Haydenella); k, bioturbation. B, holdover taxa found in Bed 178a, reducing conditions, and this is common in the Feixianguan Formation; a, Crurithyris; b, Cathaysia; c, limestone facies in the Changhsingian in South Acosarina; d, chonetids; e, Spinomarginifera. ALCHERINGA NEW PERMIAN BRACHIOPODS FROM SOUTH CHINA 279

such as Richthofenia, Perigeyerella, Leptodus of dysaerobic benthic conditions. However, in and Laterispina (Liao & Meng 1986, Shen & He the Chuanmu section pyrite crystals are 1994a). Corals are rare at the Chuanmu section commonly disseminated through the limestones and, although both Leptodus and PerigeyereUa in the upper part of the Changxing Formation, are present, they are not abundant. with little other evidence of dysaerobic Neochonetes (Huangichonetes) geniculatus conditions, as the abundant and diverse sp. nov. was only found at the very top of the brachiopod fauna otherwise indicates a relatively Changhsingian (Fig. 2) at the Chuanmu section, tolerable environment. and it appears to be one of the few species that One obvious environmental consequence of survived for a short period after the end-Permian the facies change from the Changhsing to the mass extinction. The effect of this extinction event Feixianguan Formations, aside from any on the Permian hrachiopod fauna at the Chuanmu background phenomena operating on a larger section is discussed in greater detail in Campi scale to cause the global signature of this mass (2003). Other species that also were present in extinction, is the sudden influx of argillaceous the 'holdover' fauna in the aftermath of the end- material, indicated by the argillaceous micrite of Permian mass extinction at this section were Bed 178 compared to the cleaner limestones of Cathaysia orbicularis, Spinomarginifera the Changxing Formation. This influx may have lopingensis, Linshuichonetes flatus, Neo- affected the feeding efficiency and respiration of chonetes (Huangichonetes) substropho- many of the brachiopod species adapted to the menoides, Acosarina minuta, Crurithyris sp. and less turbid conditions prevalent in the Changxing Spirigerella? sp. (Fig. 3B). Formation. There is some support for this These species were all small and the argument as the 'holdover' species that were 'holdover' fauna at this section appears to be present in the basal Feixianguan Formation were quite similar in general composition to that found all small and relatively thin-shelled. These in other areas of South China immediately above morphological adaptations have been suggested the mass extinction horizon. For example, Xu & to be a consequence of lowered oxygen levels Grant (1994) recognised a suite of Permian-type (Allison et al. 1995). Small size alone may be a survivors in the lower Griesbachian at several result of benthic environmental conditions only sites in South China that included Acosarina, being adequate for life for short periods of time,

Downloaded By: [Deakin University] At: 04:24 11 May 2009 Araxathyris, Crurithyris and Prelissorhynchia so not allowing organisms to attain their full in addition to several species of chonetids. The growth potential (Ferguson 1985). composition of the 'holdover' fauna in bed 178a Several chonetid species were also present at the Chuanmu section is quite similar in in the 'holdover' group at the Chuanmu section, composition to the Transitional Bed Lingula and are very common at this level across South fuyuanensis - Crurithyris flabelliformis China (Xu & Grant 1994). This eurytopic group is Assemblage Zone of Shen & He (1994b). often a dominant element of brachiopod There appears to have been an environmental communities which experienced less than ideal change associated with the mass extinction, as environmental conditions (Simanauskas & the Changxing Formation is dominated by cherty Cisterna 2000, Shen & Archbold 2002, Campi & and bioclastic limestone indicative of a relatively Shi 2002). clear and normal marine environment, whereas It is interesting that characteristic disaster taxa the basal Feixianguan Formation is dominated such as Lingula have not been found in the by marl and thin-bedded micrite. This facies Permian-Triassic boundary sequence at the change has been recognised from elsewhere in Chuanmu section. This genus is extremely Sichuan (Wignall & Hallam 1996, Ezaki etal. 2003), abundant above the boundary in other areas of and has been interpreted as resulting from a Sichuan and across South China (Xu & Grant transgressive event and associated development 1994, He & Shi 1996), where it often fonlas high 280 MONICA J. CAMPI and G. R. SHI ALCHERINGA

