The earliest belemnite linked with the Carnian Pluvial Episode

Introduction

Belemnites (order Belemnitida), a flourishing group of Mesozoic coleoid cephalopods, appeared in the beginning of the late Triassic (Carnian) and became extinct during the Cretaceous–Paleogene event (Zhu and Bian, 1984; Fuchs and Larson, 2011a; Fuchs and Larson, 2011b; Iba et al., 2011; Iba et al., 2012). They occupied significant niches in oceanic palaeoecosystems, acting both as predators and as important prey for fish, marine reptiles and sharks (Doyle and Macdonald, 1993; Rexfort and Mutterlose, 2006; Hoffmann and Stevens, 2020). Belemnites inhabited shallow to deep waters of epicontinental seas, with a cosmopolitan presence both in the southern and northern hemispheres, extending from subtropical to polar areas (Iba et al., 2014; Dera et al., 2016). Their diversity shifts were closely correlated with environmental changes, including extinction events and climate variations (Dera et al., 2016; Hoffmann and Stevens, 2020; Neige et al., 2021).

The Sinobelemnitidae are considered the oldest belemnite group (Iba et al., 2012; Iba et al., 2014; Hoffmann and Stevens, 2020). Their earliest occurrence is recorded from Carnian strata in South China up to Japan from lower Jurassic strata (Zhu and Bian, 1984; Iba et al., 2012; Iba et al., 2014). Iba et al. (2012, 2014) also reviewed the Sinobelemnitidae along the eastern Tethyan margin, and extended the origin of the belemnites as early as the Late Triassic, challenging the previous views related to their Jurassic origin in the western Tethys (Doyle, 1994; Weis and Delsate, 2006; Iba et al., 2012; Iba et al., 2014). The origin and palaeoecology of belemnites during pre-Jurassic times is still not fully understood. We report a new sinobelemnite species from the Carnian of southwestern China, Sichuanobelus luxiensis sp. nov., a species critical in understanding the early evolution of belemnites, and by extension, the biological shifts of marine ecosystems during the Carnian.

Geological settings

The studied outcrop occurs in the western transitional region between the Yangtze Carbonate Platform and the Nanpanjiang Basin, along the South China Block. During the Permian and Triassic, the Nanpanjiang Basin was represented by a deep-marine embayment, bordered along its western, northern, and northeastern sides by the Yangtze Carbonate Platform, a vast shallow-marine, carbonate platform that stretched across the South China Block (Minzoni et al., 2013). During the Late Triassic, the South China Block was located in eastern Tethys at ∼20°N. The deep-waters of the Nanpanjiang Basin opened to the east into the Panthalassa Ocean (Enos et al., 1998; Figure 1A), with Carnian marine strata well developed and widely exposed in the western transitional region, with continuous sedimentary successions, including the topmost sequences of the Yangliujing, Zhuganpo, Wayao, and Laishike formations respectively (Yunnan Geological Survey, 1975; Guizhou Geological Survey, 2013).

FIGURE 1

Figure 1 (A) Palaeogeographic reconstruction of study area during the Late Triassic interval (modified from Enos et al., 1998). (B) Geological map of the Luxi area and the locality with Sinobelemnites (modified from Yunnan Geological Survey, 1975). (C) Stratigraphic column of the Wayao Formation showing the distribution of Sinobelemnites at localities in the Luxi area.

The belemnites reported herein were collected from the siltic sandstones of the Wayao Formation, Baishui section, Luxi County, Yunnan Province, SW China. The Baishui section is located in the Xing’an village near Baishui town, Luxi County (coordinates: 24.644933°N, 103.828144°E, Figure 1B). The Wayao Formation (also known as the Wayao Member of the Falang Formation or as the Xiaowa Formation) is famous for its Guanling biota, yielding a rich marine reptile fauna and exceptional fossils of pelagic crinoids (Wang et al., 2008). Due to tectonic and depositional factors, the Wayao Formation gradually thickens from southeast to northwest, along the margin of the Nanpanjiang Basin (Minzoni et al., 2015).

