Volume 61,Issue 3,2022 Table of Contents

1 Advances in Late Paleozoic palaeontology and stratigraphy in China: a tribute to the achievements of Professor Liao Zhuoting

QI Yu-ping , YAO Le , WANG Qiu-lai , CHEN Zhong-qiang , WANG Xiang-dong

2022, 61(3):330-342. DOI: 10.19800/j.cnki.aps.2022301 CSTR:

[Abstract](292) [HTML](0) [PDF 3.25 M](1560)

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Through the efforts of several generations of the Chinese palaeontologists and stratigraphers, extraordinary progress has been made in the Late Paleozoic palaeontology and stratigraphy of China. Professor Liao Zhuoting, one of the pioneering palaeontologists, dedicated his lifetime to advancing the Chinese palaeontological and stratigraphical research. In his 60-year palaeontological career, Professor Liao worked on palaeontology and stratigraphy in almost all the regions across China. His research interests include palaeontology, palaeoecology, palaeobiogeography, biostratigraphy, tectonostratigraphy, and their implications in mineral, oil and gas explorations. Professor Liao’s studies focus mainly on the Carboniferous and Permian brachiopod faunas. Together with his colleagues, Professor Liao has established 22 new genera and 172 new brachiopod species. His extensive research on the Carboniferous and Permian palaeontology and stratigraphy in South China and other regions in China, including documentation of a large number of brachiopod fossils, has provided abundant first-hand palaeontological data that help define the Global Stratotype of Section and Point for the Permian-Triassic boundary at the Meishan section of Changxing County, Zhejiang Province and demonstrate the extinction-survival-recovery pattern of brachiopod faunas over the P–Tr transition in later studies. Professor Liao dedicated his 35 years of efforts to the palaeontology and stratigraphy in the Altaid, Tianshan, and Kunlun mountains, as well as the Junggar, Santanghu, Tuha, and Tarim basins in Xinjiang Province, northwest China. He, together with his colleagues, established integrated stratigraphical frameworks for the Carboniferous and Permian rocks in Xinjiang, which provide a fundamental basis for mineral and hydrocarbon explorations in Xinjiang and adjacent regions, Central Asia. This special volume, consisting of 13 papers on the Devonian through Triassic palaeontology and stratigraphy of China, is a tribute to the achievements of Professor Liao Zhuoting.

2 Recovery of brachiopods after the Permian–Triassic mass extinction: diversity, ornamental evolution, and paleobiogeography

CHEN Zhong-qiang , GUO Zhen

2022, 61(3):343-357. DOI: 10.19800/j.cnki.aps.2022014 CSTR:

[Abstract](352) [HTML](0) [PDF 6.78 M](1381)

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The Brachiopoda is one of the most important components in the Paleozoic benthic community. During the Permian-Triassic (P-Tr) mass extinction, many orders of this phylum were wiped out. Brachiopods became a marginal group in the Mesozoic and Cenozoic oceans. To investigate the effects of the P–Tr extinction on the macroevolution of this clade, we re-evaluated the diversity variations of global brachiopods during the Triassic, traced the evolutionary trend of the development of ornamentation of the Triassic and Early Jurassic elements, and analyzed the changes of brachiopod paleobiogeographic provincialisms before and after the extinction. Taxonomic diversity at the genus level abruptly declined across the P–Tr crisis, increased rapidly in the Anisian, and finally reached the highest level of the Triassic during the Norian. Compared with genera that originated in the Late Triassic, those originated in the Early Triassic are shorter in longevity. The novel ornamentation index was employed to evaluate the development of the ornamentation of brachiopods. The results show that the four main orders of the early Mesozoic brachiopods behaved differently in ornamental evolution, suggesting that their ornamental trajectories were not controlled by a common factor. Compared with the escalation-driven evolution, their evolutionary history played a more important role. The ornamentation development shows no significant relationship with the strengthened predation pressure, at least during the early Mesozoic. Network analysis indicates that the paleobiogeographic provincialism disappeared after the extinction, and re-emerged in the Olenekian. Along with the occurrence of many endemic elements during the Middle Triassic, the affinities of brachiopod faunas became much lower among various regions in the world.

