Volume 62,Issue 3,2023 Table of Contents

1 Nisusia from the Cambrian Jialao Formation in Jianhe, Guizhou

WU Kai-lun , YANG Xing-lian , CHEN Zheng-peng , WEI Bu-qing , FENG Rong

2023, 62(3):333-349. DOI: 10.19800/j.cnki.aps.2021074 CSTR:

[Abstract](333) [HTML](0) [PDF 29.65 M](713)

Abstract:
Nisusia Walcott 1905 is one of the early representatives of articulate brachiopods. This genus is widely distributed in the world. However, most reports are from the last century and the classification of many species needs further research for lack of high quality illustrations and unequivocal descriptions. Nisusia has also been reported from many places in China, but little in-depth research has been done on its classification and evolutionary trend. In 2019, Holmer et al. further studied the classification and phylogeny of Nisusioidea and proposed a phylogenetic tree for the superfamily, which contains four genera. It is noteworthy that Holmer et al. revised N. guizhouensis, erected based on specimens from the Kaili Formation in Jianhe, as Nisusia? due to missing shell spine decoration, a character initially observed in Nisusia specimens, therefore, more fossil evidence is needed for further study to confirm this assignment.Recently, many Nisusia specimens have been collected from the Cambrian Jialao Formation in Jianhe, Guizhou. Two species (N. guizhouensis and N. granosa) and one indeterminate species (Nisusia sp.) are recognized. However, the distinct spines on these shells are different from those on the shells of N. guizhouensis reported from the underlying Kaili and Tsinghsutung formations. In order to further clarify if these differences are intraspecific or interspecific, quantitative morphological analysis was conducted for the specimens from the Jialao Formation and its underlying Kaili and Tsinghsutung formations. First, the tpsDig232 program was used to obtain landmark point information. Then, the data was imported into PAST software for multivariate analysis and TPS analysis. Landmarks and semi-landmarks were taken on each contour of Nisusia after getting the complete fossil photos.The PCA results of Nisusia guizhouensis from the Tsinghsutung, Kaili and Jialao formations show that there is a large overlapping area of morphological features in the outline, indicating that the ventral and dorsal valves have little morphological differences. Therefore, these ventral and dorsal valves belong to the same species. The CVA results show that the dorsal valves of N. guizhouensis from these three stratigraphic units are dispersed, indicating that there are subtle morphological differences of N. guizhouensis from different strata. The TPS analysis shows that the overall morphological changes of ventral and dorsal valves from the Tsinghsutung, Kaili and Jialao formations are consistent. The cardinal extremities of ventral and dorsal shells expand outwards from the Tsinghsutung Formation to the Jialao Formation, while the anterior of N. guizhouensis valves shrink inward and the visceral cavity also becomes smaller. The variation of ventral valves is slightly larger than that of dorsal valves; the variation of N. guizhouensis is more significant from the Kaili Formation to the Jialao Formation than that from the Tsinghsutung Formation to the Kaili Formation.Results from the quantitative morphological analysis of Nisusia guizhouensis show little change from the Tsinghsutung Formation to the Kaili Formation, while there is a relatively significant change from the Kaili Formation to the Jialao Formation. These trends may be resulted from environmental changes. The lithology of the Tsinghsutung and Kaili formations is mainly mudstone, and the paleoenvironment is interpreted as a deep-sea with weak hydrodynamic conditions. However, the water gradually became shallow from the middle to the top part of the Kaili Formation. The overlying Jialao Formation is mainly composed of siltstone and sandstone, and the sea may become shallower with strong hydrodynamic conditions. Most energy needs of the brachiopods are provided by organic matter filtered out of water flows. The visceral cavity serves as the place where filter feeding takes place: a larger visceral cavity means access to more food. Comparing specimens of N. guizhouensis of the same size, the visceral cavity of those from the Jialao Formation is the smallest. This may be because Nisusia, as a benthic animal, lived in an environment with shallower water and stronger hydrodynamic environment of Jialao Formation and they may get enough food relying on a stronger current instead of pumping more water into the visceral cavity. In addition, the specimens of the Jialao Formation are smaller, and the shell surface has obvious spine decoration with sparser and thicker ribs. Shell spines are important fixation organs of benthic brachiopods. The sparser and thicker radiation increases the efficiency of drainage in water flow. This study not only supplements more data to the biological assemblage of the Jialao Formation in Jianhe, Guizhou, but also provides new information on the systematics and the adaptive evolution of the brachiopods.