At the Baishui section, the Wayao Formation can be divided in two parts:

a. the lower sequence, 3.6 m thick, beginning with black, calcareous shales interlayered with mudstones, with frequent fossils such as ceratitid ammonoids (Trachyceras multituberculatum), bivalves (Halobia sp.), crinoids (Traumatocrinus hsui), and less frequent gastropods (Anulifera yunnanensis);

b. the upper part, dominated by a siltic sandstone, with well preserved fossils such as ceratititd ammonoids (Austrotrachyceras triadicum and Yakutosirenites armiger), bivalves (Halobia sp.), crinoids (Traumatocrinus hsui), belemnites (Sichuanobelus luxiensis n. sp. and Sinobelemnites maantangensis), and gastropods (Anulifera yunnanensis) (Ma et al., 2023).

The new specimens were collected from the base of the upper part, about four meters above lower boundary of the unit (Figure 1C). Based on the occurrence of Austrotrachyceras triadicum and Yakutosirenites armiger, the age of the belemnites is assigned to Carnian (Julian 2).

The Carnian Pluvial Episode in South China

The Carnian Pluvial Episode (CPE), recognized as one of the most significant global climate events during the Triassic period, has garnered considerable attention in the field of geology in recent years (Dal Corso et al., 2015; Jin et al., 2015; Dal Corso et al., 2018; Dal Corso et al., 2020; Zeng et al., 2022). This event caused a substantial amount of rainfall within a short period, effectively ending the long-standing aridity that characterized the Triassic (Zeng et al., 2022). On land, sedimentary characteristics across Europe underwent a transition from widespread evaporitic salt lake and sabkha environments to fluvial and deltaic facies (Kozur and Bachmann, 2010), resulting in the formation of exceptionally large deltas during this time period in locations such as Norway (Klausen et al., 2019). In shallow marine, carbonate growth was interrupted by abundant terrigenous input, leading to a crisis for carbonate factories within the Tethys realm (Hornung and Brandner, 2005; Hornung et al., 2007; Jin, 2019). In deepwater settings, sea level rise triggered extensive deposition of chert limestone followed by a transient increase in the interface depth between chert limestone and carbonate compensation depth (Rigo et al., 2007; Sun et al., 2016). Additionally, the CPE was also a biotic turnover or extinction event (Simms and Ruffell, 1989; Simms and Ruffell, 1990; Simms et al., 1995; Sepkoski, 1996). On the one hand, certain taxa exhibited significant extinction rates, including conodonts, ammonoids, bivalves, crinoids, and bryozoans (Simms and Ruffell, 1989; Simms and Ruffell, 1990; Simms et al., 1995; Bambach, 2006), as well as terrestrial flora and fauna (Benton, 1991; Seyfullah et al., 2018). On the other hand, certain significant taxa emerged during or shortly after the CPE, including mammals, crocodylomorphs, and turtles (Li et al., 2008; Pott et al., 2008; Dal Corso et al., 2020; Funston et al., 2022), while dinosaurs and Rhynchocephalia underwent a radiation event in this interval (Benton et al., 2018; Bernardi et al., 2018; Dal Corso et al., 2020).

CPE has been also well recorded in Yangtze Carbonate Platform and in the Nanpanjiang Basin (Sun et al., 2016; Shi et al., 2018; Jin et al., 2020). The Ma’antang Formation in Sichuan Province, Yangtze Carbonate Platform, and the Wayao Formation, in Guizhou Province, Nanpanjiang Basin, record CPE influences. The Wayao Formation is dominated by black shales and by dark, thin-bedded limestones and mudstones, with rare benthic representatives (Wang et al., 2008; Minzoni et al., 2015; Ma et al., 2018). The lithological facies and absence of a benthic assemblage indicate an anoxic environment during the deposition of the Wayao Formation. Multiple negative carbonate isotope excursions and the lack of carbonate deposition in Wayao Formation were triggered by the CPE (Sun et al., 2016). The Ma’antang Formation yields marine carbonates with at least three siliciclastic interlayers in its lower succession and marine silts and sandstones with bioclastic limestone interlayers to its upper part (Shi et al., 2018). Based on lithological, biostratigraphical and on stable isotopes (Carbon and Oxygen), it was demonstrated that the Ma’antang Formation was deposited during the CPE (Jin et al., 2018; Shi et al., 2018).

Materials and methods

The specimens reported in this paper were collected from the Wayao Formation, preserved in shales and in silty mudstones. All photographs were taken with a Sony ILCE-7M3 digital camera with FE 24–105 mm f/4 G OSS lens. The photographs were processed with image editing software (CorelDRAW 2018 and Adobe Photoshop 2020). All five specimens illustrated and described in this study are deposited in the Southwest Petroleum University (SWPU), Chengdu, Sichuan Province, China, with the inventory numbers SWPU: JS 0001–JS 0005. The morphological terminology follows Hoffmann and Stevens (2020).