abundance, low-diversity shell beds with bivalve nov. (Figs 4A, G, I- O, Fig. 5) genera such as Claraia. At the Chuanmu section Triassic bivalves, such as Towaptera, first appear Etymology. From the geniculate longitudinal in Bed 180e, about 32 cm above the holdover profile of the ventral valve of this species. Permian fauna in Bed 178a, or 50 cm above the lithological boundary between the Changhsing Material. The holotype is a ventral valve external and Feixianguan Formations. Claraia appears mould and corresponding ventral internal mould, about 70 cm above the boundary (Peng et al. NMV P309550a-b. The paratype is a fragmentary 2001) and Lingula may appear above this level, ventral valve, NMV P309551. as it has been found in the Lower Triassic of nearby sections (Xu & Grant 1994), but not yet at Diagnosis. Small subquadrate Neochonetes the Chuanmu section. From our unpublished (Huangichonetes) with highly convex ventral observations in areas around Chongqing (about valve, very small ears, shallow median sulcus. 125 km from the Chuanmu section), Lingula can appear some metres above the boundary between Description. Small for genus, subquadrate to the Changhsing and Feixianguan Formations. trapezoidal in outline, moderately transverse; strongly inflated; ears very small, flat and triangular, well demarcated from body; hingeline Systematic palaeontology straight, forming maximum width, bearing hinge The material studied for this paper is deposited spines; shallow sulcus. in the Museum of Victoria (NMV P). The Visceral disk strongly inflated, maximum specimens were whitened with an aerosol of inflation anterior of midvalve, almost geniculate ammonium chloride for photography. Latex casts in longitudinal outline (Figs 4G, 5A); lateral flanks were also made from some of the specimens. steep; umbonal flanks slightly swollen; cardinal extremities acute, lateral margins tapering sharply anterior to ears; shallow median sulcus originating Suborder Muir-Wood, 1962 on umbo; about 25 fine capillae, reaching 4 per Superfamily CHONETOIDEA Bronn, 1862 mm at anterior margin, capillae distinct with steep Family RUGOSOCHONETIDAE Muir-Wood, 1962 sides in cross section, wider than interspaces, SubfamilyRUGOS~ONETINAE Muir-Wood, 1962 ornament very faint to imperceptible on ears. Downloaded By: [Deakin University] At: 04:24 11 May 2009 Ventral interior (Fig. 4M, Fig. 5B) with short Neochonetes Muir-Wood, 1962 median septum, about one-third of valve length; strong, short vascular trunks, originating close to Typespecies. Chonetesdominus King, 1938, p. 259, pl. anterior end of median septum; hinge spines not 36, figs 1-7. Late , West Texas (USA). preserved but spine canals present, pointing towards umbo; interarea shallow, triangular; Comments. Shen & Archbold (2002) discussed pseudodeltidium rounded; teeth rounded internally, in detail the various subgenera of Neochonetes tapering laterally and angled anteriorly; surface that occur in South China, and their diagnoses coarsely radially papillose, ears with much finer, are followed herein. fainter papillae. Dorsal valve unknown. Neochonetes (Huangichonetes) Shen & Specimen ml mw hw t Archbold, 2002 NMV P309550a 2.01 5.45 v NMVP309550b 4.25 6.39 6.39 2.83 v NMV P309551 2.40 3.26 v+ Type species. Chonetes substrophomenoides Huang, Table 1. Measurements of Neochonetes (Huangichonetes) (1932, p. 3, pl. 1, figs 3-7). Lopingian, South China. geniculatus in mm: ml = maximum valve length; mw = maximumvalve width; hw = hinge width; t - valvethickness; Neochonetes (Huangichonetes) geniculatus sp. v - ventral valve; + - incomplete specimen. ALCHERINGA NEW PERMIAN BRACHIOPODS FROM SOUTH CHINA 281