2 Spiriferide brachiopods from the early Famennian (Late Devonian) of western Europe

WANG Yi-ning , MA Xue-ping , Volker Ebbighausen , R. Thomas Becker

2023, 62(3):350-375. DOI: 10.19800/j.cnki.aps.2022030 CSTR:

[Abstract](150) [HTML](0) [PDF 50.80 M](716)

Abstract:
Senzeilles in the Namur province of Wallonia, Belgium, a traditional area for the study of Upper Devonian stratigraphy in western Europe, yields abundant and well-preserved Famennian spiriferide brachiopods. Nevertheless, modern studies of these brachiopods are rare; in addition, earlier taxonomic works do not meet the demands of modern thematic studies (i.e., accurate stratigraphy and correct identification of the brachiopods). Therefore, it is significant to conduct thorough investigations on brachiopod faunas across the Famennian–Frasnian (F–F) boundary in terms of modern taxonomy. This study aims at a systematic description of the early Famennian “Cyrtospirifer” brachiopods from the Senzeilles area. Our comparative study of this group of brachiopods from South China and western Europe shows that the circumscription of the genus Plicapustula should be expanded (i.e., with pustules both on plications and in grooves). The European species of Plicapustula possess pustules both on plications and in grooves as well as fewer sinal plications, whereas those from South China possess pustules only on plications as well as more, numerous sinal plications. Considering such differences, a new subgenus Paraplicapustula is proposed to include Plicapustula species from western Europe. In addition, the genus Sinospirifer is first recognized and described from western Europe in the present paper. Five species in two genera, including one new subgenus and three new species, are described based on shell size, nature of posterior shell thickenings, number and pattern of flanks and sinal plications, and microornamentation. These taxa are: Plicapustula (Paraplicapustula) eleganta subgen. et sp. nov. Ma and Wang, Plicapustula (Paraplicapustula) magna subgen. et sp. nov. Ma and Wang, Sinospirifer transversus sp. nov. Ma and Wang, Plicapustula (Paraplicapustula) sp., and Sinospirifer subextensus (Martelli, 1902).

3 Changhsingian (Late Permian) calcareous algae and foraminifers from Shuicheng, western Guizhou, South China and their paleoenvironmental significance

DANG Zhi-ying , SHEN Yu-lin , CHEN Fei-yang , ZHANG Li-jun , YANG Tian-yang , JIN Jun , ZHAO Yong

2023, 62(3):376-389. DOI: 10.19800/j.cnki.aps.2023006 CSTR:

[Abstract](179) [HTML](0) [PDF 8.28 M](693)

Abstract:
Three species in three genera of calcareous algae, including Gymnocodium bellerophontis, Permocalculus sp., and Tauridium kurdistanensis and ten species in eight genera of foraminifers, including two species in two genera of fusulinids, are identified from Well K576 near Shuicheng, western Guizhou, South China. The foraminifers mainly consist of Reichelina sp. indet., Nankinella sp., Pachyphloia schwageri, Pachyphloia sp., Geinitzina sp., Nestellorella sp. indet., Howchinella sp., Hemigordius aff. saranensis, Hemigordius sp., and Midiella sp. indet. The fossil fauna is divided into seven assemblages: Foraminifer-Calcareous alga-Ostracod, Foraminifer-Brachiopod-Ostracod, Ostracod-Bivalve-Gastropod, Foraminifer-Calcareous alga-Bivalve, Foraminifer-Brachiopod-Ostracod, Foraminifer-Calcareous alga-Bivalve, and Ostracod assemblages. Based on sequence stratigraphy of the study area, characteristics of the vertical sedimentary sequence, and analysis of log data, the palaoenvironment of the deposits that yield the Foraminifer-Calcareous alga-Ostracod assemblage (SQ3-3) and Foraminifer-Brachiopod-Ostracod assemblage (SQ3-4) is a delta front. The strata that yields the Ostracod-Bivalve-Gastropod assemblage (SQ4-1) was deposited on a tidal flat. The paleoenvironment of the Foraminifer-Calcareous alga-Bivalve (SQ4-2), the Foraminifer-Brachiopod-Ostracod (SQ4-3), and the Foraminifer-Calcareous alga-Bivalve (SQ4-3) assemblages is a lagoon. The strata that yields the Ostracoda assemblage (SQ4-3) was deposited on a sand flat. A low-energy, calm, and warm nearshore delta and barrier lagoon depositional systems were generally developed during the Changhsingian in Shuicheng area of western Guizhou. In addition, analysis of the fossil assemblages indicates that biodiversity increases significantly when both foraminifers and calcareous algae are present in the lagoon environment.