3 A reinvestigation of the conodonts from the boundary interval between the Permian Kuhfeng and Chihsia formations at the Zhengpanshan section in Longtan, Nanjing City

YUAN Dong-xun , SHEN Shu-zhong

2022, 61(3):358-370. DOI: 10.19800/j.cnki.aps.2022020 CSTR:

[Abstract](258) [HTML](0) [PDF 8.58 M](1621)

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The base of the Guadalupian Series (Middle Permian) has been defined by the first appearance datum (FAD) of the conodont Jinogondolella nankingensis at the Stratotype Canyon section in the Guadalupe Mountains National Park in Texas, USA. However, both the holotype of the conodont J. nankingensis and the geochronologic age of the Cisuralian-Guadalupian boundary (CGB) in the official International Chronostratigraphic Chart were obtained from the base part of the Kuhfeng Formation at the Zhengpanshan section in Longtan in the suburb of Nanjing City, southeast China. Therefore, the Zhengpanshan section has actually become one of the international standard sections for the correlation around the CGB. However, conodont succession from the section has not been studied in detail so far. In this paper, we review previously reported data and report our newly recovered conodont specimens from the Zhengpanshan section. In total, nine species of five genera have been identified here from the topmost part of the Chihsia Formation to the middle part of the Kuhfeng Formation at the section, and they are Hindeodus minutus, H. typicalis, H. sp., Jinogondolella nankingensis, J. cf. aserrata, J. sp., Pseudohindeodus ramovsi, Pseudosweetognathus costatus and Sweetognathus subsymmetricus. Three conodont zones, the S. subsymmetricus–Pseudosweetognathus costatus Assemblage Zone in the topmost part of the Chihsia Formation and the Jinogondolella nankingensis and J. cf. aserrata zones in the lower-middle part of the Kuhfeng Formation, have been recognized at this section. The updated conodont biostratigraphy shows that the CGB should be placed at the base of the Kuhfeng Formation at the Zhengpanshan section, and the lower part of the Kuhfeng Formation belongs to the Roadian Stage. The middle part of the Kuhfeng Formation has yielded the conodont J. cf. aserrata, which probably indicates a Wordian age. The precise Roadian-Wordian boundary (RWB) is not well identified, because a nearly 2 m interval between the last occurrence (LO) of J. nankingensis and the first occurrence (FO) of J. cf. aserrata is barren of conodonts. The radiolarian, ammonoid and brachiopod horizons, including Altudoceras, Neoplicatifera huangi, Pseudoalbaillella scalprata-P. longtanensis and Hegleria mammilla in ascending order, reported from the Zhengpanshan section are well correlated with the updated conodont zones

4 Progress of studies on Carboniferous conodonts in China

HU Ke-yi , QI Yu-ping , WANG Qiu-lai , WANG Zhi-hao , WANG Xiang-dong

2022, 61(3):371-396. DOI: 10.19800/j.cnki.aps.2022015 CSTR:

[Abstract](227) [HTML](0) [PDF 20.56 M](1895)

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This paper summarizes the history and current situation of the Carboniferous conodont research in China from the 1970s to the present. The taxa of Carboniferous conodonts established based on the Chinese materials are reviewed. Only two of the five genera/subgenera are valid. Among the 99 species, 48 species are valid, seven species are invalid, and 44 species still need further study. The conodont biostratigraphy of 10 stratigraphic subprovinces in four Carboniferous stratigraphic provinces in China is discussed and revised. In the South China Subprovince, the most complete zonation (43 zones) is recognized. In other stratigraphic subprovinces, 10 zones in Junggar, eight zones in Qiangtang, eight zones in Tarim, eight zones in Qilian–Helan Mountains, and six zones in Inner Mongolia–Jilin are also recognized. There are only three zones in North China, six zones in western Yunnan, five zones in southern Tibet, and two zones in Gangdise. The Global Boundary Stratotype Section and Point (GSSP) of four Carboniferous stages, namely Serpukhovian, Moscovian, Kasimovian, and Gzhelian stages, has not yet been established. Conodonts have the most potential as boundary markers for these four stages. The Naqing section in Luodian, Guizhou, South China is a candidate stratotype section for these four stages. The section yields abundant conodont fossils, including potential boundary markers for those four stages. Great progress has been made in the study of the conodont evolutionary lineages in recent years.