Systematic palaeozoology

Order Belemnitida MacGillivray, 1840

Family Sinobelemnitidae Zhu and Bian, 1984

Based on the diagnosis of Sinobelemnitidae made by Zhu and Bian (1984) and by Iba et al. (2015), the surface of Sinobelemnitidae are smooth, lacking any ornaments. However, the surface of specimens from the Luxi country are ornamented, with dense and small knobs. The diameter of a single, small knob is about 10 μm, growing from the rostrum to the proostracum. Besides the Sinobelemnites, similar rostral ornamentation can only be found in some large Megateuthis specimens, but only along lateral rostral sides (Hoffmann and Stevens, 2020). For Megateuthis, two main explanations were given for this type of ornamentation: a. color markings, providing camouflage coloration or muscle attachment structures (Jordan et al., 1975; Spaeth, 1983), and b. these patterns simply reflect a heterogeneous distribution of organic matter within the rostrum (Keupp, 2012; Hoffmann and Stevens, 2020). In Sinobelemnites, the small knobs cover the whole rostrum, densely and uniformly, each small knob being united by three to four minor points. This ornamentation may related to family characters, but it also could serve some special function.

Genus Sichuanobelus Zhu and Bian, 1984

Type species: Sichuanobelus longmenshanensis Zhu and Bian, 1984, from Ma’antang section in Jiangyou, Sichuan Province, China

Sichuanobelus luxiensis sp. nov.

Figure 2A

Diagnosis

Rostrum with a single alveolar groove along the dorsal side, the rostral diameter quickly increasing along the rostrum cavum region. Groove deep and V-shaped. The lateral part of rostrum slightly compressed with no lateral lines, the surface of rostrum with well arranged knobs.

Stratum tipicum

The base of the upper part of the Wayao Formation.

Locus tipicus

Baishui section in Luxi Country, Yunnan Province, Southwest China.

Material

Specimen SWPU: JS 0001 (Figure 2A).

FIGURE 2

Figure 2 The Sinobelemnites from the lower sequence of the Wayao Formation in Luxi, Baishui section. (A) holotype SWPU: JS 0001 of Sichuanobelus luxiensis. (A1) Ventral view. (A2) Dorsal view. (B–E) SWPU: JS 0001–JS 0005, Sinobelemnites maantangensis. (B1) Ventral view. (B2) Dorsal view. (C1) Ventral view. (C2) Dorsal view. (D1) Ventral view. (D2) Dorsal view. (D3) details of the small knobs. (D4) scanning electron micrographs (SEM) of the small knobs covering the belemnite rostra. (E1) longitudinal section showing the organic matter enriched to the primordial rostrum region.

Holotype

SWPU: JS 0001.

Derivatio nominis

luxiensis, from the local name of the study area, Luxi.

Description

The length of the preserved rostra is 23.1 mm, medium in size. The diameter of the rostrum increases quickly along the rostrum cavum region. The largest diameter of the rostrum occurs in the anterior region. The maximum dorsoventral width (Dv) and the lateral width (Dl) are 5.0 mm and 3.0 mm, respectively, with a Dv/Dl ratio of 1.66. The rostra are covered by small knobs which become larger from the posterior area to the anterior area. The average diameter of the knobs is 10 μm. The rostrum apex is missing.

Remarks

Sichuanobelus includes three other species, Sic. longmenshanensis, Sic. yangi and Sic. utatsuensis. The Sic. utatsuensis from the upper Hettangian (Lower Jurassic) of Japan has a shorter rostrum with a very deep groove (Iba et al., 2012), such a feature missing in Sic. luxiensis. Sic. longmenshanensis and Sic. yangi, are both reported from the Carnian of the Ma’antang section in Jiangyou, with Sic. luxiensis n. sp. having a longer and slender rostrum, different from Sic. longmenshanensis and Sic. yangi.

Genus Sinobelemnites Zhu and Bian, 1984.