r-

G Downloaded By: [Deakin University] At: 04:24 11 May 2009

:~ R Fig. 4A-R. A, G, l, J, N-O, Neochonetes (Huangichonetes) geniculatus sp. nov. A, G, I, J, N, NMV P309550a, holotype, ventral external mould and latex cast. A, N, ventral views of latex cast, xl, x7. G, lateral view of latex cast, x7. I, anterior view of latex cast, x7. J, posterior view of latex cast, x7. O, ventral valve external mould, x7, from Bed 174. K-M, Neochonetes (Huangichonetes) geniculatus sp. nov., NMV P309550b, holotype, counterpart of NMV P309550a, ventral internal mould and corresponding latex cast. K-L, posterior and ventral views of ventral internal mould, x7. M, latex cast of ventral internal mould, x7, from Bed 174. B-F, H, P-R, Neochonetes (Zhongyingia) linshuiensis sp. nov.. B, H, NMV P309559, holotype, ventral views of dorsal interior, xl and x4 respectively, from Bed 172. C, R, NMV P309556, ventral view of an incomplete dorsal interior, xl and x4 respectively, from Bed 150. D, NMV P309562, paratype, ventral view of an incomplete dorsal interior, x5, from Bed 174. E, NMV P309558, ventral view of a dorsal interior with some of internal shell layer removed, x3, from Bed 161. F, NMV P309561, paratype, dorsal view of a ventral interior, x5, from Bed 174. P, NMV P309553a, ventral view of a ventral internal mould, x6, from Bed 134. Q, NMV P309557, ventral view of a ventral internal mould, x4, from Bed 152. All specimens from the Changxing Formation at the Chuanmu Section. 282 MONICA J. CAMPI and G. R. SHI ALCHERINGA

Discussion. This new species is placed in other figured specimens of this species. This may Neochonetes (Huangichonetes) on the basis of simply be a reflection of the close generic the highly convex ventral valve, small ears, fine relationship, or these two specimens may actually capillation, well defined ventral median sulcus, belong to the new species, but the figures are and ventral internal structure, especially the not clear enough to be certain at this stage. presence of vascular trunks and a short median Shen et al. (2002) illustrated a very similar septum. The highly convex ventral valve is also new species of this genus from the Yongde characteristic of Tethyochonetes Chen et al., 2000, Formation of Yunnan. This species, N. (H.) and specimens described as Tethyochonetes sp. inflatus Shen et al. (2002, p. 671, fig. 3: 1-9) by Shen & Archbold (2002, figs 60-P) appear appears very similar to the Sichuan species as quite similar to the new species in terms of both are finely capillate, have a conspicuous convexity and outline of the ventral valve, median sulcus, small ears and are strongly although Shen & Archbold's specimens do not convex. They can be differentiated by the detail seem to have vascular trunks and are more finely of the inflation of the ventral valve, as N. (H.) papillose internally than the new species. inflatus is most strongly inflated at the umbo, Two other subgenera of Neochonetes are while N. (H.) geniculatus is most inflated at the present in the Permian of South China, and the midvalve. The latter species also has acute, well- new species can be separated from these. The defined ears whereas the former has obtuse, ill- highly convex dorsal valve and small ears clearly defined ears. differentiate the new species from Neochonetes (Zhongyingia) Shen & Archbold, 2002, which has Occurrence. Specimens were collected from Bed a gently to moderately convex ventral valve and 174 in the Changxing Formation and Bed 178 of the extended, acute cardinal extremities. Neochonetes Feixianguan Formation at the Chuanmu section. (Sommeriella) Archbold, 1982 can be separated Neochonetes (Zhongyingia) Shen &Archbold, 2002 from the new species as the former is widest at the midvalve rather than the hingeline and has Type species. Neochonetes zhongyingensis Liao, larger ears. The new species can be differentiated 1980a, p. 257, pl. 5, figs 10-13. Changhsingian from Neochonetes convexa Liao (1980b, pl. 5, figs (Lopingian), South China. 18-22) which Shen & Archbold (2002) placed in Neochonetes ( Sommeriella) strophomenoides Downloaded By: [Deakin University] At: 04:24 11 May 2009 Neochonetes (Zhongyingia) linshuiensis sp. nov. (Waagen, 1884) as Liao's material is more (Figs 4B - F, H, P- R) transverse than the new species and has rounder rather than acute cardinal extremities. Etymology. From Linshui county, in which the Neochonetes (Huangichonetes) geniculatus Chuanmu section is located. sp. nov. can be easily differentiated from N. (H.) substrophomenoides (Huang, 1932) and N. (H.) Material. Twelve specimens from the Changxing cursothornia (Xu & Grant, 1994) as the latter two Formation. The holotype is a dorsal valve, NMV species have more moderately convex ventral P309559; the paratypes are a ventral valve, NMV valves and much larger ears. Neochonetes (H.) P309561, and a dorsal valve, NMV P309562; and cursothornia also has square cardinal extremities the remaining material consists of one conjoined rather than the more acute cardinal extremities of specimen, NMV P309553a and counterpart, NMV the new species and some specimens of N. (H.) P309553b; two ventral valves, NMV P309557, substrophomenoides. However, two specimens NMV P309554a and counterpart, NMV P309554b; of N. (H.) cursothornia figured by Xu & Grant four dorsal valves, NMV P309552, NMV P309555, (1994, figs 15.4, 15.8) appear similar in general NMV P309556 and NMV P309558 and two shape and convexity to the new species, as they fragmentary specimens, NMV P309560 and NMV are less transverse and more inflated than the P309563. ALCHERINGA NEW PERMIAN BRACHIOPODS FROM SOUTH CHINA 283