5 The discovery of Adoketophyton subverticillatum from the Devonian Mangshan Group of Duyun, Guizhou Province and its significance

HUANG Pu , WANG Yao , ZHANG Xiao-le

2022, 61(3):397-406. DOI: 10.19800/j.cnki.aps.2021057 CSTR:

[Abstract](233) [HTML](0) [PDF 8.31 M](1572)

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Adoketophyton subverticillatum (Li et Cai) Li et Edwards is one of the representative and well-known plants in the Early Devonian Posongchong flora of Yunnan, and its external morphology and anatomy have been described in detail. One specimen collected from the lower part of the Mangshan Group at the Baoyang section of Duyun, Guizhou Province is assigned herein to A. subverticillatum, based on features revealed by serial dégagement. The specimen, broken at both ends, is ca. 19.3 mm long, and shows a fertile axis comprising of sporophylls and sporangia. The fertile axis is slender, ca. 0.9 mm wide. The fan-shaped sporophylls, 6.2 mm in length and 6.4 mm in maximum width when complete, are oppositely and decussately inserted on the fertile axis to form four vertical rows. An incomplete sporangium reaches up to 4.1 mm high and 1.1 mm wide. The temporal and spatial distribution of Adoketophyton indicates that this genus is a representative element of the Early Devonian floras, thus suggesting an Early Devonian age for the lower part of the Mangshan Group. The discovery of A. subverticillatum in Duyun confirms the dispersal and interrelationship at the genus and species levels between the Posongchong flora of Yunnan and plants from the Mangshan Group of Duyun, Guizhou. The studies of Adoketophyton and other related taxa suggest that leafy structures, such as sporophylls, bracts, and branch-leaf complexes, were widespread in the South China Block during the Early Devonian, when the atmospheric CO₂ level was high.

6 Discovery of the micromorphic brachiopod Lambdarina from the Lower Carboniferous of Long’an, Guangxi, with remarks on the evolutionary history of the superfamily Lambdarinoidea

GUO Wen , NIE Ting , QIE Wen-kun , LU Gang , HUANG Xiang-lin , SUN Yuan-lin

2022, 61(3):407-417. DOI: 10.19800/j.cnki.aps.2022016 CSTR:

[Abstract](116) [HTML](0) [PDF 10.78 M](1546)

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A species of the superfamily Lambdarinoidea (Brachiopoda), namely Lambdarina cf. sinensis, from the Tournaisian of Long’an, Guangxi is described herein. The evolutionary history of this micromorphic rhynchonellid group is discussed based on their geologic and palaeogeographic distribution from the reported fossil records. The early lambdarinoids seem to appear first in central Europe during the Famennian and then migrated to the South China Plate where they experienced a biodiversification during the Tournaisian. The late Tournaisian and Visean witnessed the dispersal of succeeding lambdarinoid taxa to Europe of the western Paleo-Tethys Ocean and Australia on the northeastern margin of the Gondwana. Two evolutionary lineages are proposed here according to shell outlines and development of median ridge-like folds on the ventral valve. The lambdarinoids are supposed to favor warm and low-energic habitats at low latitude.