Sinobelemnites maantangensis Zhu and Bian, 1984

Figures 2B–E

1984 Sinobelemnites maantangensis Zhu and Bian, 1984, p. 307, Figure 1.

Description

The lengths of the preserved rostra vary between 20.0–35.0 mm, middle sized (Figures 2B–E), with missing apices. The largest diameter of the rostrum occurs along the alveolar region. The maximum dorsoventral width (Dv) and the lateral width (Dl) occur in samples JS0002 and JS0003, 15.0 and 12.6 mm, respectively. The Dv/Dl ratio is 1.14–1.19. The rostrum has a deep, singular, alveolar groove along the dorsal side, with two well-developed lateral lines occurring on both lateral rostral sides. All rostra are covered with small knobs which have an average diameter of 20 μm (Figures 2D3, D4). Based on longitudinal sections, the apical angle of the phragmocone is about 22° (Figure 2E1).

Remarks

This species was previously described from the Ma’antang section in Jiangyou County, Sichuan Province, Southwest China (Zhu and Bian, 1984), from the lower part of the Ma’antang Formation (Zhu and Bian, 1984). Our material from Luxi was collected from the base of the upper part of the Wayao Formation. The deep, single, alveolar groove on the dorsal side and the occurrence of lateral lines on both lateral sides of the rostra undoubtedly points to genus Sinobelemnites. The material has affinities with Sin. maantangensis Zhu and Bian 1984, with only one lateral line on each side of the rostrum, a unique character for Sin. maantangensis. The small rostral knobs of Sin. maantangensis are larger than those of Sic. luxiensis sp. nov., a species difference which could also be linked to different ontogenetic stages.

Material

Four specimens (SWPU: JS0002–JS0005, Figures 2B–E).

Discussions

The significance of the Carnian belemnites

The Sinobelemnitidae exhibit a typical belemnite morphology characterized by long rostra and a well-developed alveolar groove. However, belemnites with these characteristics were not found in Europe until the Middle Jurassic, leading to the erroneous identification of their age as belonging to the Middle Jurassic (Doyle, 1993; Doyle, 1994; Weis and Delsate, 2006). Our newly discovered specimens from Luxi county, Yunnan Province co-occurred with Austrotrachyceras triadicum and Yakutosirenites armiger provide strong evidence supporting the age of Chinese sinobelemnitids should be assigned to Carnian (Julian 2) and the Sinobelemniteidae is likely one of the most ancient clades of belemnites. Moreover, the presence of sinobelemnitids in both the northwest and southwest regions of the Yangtze Block suggests a wide distribution of sinobelemnitids in south China during the Carnian. As the Yangtze Block continues to uplift, the receding of water into the Panthalassa Ocean facilitated the migration of sinobelemnitids to the Panthalassa realm and even to the southern hemisphere, where they thrived during the early Early Jurassic (Iba et al., 2012; Iba et al., 2014; Iba et al., 2015; Figure 3).

FIGURE 3

Figure 3 Belemnites and the Carnian Pluvial Episode records from Luxi area, Baishui section, western Nanpanjiang Basin, Ma’antang section, western Sichuan Basin (from Shi et al., 2018) and the Long Chang section (northern Nanpanjiang Basin; from Sun et al., 2016).

The Carnian Pluvial Episode and the early belemnite

The earliest records of Sinobelemnitidae were made by Zhu and Bian (1984) in Jiangyou County, Sichuan Province, with all specimens collected from the middle-lower part of the Ma’antang Formation. Based on the co-occurring ammonoids, the age of the specimens was considered Carnian. Later, the Ma’antang Formation was considered to belong to Julian 2 (Shi et al., 2018; Jin et al., 2020), while the age of the Wayao Formation was considered as Julian 2 (Sun et al., 2016; Zhang et al., 2020). In our study, ammonoids such as Austrotrachyceras triadicum, and Yakutosirenites armiger co-occurred with the newly collected Sinobelemnitidae specimens, supporting the Julian 2 age.

A remarkable fact is that the Carnian belemnites occurred both in the Wayao and Ma’antang formations and the age of the strata coincides with the age of the first time terrigenous input caused by the CPE (Sun et al., 2016; Jin et al., 2018; Shi et al., 2018; Figure 4), clearly pointing to a link between the the Carnian belemnites and the CPE. The heavy rainfall transported a substantial amount of terrigenous sediments into the basin, resulting in seafloor flattening (Dal Corso et al., 2015). This process led to the expansion of the continental shelf and triggered the transition from a deep, inner carbonate platform to a shallow, nearshore sea during the Carnian period. The enlarged continental shelf provides an extensive habitat for beleminites, which thrive in this particular environment (Dzyuba, 2013; Dera et al., 2016). Meanwhile, the anoxic event triggered by the CPE (Sun et al., 2016) resulted in extensive belemnite mortality, as they were rapidly buried in sediment due to precipitation. This process facilitated the preservation of their delicate surface ornamentation on their rostrum. Although the exact explanation for this ornamentation remains unclear, this discovery provides potential evidence regarding the early morphology, ecology, and systematic evolution of belemnites.