Diagnosis. Small Neochonetes (Zhongyingia), Specimen # ml mw hw t subquadrate to trapezoidal in outline, with NMV P309552 6.57 7.66 d flattened ears, cardinal extremities acute. Ventral NMVP309553a 6.60 10.36 10.36 1.8 c valve moderately inflated for genus with a NMV P309553b 6.65 14.88 14.88 v shallow, broad median sulcus and coarse NMV P309554a 7.54 10.61 1.58 v capillation. Dorsal interior with indistinct median NMV P309554b 5.19 5.35 v NMV P309555 6.83 8.05 d septum. NMV P309556 4.84 11.74 10.67 0.88 d NMV P309557 7.54 9.13 1.6 v Description. Small for subgenus, subquadrate NMV P309729 4.49 8.8 8.8 0.75 d to trapezoid outline, hingeline close to straight, NMV P309730 5.57 9.39 9.39 1.05 d forming maximum width; 4 pairs of hinge spines; NMV P309731 2.84 7.73 7.73 + corpus cavity moderately inflated, ears flattened; NMV P309732 3.75 10.06 10.06 2 d longitudinal profile steeper at umbo and close to NMV P309733 3.08 8 1.24 v anterior margin, flatter medianly. NMV P309734 8.39 10.21 2 + Ventral umbo pointed, extending short Table 2. Measurements of Neochonetes (Zhongyingia) distance posterior to hinge, slightly elevated; linshuiensis in mm: c = conjoined valves; d = dorsal shallow median sulcus extending from umbo; valve; other abbreviations as above. coarsely capillate (2 per mm at anterior margin), alveolus small and rounded (Figs 5D, H); hinge capillae low and rounded, equal to interspaces sockets shallow, socket ridges short, extending and bearing elongate spinule apertures, laterally, diverging at about 20 ° from hingeline. sometimes bifurcating. Dorsal valve flat or Internal surface ornamented with radial rows of slightly convex; coarsely capillate (2 per mm); fine rounded papillae, with short rows of finer low median fold sometimes developed, papillae intercalating close to anterior margin containing 3 capillae. (Figs 5E, H), 5-6 rows on each interspace at Ventral interior with median septum and anterior margin, internal surface of ears smooth. vascular trunks; vascular trunks originating anterior to umbo, shorter than median septum; Discussion. This new species is closest to surface radially papillose, including ears, papillae Neochonetes (Zhongyingia) zhongyingensis fine, except for coarse papillae ornamenting Liao, 1980b, especially in internal features, median septum and vascular trunks. although the dorsal median septum and ventral Downloaded By: [Deakin University] At: 04:24 11 May 2009 Dorsal interior with indistinct median septum; vascular trunks are more prominent in Liao's cardinal process small, rounded, internally bifid; species. Both species have quite fine papillae ventral median papillae interareaseptum

posterior anterior .~/..inter~\ naldetails ~ not shown

0 6mm 4 mm , 1 ~ anterior Fig. 5. Neochonetes (Huangichonetes) geniculatus sp. nov. A, longitudinal profile of ventral valve. B, details of hinge region and internal ear ornament, (other internal features omitted) based on latex casts of holotype, NMV P309550a (Fig. 4G) and NMV P309550b (Fig. 4M). 284 MONICA J. CAMPI and G. R. SHI ALCHERINGA