4 Discovery of important taxonomic characters of the type species of the Late Triassic clam shrimp Anyuanestheria (Crustacea: Spinicaudata) from Jiangxi

LI Gang

2023, 62(3):390-397. DOI: 10.19800/j.cnki.aps.2022031 CSTR:

[Abstract](113) [HTML](0) [PDF 25.90 M](648)

Abstract:
Re-examination of the type specimens of the clam shrimp type species Anyuanestheria subquadrata Zhang and Chen in Zhang et al., 1976 using a scanning electron microscope (SEM) has revealed details of carapace characters that are of important taxonomic value. These characters have hitherto never been documented. The new characters include medium to large polygonal reticulations on growth bands on the dorsal part of the carapace. Two types of ornaments are present on the growth bands of the ventral and posterior parts of the carapace: fine and widely spaced radial lirae on the upper part and medium to large reticulations on the lower part of each growth band. The walls of the polygonal reticulations are thin with shallow depressions, resulting in flat polygonal protrusions on the external mould. This ornamentation pattern is different from the original description. Based on these observations, the diagnosis of Anyuanestheria is emended.

5 A new species of Trigonostemon Blume (Euphorbiaceae) from the middle Miocene of Fujian, Southeast China and its paleoclimatic and paleoecological significance

ZHENG Qing-dan , DONG Jun-ling , ZHENG Dong-yu , SUN Bai-nian

2023, 62(3):398-409. DOI: 10.19800/j.cnki.aps.2022044 CSTR:

[Abstract](187) [HTML](0) [PDF 13.00 M](858)

Abstract:
A new species of Trigonostemon Blume, namely Trigonostemon zhangpuensis sp. nov. J.L. Dong et B.N. Sun, is established based on well-preserved leaf fossils from the Fotan Group in Zhangpu County, Fujian, Southeast China. The new species is characterized by oblanceolate lamina with cuneate and narrow base, acuminate apex, serrate margin at the distal part and entire margin at the proximal 1/3 part of the lamina, pinnate brochidodromous venation with a stout primary vein thinning gradually toward the apex, anomocytic stomatal complex, sparsely distributed and randomly arranged stomata, and normal epidermal cells with striate periclinal walls. The morphological characters of the fossil leaves closely resemble those of extant Trigonostemon xyphophyllorides (Croiz.) L. K. Dai et T. L. Wu., which is endemic to Hainan Province. The ecological amplitude of the fossil species is similar to that of its nearest living relatives; the paleoclimate of the Zhangpu area during the middle Miocene might be similar to that of Hainan Province today. We recognize thirteen insect-mediated damage types belonging to six functional feeding groups on the leaves of T. zhangpuensis. The damage types include hole feeding, margin feeding, skeletonization, surface feeding, galling, and piercing and sucking. Moreover, our observation suggests that phytophagous insects feeding on leaves of T. zhangpuensis might be relatively diverse.

6 Neogene Picea leaves from the Baoshan Basin of Yunnan and their implication for paleoaltimetry

ZONG Hao-ran , WANG Nian , HUANG Yong-jiang

2023, 62(3):410-423. DOI: 10.19800/j.cnki.aps.2023011 CSTR:

[Abstract](243) [HTML](0) [PDF 2.56 M](753)

Abstract:
Picea A. Dietrich. (Pinaceae) has a rich fossil pollen record. Macrofossils of the genus known to date are mainly represented by wood and seed cones, while leaves are scarce. Moreover, reported fossil leaves lack microscopic examinations, limiting their taxonomic resolution. In this study, we report a large number of three-dimensional preserved leaves and leaf fragments from the Neogene (latest Miocene to early Pliocene) Yangyi Formation of the Baoshan Basin, western Yunnan Province. These fossil leaves are compressed and flattened. The leaf apex is acuminate and its base is truncated with no petiole. The midvein is obviously raised on both the adaxial and abaxial surfaces. On the adaxial surface, a stomatal band is present on both sides of the midvein, each comprising five or six, mostly continuous stomatal files. The epidermal cells are rectangular to elongate with undulate anticlinal walls. On the abaxial surface, stomata are absent, and the epidermal cells are rectangular to elongate with undulate anticlinal walls. Based on comparisons of gross leaf morphology and micromorphology with extant species of Picea, we found that the fossil leaves are most similar to Picea brachytyla var. complanata (Mast.) W. C. Cheng ex Rehder. Due to the absence of other associated plant organs, it is uncertain if these leaves truly represent this living species. In consequence, we tentatively assign these fossil leaves to Picea brachytyla var. complanata (Mast.) W. C. Cheng ex Rehder. Based on modern altitudinal range (2000–4000 m) of Picea in Yunnan along the southeastern margin of the Tibetan Plateau, the Baoshan Basin might have reached a considerably high elevation, at least higher than the modern level (1650 m), during the latest Miocene and the early Pliocene. Given the warmer climate of that period, Picea may have occupied even higher elevations than their modern counterparts. This finding enriches our knowledge on the elevation history of the southeastern margin of the Tibetan Plateau.