FIGURE 4

Figure 4 Global distributions of belemnites in the early phase of their evolution. Global palaeogeographic reconstruction for Late Triassic (220 Ma) is after Blakey (2022). Red represents the age of belemnites assigned to Late Triassic; yellow represents the age of belemnites assigned to Earliest Jurassic; star represents the localities of Sinobelemnildae; square represents the localities of putative putative; circle represents the localities of Belemnitina.

Data availability statement

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author.

Author contributions

TZ, ZM and JZ designed the project. TZ supported the project. ZM wrote the manuscript. MP, TZ and XZ revised the draft. ZM, JC, HL and SL contributed to field work and provided specimens. ZM, JC prepared the figures. ZM, MP, JZ and TZ contributed to the discussion. All authors contributed to the article and approved the submitted version.

Acknowledgments

We are grateful to Ms. Fu Lihong (付丽红) for helping us with figures and her assistance during our field trips. This work is supported by the National Natural Science Foundation of China (No. 41972120; 42172129) and by the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS) (No. 173131).

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Publisher's note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

References

Bambach R. K. (2006). Phanerozoic biodiversity mass extinctions. Annu. Rev. Earth And Planetary Sci. 34, 127–155. doi: 10.1146/annurev.earth.33.092203.122654

CrossRef Full Text | Google Scholar

Benton M. J. (1991). What really happened in the Late Triassic? Hist. Biol. 5, 263–278.

Google Scholar

Benton M. J., Bernardi M., Kinsella C. (2018). The Carnian Pluvial Episode and the origin of dinosaurs. J. Geol. Soc London 175 (6), 1019–1026. doi: 10.1144/jgs2018-049

CrossRef Full Text | Google Scholar

Bernardi M., Gianolla P., Petti F. M., Mietto P., Benton M. J. (2018). Dinosaur diversification linked with the Carnian Pluvial Episode. Nat. Commun. 9, 1499. doi: 10.1038/s41467-018-03996-1

CrossRef Full Text | Google Scholar

Blakey R. (2022). Maps of ancient Earth – 600 Ma to present (Colorado: Plateau Geosystem Inc). Available at: http://deeptimemaps.com/.

Google Scholar

Dal Corso J., Bernardi M., Sun Y., Song H., Seyfullah L. J., Preto N., et al. (2020). Extinction and dawn of the modern world in the Carnian (Late Triassic). Sci. Adv. 6, eaba0099. doi: 10.1126/sciadv.aba0099

CrossRef Full Text | Google Scholar

Dal Corso J., Gianolla P., Newton R. J., Franceschi M., Roghi G., Caggiati M., et al. (2015). Carbon isotope records reveal synchronicity between carbon cycle perturbation and the “Carnian Pluvial Event” in the Tethys realm (Late Triassic). Global Planetary Change 127, 79–90. doi: 10.1016/j.gloplacha.2015.01.013

CrossRef Full Text | Google Scholar

Dal Corso J., Gianolla P., Rigo M., Franceschi M., Roghi G., Mietto P., et al. (2018). Multiple negative carbon-isotope excursions during the Carnian Pluvial Episode (Late Triassic). Earth-Science Rev. 185, 732–750. doi: 10.1016/j.earscirev.2018.07.004

CrossRef Full Text | Google Scholar

Dera G., Toumoulin A., Baets K. D. (2016). Diversity and morphological evolution of Jurassic belemnites from South Germany. Palaeogeography Palaeoclimatology Palaeoecol. 457, 80–97. doi: 10.1016/j.palaeo.2016.05.029

CrossRef Full Text | Google Scholar

Doyle P. (1993). “Mollusca: Cephalopoda (Coleoidea),” in The fossil record 2. Ed. Benton M. J. (London: Chapman and Hall), 229–236.

Google Scholar

Doyle P. (1994). “Aspects of the distribution of Early Jurassic belemnites,” in Palaeopelagos Special Publication. Roma 1, 109–120.