7 The origin of primitive archaediscacean foraminifers and the development of their wall structure

The origin of primitive archaediscacean foraminifers , the development of their wall structure

2022, 61(3):418-426. DOI: 10.19800/j.cnki.aps.2021077 CSTR:

[Abstract](142) [HTML](0) [PDF 2.01 M](1391)

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Archaediscacean foraminifers are perhaps one of the most useful biostratigraphic marker microfossils from the early Visean to early Serpukhovian. They are widespread in normal marine, benthic, shallow-shelf carbonate environments. The subfamily Planoarchaediscinae of the superfamily Archaediscacea appeared in the middle early Visean. The wall structure of the planoarchaediscins is the most primitive in this superfamily, which provides an evolutionary link between the Archaediscidae and the ancestral pseudoammodiscids. The wall of the archaediscaceans is predominantly dark and microgranular. The hyaline-radial layer is either limited to the umbilical area, or it forms a thin covering over the sides, or sometimes it covers the periphery of the interior volutions. Regarding the ancestor of the archaediscids, this topic is still open to debate. This study focuses on the specimens of Lapparentidiscus and Viseidiscus with continuous stratigraphic appearances from the Shidengzi Formation at the Malanbian section in South China. The result suggests that Lapparentidiscus is a unique pseudoamrnodiscid that gives rise to the archaediscids. The evolution trend of primitive archaedis cacean foraminifers is: Lapparentidiscus talasicus–Viseidiscus eospirillinoides–Viseidiscus monstratus. The hyalin-radial layer of the archaediscids appears as a replacement instead of a filling. It replaces the dark, microgranular layer gradually, and extends from the umbilical area to the sides.

8 Late Visean (Early Carboniferous) reef-building rugose corals from Yashui, Guizhou Province

WANG Hui , YAO Le , LIN Wei , WANG Qiu-lai

2022, 61(3):427-438. DOI: 10.19800/j.cnki.aps.2022041 CSTR:

[Abstract](163) [HTML](0) [PDF 9.02 M](1575)

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Accompanying the Late Devonian Frasnian-Famennian Kellwasser and the Devonian–Carboniferous Hangenberg mass extinction events, the main Devonian reef builders of stromatoporoid sponges and corals disappeared and dramatically declined, respectively. The subsequent Early Carboniferous is the critical recovery period for the reef-building corals. The Visean (Lower Carboniferous) Shangsi Formation at the Yashui section, Guizhou Province has yielded abundant coral autobiostromes (biostromal reefs). Based on foraminiferal biostratigraphy, the age of these coral biostromes was proposed to be late Visean. The Shangsi coral biostromes mainly consist of colonial rugose corals with low diversity. Two species belonging to two genera are identified: Siphonodendron pauciradiale and Stylostrotion pentalaxoidea. Siphonodendron pauciradiale is widely distributed in the late Visean coral reefs in Western Europe and North Africa, while S. pentalaxoidea is limited locally in the South China Block. The occurrence of the cosmopolitan reef-building coral S. pauciradiale during the late Visean at the Yashui section indicates that the recovery and evolutionary process of the coral reefs in South China is consistent with that of the globe after the Late Devonian extinctions.

9 Rugose corals from the Hezhou Formation (Mississippian, Carboniferous) in Chaohu, Anhui Province

LIN Wei , YAO Le , HUANG Xing

2022, 61(3):439-457. DOI: 10.19800/j.cnki.aps.2022048 CSTR:

[Abstract](190) [HTML](0) [PDF 26.54 M](1753)