7 Classic formula, updated algorithm and application of rarefaction: bias correction in fossil diversity through subsampling

HUANG Hao , WEN Die , HUANG Han-hui

2023, 62(3):424-435. DOI: 10.19800/j.cnki.aps.2023004 CSTR:

[Abstract](117) [HTML](0) [PDF 1.27 M](670)

Abstract:
Taxonomic diversity of paleocommunities is a key metric for tracing the evolution of life and underlying geological events. However, the taxonomic richness of fossil collections or compiled data is easily biased by differences in sampling size. Rarefaction is a routine statistical method to mitigate such biases by reducing larger collections to a consistent sample size with the smaller ones. Traditional individual-based rarefaction has been increasingly superseded in the literature by coverage-based rarefaction (or SQS, shareholder quorum subsampling as named by some paleontologists). However, some case studies still show certain misunderstanding of this longstanding method, and coverage-based rarefaction has rarely been clarified in the Chinese literature. In order to better apply this method, this paper introduces the principle, details of calculation and suggestions for application of the rarefaction techniques. The core idea of rarefaction is to randomly resample from the original samples until the subsamples reach a consistent sample level, then the mathematical expectation of the taxonomic richness of these subsamples is calculated for comparison. Traditional rarefaction method evaluates such consistency by the same sample size, such as the number of specimens or fossil occurrences in literature. One major drawback of this traditional method is that the information of larger samples is often severely compressed. To address this problem, an updated method, i.e., coverage-based rarefaction, requires resampling until the equal sample coverage is achieved. The degree of coverage is measured by the sum of the individual frequencies in the community covered by the taxa in the subsamples. It has been well demonstrated that the updated method could more faithfully reflect the true ratio of taxonomic richness among communities. Both the traditional and updated rarefaction methods can be implemented by algorithmic simulation or analytical derivation, and software such as PAST or iNext is convenient for implementation. The primary requirement for applying rarefaction is that the samples at hand are as representative of the paleocommunity as possible. We also suggest several potential directions to further develop the rarefaction techniques in the field of quantitative paleontology.

8 Chitinozoan study: review and prospect

LIANG Yan , TANG Peng , Olle HINTS , Jaak N?LVAK , ZHANG Lin-na , CHEN Yan-sen

2023, 62(3):436-450. DOI: 10.19800/j.cnki.aps.2023003 CSTR:

[Abstract](126) [HTML](0) [PDF 4.70 M](676)

Abstract:
Chitinozoans are one of the vital groups of microfossils in the Palaeozoic ocean. Because of their rapid evolution at species level and high abundance in different types of sediments, chitinozoans have been used as one of the most essential and valuable tools in Ordovician and Silurian biostratigraphy. Related research results have provided crucial insights into major geological, environmental, and biological events. However, with the development in modern palaeontology and the accumulation of data during the last decades, chitinozoan study has been experiencing a period of stagnation. First, due to limited techniques applied and lacking standard research criteria in early chitinozoan study, poorly presented images and simple descriptions have resulted in confusion on the taxonomy of some early established chitinozoan taxa, which in turn has a negative influence on the chitinozoan systematics, as well as further study that relies on taxonomy, preventing a better survey of major biological events such as the Great Ordovician Biodiversification Event. Second, digitalisation and visualisation of palaeontological data have become two of the most important and hottest subjects in palaeontology. However, chitinozoan research in this aspect is proceeding quite slowly. Until now, the lack of an efficient database collecting and analysing global chitinozoan data has seriously weakened the usage of the chitinozoan data and has failed to provide data support on the study of chitinozoan macroevolution, biostratigraphy, palaeobiogeography, and palaeoecology. Third, recent studies have provided new insights into the biological affinity of chitinozoans, albeit with some controversies. New evidence is required to further discuss this issue. ＜br＞To effectively facilitate chitinozoan studies, we suggest designing and constructing an open-access chitinozoan database for all the 1311 documented chitinozoan species on the Geobiodiversity Database platform. This database will present combined information such as taxonomy, morphological features, high-resolution images, locality data, formations, ages, section information, and related references. Registered users can search, input, revise, import and analyse the data. In the meantime, revisions on poorly documented chitinozoan records from China and the Baltic will be carried out as a first step to make the data more reliable. Advanced imaging techniques, such as Near-Infrared Microscopy, focused ion beam scanning electron microscopy, field emission scanning electron microscope, and X-ray computed micro-tomography will be applied to update the morphological information of all poorly imaged types, including their interior structures and ultrastructures. Revisions on controversial chitinozoan records from the Ordovician as well as those from the Silurian and the Devonian in other areas will be carried out in the future.
Creating a global chitinozoan database and revising poorly presented chitinozoan records based on extensive data analysis and high-resolution imaging techniques will be a significant advance for the field. This will substantially facilitate the study of the morphology, microevolution, biostratigraphy, palaeogeography and palaeoecology of chitinozoans. This project will bring chitinozoan research into a new era and will be an essential case study to show how far basic research can go with the support of big data and advanced techniques.

Volume 62,Issue 3,2023 Table of Contents

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