internally, which become much finer anteriorly. References Neochonetes (Z.) zhongyingensis also has AB~CH, O.W.H., 1878. Geologische Forschungen in den coarser costellation, and is slightly more Kaukasischen Lginden. Theil 1. Eine Bergkalkfauna semicircular in outline. aus der Araxesenge bei Djoulfa in Armenien. A. HOlder, Wien, 126 pp. (German) The new species appears similar to ALL~SON, P.A., WIGNALL, P.B. & BRErr, C.E., 1995. Palaeo- Neochonetes (Huangichonetes) substropho- oxygenation: effects and recognition, 97-112. In menoides (Huang, 1932) in general size, shape Marine Palaeoenvironmental Analysis from Fossils. and external ornament. However, there are several D.W.J. BOSENCE& P.A. ALLISONeds, Geological Society of London, Special Publication 83. significant differences, as N. (H.) substropho- ARCHBOLD, N.W., 1982. Sommeriella, a new name for menoides has a more strongly inflated ventral the Permian chonetacean brachiopod subgenus valve compared to the moderate inflation of N. Sommeria Archbold. Proceedings" of the Royal (Z.) linshuiensis, and the median sulcus of the Society of Victoria 94, 10. BRONN, H.G., 1862. Die Klassen und Ordnungen der former is deep and broad, while the new species Weichthiere (Malacozoa): Dritte Klasse - Armkiemen- has a shallower sulcus. Internal features are Muschelm: Branchio-cephala. C.F. Winter'sche similar, reflecting the generic level relationship, Verhandlung, Leipzig & Heidelberg. 224-316. although N. (H.) substrophomenoides has coarser CAMPI, M.J., 2003. Permian Brachiopoda from China and Malaysia: Palaeoecology, palaeobiogeography papillae on the internal surface. and extinctions. Unpublished PhD thesis, Deakin Tethyochonetes Chen et al., 2000 is another University, 430 pp. genus common to the Lopingian of South China, CAMPI, M.J. & SHI, G.R., 2002. Linshuichonetes gen. nov., and the new species can be easily distinguished a new rugosochonetid (Brachiopoda) genus from the Liangshan Formation (Early Permian) in Sichuan, from members of this genus. Tethyochonetes has China, and its ecology. Acta Palaeontologica Sinica different internal features, as it possess much 41, 105-118. stronger median and lateral septa in the dorsal CHEN, Z.Q., JIN, Y.G. & Sin, G.R., 1998. Permian valve, while vascular trunks are absent from the transgression-regression sequences and sea-level changes of South China. Proceedings of the Royal ventral interior. Society of Victoria 110, 345-367. CHEN, Z.Q., SHI, G.R., SHEN, S. & ARCHBOLD,N.W., 2000. Occurrence. Specimens were collected from Beds Tethyochonetes gen. nov. (Chonetida, Brachiopoda) fi"om 126, 134, 150, 152, 161,172, and 174 from the the Lopingian (Late Permian) of China. Proceedings of Changxing Formation, Chuanmu section, the Royal Society of Vietoria 112, 1-15. EZAKI, Y., LJu, J.B. & ADACHI, N., 2003. Earliest Triassic Sichuan.

Downloaded By: [Deakin University] At: 04:24 11 May 2009 microbialite micro- to megastructures in the Huaying area of Sichuan Province, South China: Implications for the nature of oceanic conditions after the end- Acknowledgements Permian extinction. Palaios 18, 388-402. We thank Dr. Chen Zhongqiang (University of FERGUSON, L., 1985. Size diminution in vertically inclined lingulid shells as an indicator of proximity of Western Australia) and Mr. Peng Yuanqiao (Deakin conditions near the end of their tolerance range; an University) for assistance with collection of material example from the shale underlying the Second Abden in the field, Prof. Tong Jinnan (China University of Limestone (Visean), Kinghorn, Fife, Scotland. In 1st Geosciences) for access to fossil material, Dr. Zhu International Congress on Brachiopods: Les brachiopodes fossiles et actuels: le point des Qi (Southwest Petroleum College, Chongqing) for connaissances sur 600 M. A. de l'histoire de la vie assistance with logistics in the field and Professors marine, Abstracts. Universit6 de Bretagne Neil Archbold (Deakin University) and Shen Occidentale, Brest, France, 33. Hz, X.L. & Sin, G.R., 1996. The sequence of brachiopod Shuzhong (Nanjing Institute of Geology and assemblages in South China, 111-115. In Palaeontology) for helpful discussion regarding Brachiopods. Proceedings of 3rd International these species. This work was partially undertaken Brachiopod Congress, P. COPPER & J.S. Jly eds, A.A. while the principle author was the recipient of a Balkema, Rotterdam. HUAN~, T.K., 1932. Late Permian Brachiopoda of Deakin University Postgraduate Award. We also southwestern China, Part 1. Geological Survey of thank Drs Jun-ichi Tazawa and Patrick Rachboeuf China, Palaeontologia Sinica, Series B 9, 1-107. for their critical comments. HUANG, T.K., 1933. Late Permian Brachiopoda of ALCHERINGA NEW PERMIAN BRACHIOPODS FROM SOUTH CHINA 285

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