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The Mississippian is a subperiod of the Carboniferous during which the rugose corals prospered. During the late Visean and Serpukhovian, the onset of the Late Palaeozoic Ice Age, coupled with the Hercynian Orogenic movement, resulted in rapid sea-level changes. Regions in the western Palaeotethys, such as the present-day western Europe, were affected by the sea-level fluctuations and facies changes recorded in marine sedimentary successions, resulting in the punctuation and scarcity of rugose coral fossil record. South China, however, was less affected, and shallow marine carbonate successions with abundant rugose coral fossils were widespread and well-developed. From a collection of coral fossils recovered from the Hezhou Formation near Chaohu, Anhui Province, Lower Yangtze region, a total of 15 species belonging to 11 genera, including two species conformis, one species affinis and three species indeterminabilis are recognized. The coral fauna consists of elements of the late Datangian type from South China, and can be assigned to the Aulina rotiformis Zone (the upper part of the former Yuanophyllum Zone), inferring the Serpukhovian age for the Hezhou Formation. The assemblage correlates well with the Neoclisiophyllum yengtzeense–oninckophyllum stellatum subzone and Lophophyllum lophyphylloidea subzone of the Zimenqiao Formation in central Hunan, and to the Palaeosmillia militaris Zone and Lithostrotion decipiens–Dibunophyllum bipartitum Zone of the Zhaojiashan Formation in western Guizhou. The size and growth-form, the symbiotic relationship of the corals, and the taxonomical composition of the coral assemblage indicate that the fauna lived in a relatively turbid, restricted and eutrophic nearshore shallow marine environment.

10 The Pennsylvanian conodont genus Neognathodus from southern Guizhou, South China: evolution and its stratigraphic significance

WANG Qiu-lai , QI Yu-ping

2022, 61(3):458-471. DOI: 10.19800/j.cnki.aps.2022047 CSTR:

[Abstract](100) [HTML](0) [PDF 14.68 M](1590)

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Early to Middle Pennsylvanian conodonts from the Naqing section in Luodian, Guizhou, South China are well-documented. Based on several conodont genera from deep-water facies, their successions are applied as the standard conodont zonation from China for global correlation. Meanwhile, Neognathodus, a shallow-water genus that is rare in slope facies in South China, is of great importance in stratigraphic correlations especially in shallow-water facies in North America, East Europe, and North China. In this report, with reference to the standard zonation, successions of Neognathodus from three sections of slope facies (Naqing, Narao, and Luokun) in Luodian are studied and discussed. Furthermore, based mainly on the platform symmetry of the P₁ elements, an evolutionary trend divided into five stages is preliminarily introduced. The resurgence of symmetric elements of this genus in Stage 4 (e.g., N. bothrops, N. colombiensis, and N. nataliae) is suggested as an auxiliary index for the base of the global Moscovian Stage.

11 New discovery of Permochoristidae (Insecta, Mecoptera) from the Guadalupian of Chaohu City, Anhui Province, China

LIAN Xin-neng , CAI Chen-yang , HUANG Di-ying

2022, 61(3):472-478. DOI: 10.19800/j.cnki.aps.2022029 CSTR:

[Abstract](218) [HTML](0) [PDF 13.69 M](1650)

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Permochoristidae is a representative of the mecopterans in the Permian and a stem group of Mecoptera. The Permian is a period during which mecopterans were prosperous, however, Permian mecopterans from China are extremely rare, with only one previously reported species. We establish a new genus and species, Chaohuchorista liaoi gen. et sp. nov., based on one specimen collected from the Guadalupian Yinping Formation of Chaohu City, Anhui Province, eastern China. The diagnostic characters of the genus include small and elongate forewing, Rs₁ and Rs₂ combined into a single vein, Rs₄ with two branches, M with five branches and M₂ with two branches. Our new finding highlights the mecopteran diversity during the Permian in China.

12 Discovery of gigantopterid plants from the upper Permian of Daha Coalfield, Qinghai Province

SUN Wei , CHEN Ming , SONG Chun-yan , XIE Shang-ke , LI Xue-ren , ZHOU Wei-ming

2022, 61(3):479-493. DOI: 10.19800/j.cnki.aps.2020061 CSTR:

[Abstract](150) [HTML](0) [PDF 5.76 M](1558)