Google Scholar

Doyle P., Macdonald D. I. M. (1993). Belemnite battlefield. Lethaia 26, 65–80. doi: 10.1111/j.1502-3931.1993.tb01513.x

CrossRef Full Text | Google Scholar

Dzyuba O. S. (2013). Belemnites in the Jurassic-Cretaceous boundary interval of the Mauryn’ya and Yatriya River sections, Western Siberia: Biostratigraphic significance and dynamics of taxonomic diversity. Stratigraphy Geological Correlation 21, 189–214. doi: 10.1134/S0869593813020020

CrossRef Full Text | Google Scholar

Enos P., Wei J.-y., Lehrmann D. J. (1998). Death in Guizhou — Late Triassic drowning of the Yangtze carbonate platform. Sedimentary Geology 118, 55–76. doi: 10.1016/S0037-0738(98)00005-0

CrossRef Full Text | Google Scholar

Fuchs D., Larson N. L. (2011a). Diversity, morphology, and phylogeny of coleoid cephalopods from the upper cretaceous plattenkalks of Lebanon–part I: prototeuthidina. J. Paleontology 85, 234–249. doi: 10.1666/10-089.1

CrossRef Full Text | Google Scholar

Fuchs D., Larson N. L. (2011b). Diversity, Morphology, and Phylogeny of Coleoid Cephalopods from the Upper Cretaceous Plattenkalks of Lebanon–Part II: Teudopseina. J. Paleontology 85, 815–834. doi: 10.1666/10-159.1

CrossRef Full Text | Google Scholar

Funston G. F., dePolo P. E., Sliwinski J. T., Dumont M, Shelley S. L., Pichevin L. E., et al. (2022). The origin of placental mammal life histories. Nature 610, 107–111. doi: 10.1038/s41586-022-05150-w

CrossRef Full Text | Google Scholar

Guizhou Geological Survey (2013). Geology special report of the Ministry of Geology and Mineral Resources of People’s Republic of China (PRC) number 21 (Beijing: Geological Publishing House), 179–201.

Google Scholar

Hoffmann R., Stevens K. (2020). The palaeobiology of belemnites – foundation for the interpretation of rostrum geochemistry. Biol. Rev. 95, 94–123. doi: 10.1111/brv.12557

CrossRef Full Text | Google Scholar

Hornung T., Brandner R. (2005). Biochronostratigraphy of the Reingraben Turnover (Hallstatt Facies Belt): Local black shale events controlled by regional tectonics, climatic change and plate tectonics. Facies 51, 460–479. doi: 10.1007/s10347-005-0061-x

CrossRef Full Text | Google Scholar

Hornung T., Krystyn L., Brandner R. (2007). A Tethys-wide mid-Carnian (Upper Triassic) carbonate productivity crisis: Evidence for the Alpine Reingraben Event from Spiti (Indian Himalaya)? J. Asian Earth Sci. 30, 285–302. doi: 10.1016/j.jseaes.2006.10.001

CrossRef Full Text | Google Scholar

Iba Y., Mutterlose J., Tanabe K., Sano S., Misaki A., Terabe K. (2011). Belemnite extinction and the origin of modern cephalopods 35 m.y. prior to the Cretaceous-Paleogene event. Geology 39, 483–486. doi: 10.1130/G31724.1

CrossRef Full Text | Google Scholar

Iba Y., Sano S., Mutterlose J. (2014). The early evolutionary history of belemnites: new data from Japan. PloS One 9 (5), e95632. doi: 10.1371/journal.pone.0095632

CrossRef Full Text | Google Scholar

Iba Y., Sano S., Mutterlose J., Kondo Y., Prefectural F. (2012). Belemnites originated in the Triassic: A new look at an old group. Sci. Rep. Niigata Univ. 29, 95–96. doi: 10.1130/G33402.1

CrossRef Full Text | Google Scholar

Iba Y., Sano S., Rao X., Fuchs D., Chen T., Weis R., et al. (2015). Early Jurassic belemnites from the Gondwana margin of the Southern Hemisphere —Sinemurian record from South Tibet. Gondwana Res. 28, 882–887. doi: 10.1016/j.gr.2014.06.007

CrossRef Full Text | Google Scholar

Jin X. (2019). The Carnian (Late Triassic) extreme climate event: comparison of the Italian tethys and South China geological records (Università degli Studi di Padova, Department of Geosciences), 1–122.

Google Scholar

Jin X., Gianolla P., Shi Z., Franceschi M., Caggiati M., Du Y., et al. (2020). Synchronized changes in shallow water carbonate production during the Carnian Pluvial Episode (Late Triassic) throughout Tethys. Global Planetary Change 184, 103035. doi: 10.1016/j.gloplacha.2019.103035

CrossRef Full Text | Google Scholar

Jin X., Shi Z., Rigo M., Franceschi M., Preto N. (2018). Carbonate platform crisis in the Carnian (Late Triassic) of Hanwang (Sichuan Basin, South China): Insights from conodonts and stable isotope data. J. Asian Earth Sci. 164, 104–124. doi: 10.1016/j.jseaes.2018.06.021

CrossRef Full Text | Google Scholar

Jin X., Shi Z., Wang Y., Duan X., Cheng M. (2015). Mid-carnian (Late triassic) extreme climate event: advances and unsolved problems. Acta Sedimentologica Sin. 33, 105–115. doi: 10.14027/j.cnki.cjxb.2015.01.011

CrossRef Full Text | Google Scholar

Jordan R., Scheuermann L., Spaeth C. (1975). Farbmuster auf jurassischen Belemniten-Rostren. Paläontologische Z. 49, 332–343. doi: 10.1007/BF02987667

CrossRef Full Text | Google Scholar

Keupp H. (2012). Atlas zur Paläopathologie der Cephalopoden. Berliner Paläobiologische Abhandlungen 12, 1–392.

Google Scholar

Klausen T. G., Nyberg B., Helland-Hansen W. (2019). The largest delta plain in Earth’s history. Geology 47, 470–474. doi: 10.1130/G45507.1

CrossRef Full Text | Google Scholar

Kozur H. W., Bachmann G. H. (2010). The middle carnian wet intermezzo of the stuttgart formation (Schilfsandstein), germanic basin. Palaeogeogr. Palaeoclimatol. Palaeoecol. 290, 107–119. doi: 10.1016/j.palaeo.2009.11.004

CrossRef Full Text | Google Scholar

Li C., Wu X., Rieppel O., Wang L., Zhao L. (2008). An ancestral turtle from the Late Triassic of southwestern China. Nature 456, 497–501. doi: 10.1038/nature07533

CrossRef Full Text | Google Scholar

Ma Z., Yu M., Yan B., He B. (2018). Primary discussion of wayao section sedimentary characteristics of falang formation in wusha area of xingyi, guizhou. Guizhou Geology 35, 37–43.

Google Scholar

Ma Z., Zhang T., Chen J., Zeng J., Zhang X., Li H., et al. (2023). Ammonites from the carnian (Upper triassic) xiaowa formation, southeast yunnan province. Acta Palaeontologica Sin. 62 (2), 282–300. doi: 10.19800/j.cnki.aps.2022010

CrossRef Full Text | Google Scholar

MacGillivray W. (1840). A manual of geology (London, Scott: Webster and Geary), 239.

Google Scholar

Minzoni M., Lehrmann D. J., Payne J., Enos P., Yu M., Wei J., et al. (2013). “Deposits, Architecture, and Controls of Carbonate Margin, Slope, and Basinal Settings: SEPM,” in Triassic tank: platform margin and slope architecture in space and time, Nanpanjiang Basin,south China, vol. 105 . Eds. Playton T., Harris M., Verwer K. (Tulsa: Special Publication), 84–113.

Google Scholar

Minzoni M., Lehrmann D. J., Dezoeten E., Enos P., Montgomery P., Berry A. K., et al. (2015). Drowning of the triassic yangtze platform, south China, by tectonic subsidence into toxic deep waters of an anoxic basin. J. Sedimentary Res. 85, 419–444. doi: 10.2110/jsr.2015.32

CrossRef Full Text | Google Scholar

Neige P., Weis R., Fara E. (2021). Ups and downs of belemnite diversity in the Early Jurassic of Western Tethys. Palaeontology 64, 263–283. doi: 10.1111/pala.12522

CrossRef Full Text | Google Scholar

Pott C., Krings M., Kerp H. (2008). The Carnian (Late Triassic) flora from Lunz in Lower Austria: Paleoecological considerations. Palaeoworld 17, 172–182. doi: 10.1016/j.palwor.2008.03.001

CrossRef Full Text | Google Scholar

Rexfort A., Mutterlose J. (2006). Stable isotope records from Sepia officinalis—a key to understanding the ecology of belemnites? Earth Planetary Sci. Lett. 247, 212–221. doi: 10.1016/j.epsl.2006.04.025

CrossRef Full Text | Google Scholar

Rigo M., Preto N., Roghi G., Tateo F., Mietto P. (2007). A rise in the Carbonate Compensation Depth of western Tethys in the Carnian (Late Triassic): Deep-water evidence for the Carnian Pluvial Event. Palaeogeogr. Palaeoclimatol. Palaeoecol. 246, 188–205. doi: 10.1016/j.palaeo.2006.09.013

CrossRef Full Text | Google Scholar

Sepkoski J. J. (1996). “Patterns of Phanerozoic Extinction: a Perspective from Global Data Bases,” in Global Events and Event Stratigraphy in the Phanerozoic. Ed. Walliser O. H. (Berlin, Heidelberg: Springer).