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The Tibetan Plateau contains part of the boundary between the Permian Cathaysian and Gondwanan floras. In this paper, we report a new gigantopterid fossil locality from the upper Permian of Daha Coalfield, an area close to the Wuli Coalfield in the Tuotuohe District of Qinghai Province, China. We describe five species in four genera of gigantopterid leaves collected from a mudstone bed. Among these fossils, the discovery of double-meshed gigantopterids such as Gigantopteris cf. dictyophylloides、Filigigantopteris asymmetrica and F. dahaia further demonstrates the palaeogeographic affiliation of the fossil locality with the Cathaysian Realm, and suggests the existence of a land bridg for the westward expansion of the Cathaysian flora to the Middle East. In addition, the leaves vary greatly in size. The lack of long petioles, hooks and prickles suggests that these gigantopterid plants may not be climbers.

13 Dissepimental Characteristics of the Late Permian Rugose Coral Ipciphyllum naoticum with Discussion on the Taxonomic Significance of Naotic Structure

WANG Xiao-juan

2022, 61(3):494-505. DOI: 10.19800/j.cnki.aps.2022009 CSTR:

[Abstract](128) [HTML](0) [PDF 22.50 M](1570)

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Ipciphyllum, which is a dissepimental, colonial rugose coral representative of the Waagenophyllinae, is one of the most diversified and abundant genera in the Permian Maokou Formation in South China. Ipciphyllum naoticum from the Late Permian Gyanyima section in Tibet, southwest China is the only species in Ipciphyllum that develops naotic dissepiments. In this study, the dissepimental shapes of Ipciphyllum naoticum in different ontogeny stages can be observed in detail with serial transverse sections. In the embryonic and neanic stage, septa strongly thickened in peripheral part. Concentric dissepiments inconspicuously developed in the middle part in transverse section. In the adult stage, septa become thin in peripheral part and obviously thinner than that of the central part within the tabularium. Dissepiments developed at the base of partial septa. Firstly, these dissepiments partly appear as arc-shaped plates, and then some arc-shaped plates overlap each other and form naotic structure. The lateral dissepiments sporadically appear in this later stage. The dissepiments of the same species are various in shape, so it is not suitable to regard one special shape of dissepiments such as arc-shaped plates, or naotic structure as species character or even higher classification standard. Pervious study about the Permian naotic-dissepiment rugose corals in China implies that naotic-dissepiment development was probably related to enhanced hydrodynamic force. Naotic dissepiments are not of higher taxonomic significance than species level. On this basis, combined with various shape of Ipciphyllum naoticum, this paper suggests that Protonaoticophylidae and Protonaoticophylinae with naotic structure (dissepiments) as the important classification characteristics should be abandoned.

14 Quantitative evaluation of the taxonomy of Pseudoschwagerina species from South China

WANG Juan , WANG Yue

2022, 61(3):506-522. DOI: 10.19800/j.cnki.aps.2021028 CSTR:

[Abstract](152) [HTML](0) [PDF 4.70 M](1449)

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This paper reclassify 48 species of Pseudoschwagerina previously reported from South China. The measurements of the length, width of the shells, form ratios, radius of the proloculus, number of inner tightly-coiled and outer loosely-coiled volutions are entered into a database for quantitative analysis. Some holotype specimens stored in Nanjing Institute of Geology and Palaeontology have been re-measured. All species are projected onto a 3-D map using the software Grapher to show inter-species distances. Two groups can be recognized. One group is characterized by having fusiform to long fusiform shells, strong septal folding, and a form ratio greater than 2.0. The other group is characterized by having subspherical or thickly fusiform shells, weak to strong septal folding, and a form ratio less than 2.0. A further comparison of the inner volutions as well as the length of the shells of all species is made. Our analysis shows that species with longer shells usually contain long fusiform to fusiform inner shells and have less, normally 1–1.5, inner volutions, whereas those species with shorter shells contain short fusiform to subspheric younger shells and have relatively more, usually 2–4, inner volutions. Therefore, four and five subgroups are recognized in the first and second group, respectively, based on a comprehensive evaluation of the features of the younger shells, the size of the proloculous and the length of the shells.

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