Google Scholar

Seyfullah L. J., Roghi G., Dal Corso J., Schmidt A. R. (2018). The Carnian Pluvial Episode and the first global appearance of amber. J. Geol. Soc Lond 175 (6), 1012–1018. doi: 10.1144/jgs2017-143

CrossRef Full Text | Google Scholar

Shi Z., Jin X., Preto N., Rigo M., Du Y., Han L. (2018). The Carnian Pluvial Episode at Ma’antang, Jiangyou in Upper Yangtze Block, Southwestern China. J. Geological Soc. 176, 197–207. doi: 10.1144/jgs2018-038

CrossRef Full Text | Google Scholar

Simms M. J., Ruffel A. H., Johnson L. A. (1995). “Biotic and climatic changes in the Carnian (Triassic) of Europe and adjacent areas,” in In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods. Eds. Fraser N. C., Sues H. D. (London: Cambridge University Press), 352–365.

Google Scholar

Simms M. J., Ruffell A. H. (1989). Synchroneity of climatic change and extinctions in the Late. Geology 17(3), 265–268. doi: 10.1130/0091-7613(1989)017<0265:SOCCAE>2.3.CO;2

CrossRef Full Text | Google Scholar

Simms M. J., Ruffell A. H. (1990). Climatic and biotic change in the late Triassic. J. Geol. Soc Lond 147 (2): 321–327. doi: 10.1144/gsjgs.147.2.0321

CrossRef Full Text | Google Scholar

Spaeth C. (1983). Ergänzende Beobachtungen zu Farbmustern auf Belemniten-Rostren. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 165, 438–449.

Google Scholar

Sun Y., Wignall P. B., Joachimski M. M., Bond D. P., Grasby S. E., Lai X., et al. (2016). Climate warming, euxinia and carbon isotope perturbations during the Carnian (Triassic) Crisis in South China. Earth Planetary Sci. Lett. 444, 88–100. doi: 10.1016/j.epsl.2016.03.037

CrossRef Full Text | Google Scholar

Wang X., Bachmann G. H., Hagdorn H., Sander P. M., Cuny G., Chen X., et al. (2008). The late Triassic Black Shales of the Guanling area, Guizhou Province, South-West China: a unique marine reptile and pelagic crinoid fossil Lagerstätte. Palaeontology 51, 27–261. doi: 10.1111/j.1475-4983.2007.00735.x

CrossRef Full Text | Google Scholar

Weis R., Delsate D. (2006). The earliest belemnites: New records from the Hettangian of Belgium and Luxembourg. Acta Universitatis CarolinaeGeologica 24, 181–184.

Google Scholar

Yunnan Geological Survey (1975). 1: 200000 Mile Area Regional Geological Survey Report (Beijing: Geology Press), 1–225.

Google Scholar

Zeng J., Zhang T. Y., Ma Z., Li S., Zhang X. (2022). Carnian Pluvial Episode: advances in climate-environment change and marine ecological effects. Acta Geologica Sin. 96, 729–743. doi: 10.19762/j.cnki.dizhixuebao.2022264

CrossRef Full Text | Google Scholar

Zhang Y., Ogg J. G., Franz M., Bachmann G. H., Szurlies M., Röhling H., et al. (2020). Carnian (Late Triassic) magnetostratigraphy from the Germanic Basin allowing global correlation of the Mid-Carnian Episode. Earth Planetary Sci. Lett. 541, 116275. doi: 10.1016/j.epsl.2020.116275

CrossRef Full Text | Google Scholar

Zhu K., Bian Z. (1984). Sinobelemnitidae, a new family of Belemnitida from the Upper Triassic of Longmenshan, Sichuan. Acta Palaeontologica Sin. 23, 300–319. doi: 10.19800/j.cnki.aps.1984.03.005

CrossRef Full Text | Google Scholar