The evolution and initial rise of pelagic caryocaridids in the Ordovician (2022)

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Earth-Science Reviews Abstract Introduction Section snippets Phylogenetic analyses Research status of systematic classification British Isles Paleogeographical distribution of caryocaridids Discussion Conclusions Credit author statement Declaration of Competing Interest Acknowledgments References (223) Palaeogeogr. Palaeoclimatol. Palaeoecol. Palaeogeogr. Palaeoclimatol. Palaeoecol. Sediment. Geol. Earth-Sci. Rev. Earth-Sci. Rev. Gondwana Res. Palaeogeogr. Palaeoclimatol. Palaeoecol. Filocáridos (Crustacea) en el Tremadociano del Sistema de Famatina, Provincia de La Rioja, Argentina El Ordovícico del río La Alumbrera, Departamento Tinogasta, Provincia de Catamarca Ameghiniana Chapter 20 A synopsis of Ordovician trilobite distribution and diversity Geol. Soc. London Mem. The apparatusarchitecture and function of Promissum pulchrum Kovacs-Endrody (Conodonta, Upper Ordovician), and the prioniodontidplan Philos. Trans. R. Soc B Ecospace utilization and guilds in marine communities through the Phanerozoic Autecology and the filling of ecospace: key metazoan radiations Palaeontol. Was there an ordovician superplume event? Protocimex: a phyllocarid crustacean, not an Ordovician insect J. Paleontol. Report on the Cambrian and Ordovician formations of Maryland Md. Geol. Surv. Special Paper Parasitism on graptoloid graptolites Palaeontol. Vorpommern und angrenzende Ostsee (Ruögen–Kaledoniden) The ordovician rocks of New Zealand Geol. Mag. The Ordovician graptolites of north-west Nelson, NZ, second paper; with notes on other Ordovician fossils Trans. R. Soc. N. Z. Morphology, palaeobiology and phylogeny of Oryctocaris balssi gen. nov. (Arthropoda), a phyllocarid from the Lower Devonian Hunsrück Slate (Germany) J Syst. Palaeontol. The Almelund Shale, a replacement name for the Upper Didymograptus Shale and the Lower Dicellograptus Shale in the lithostratigraphical classification of the Ordovician succession in Scania, Southern Sweden Bull. Geol. Soc. Den. Eurypterids, phyllocarids, and ostracodes Los fosiles guías de Bolivia I. Paleozoico. Ser. Geol. Bolivia Extinction and the fossil record of the arthropods Bivalved arthropods from the Middle Ordovician Winneshiek Lagerstätte, Iowa, USA J. Paleontol. Decoupled evolution of soft and hard substrate communities during the Cambrian explosion and Great Ordovician Biodiversification Event Proc. Natl. Acad. Sci. U. S. A. South American graptolites with special references to the Nordenskiöld collection Arkiv för Zoologi Evolution of shallow-water level-bottom communities Ecology and evolution of the Cambrian plankton Invertebrados Fósiles XII.—New or Little-known Victorian Fossils in the National Museum, Melbourne. Part I. — Some Paleozoic Species R. Soc. Victoria On some phyllocarids from the Ordovician of Preservation Inlet and Cape Providence, New Zealand Trans. Proc. R. Soc. N. Z. Larval ecology, life history strategies, and patterns of extinction and survivorship among Ordovician trilobites Paleobiology Arenig to Llanvirn Graptolite provincialism of South China Ordovician graptolite evolutionary radiation: a review Geol. J. Phyllocarid crustaceans of the Bohemian Ordovician Sborník geologických věd, Paleontologie Phyllocarid crustaceans from the Middle Ordovician Šárka Formation at Praha-Vokovice Bull. Geosci. Morphology and stratigraphic range of the phyllocarid crustacean Caryocaris from Alaska and the Great Basin Palaeontol. Notes on Paleozoic Crustaceans, 2, Phyllocarida from the black shales at the base of the Salina beds in western New York. 3, Some Devonic Phyllocarida from New York Über den Organismus der Nebaliden und die systematische Stellung der Leptostraken Arbeiten aus dem zoologische Institut der Universität Wien und der zoologischen Station in Triest Lower Palaeozoic facies and faunas around Gondwana Early ordovician brachiopods from south-west Wales Proc. Geol. Assoc. Early evolution of phyllocarid arthropods: phylogeny and systematics of cambrian-devonian archaeostracans J. Paleontol. Three-dimensionally preserved arthropods from Cambrian Lagerstätten of Quebec and Wisconsin J. Paleontol. Sequence and correlation of Tremadoc graptolite assemblages Alcheringa Towards a general model for the depth ecology of graptolites Ecostratigraphy, zonation and global correlation of earliest Ordovician planktic graptolites Lethaia Latitudinal and depth zonation of early Ordovician graptolites Lethaia Cited by (0) Recommended articles (6)
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Earth-Science Reviews

Volume 231,

August 2022

, 104097

Abstract

The ‘Great Ordovician Biodiversification Event’ represents the rapid diversification of marine organisms during the Middle Ordovician. As the initial stage of the ‘Great Ordovician Biodiversification Event’, the ‘Ordovician Plankton Revolution’ played a crucial role in the increase of biodiversity in the Ordovician. The occurrence, phylogeny, diversity and abundance of caryocaridids, an important and representative group of planktonic arthropods in the Ordovician, may provide important information for studying the ‘Ordovician Plankton Revolution’ and the evolution of pelagic food webs. Based on systematic collection of paleontological data of caryocaridids, combined with phylogenetic methods, we determined the taxonomic position of caryocaridids, discussed the evolutionary path of the group, reconstructed the biological migration process of caryocaridids and its paleoecological and paleogeographical setting. Our results show that caryocaridids were a clade of planktonic arthropods, which are commonly preserved in black shales along with graptolites and pelagic trilobites. The caryocaridids evolved from the late Cambrian phyllocarids. They first appeared in the Tremadocian, rapidly diversified in the Floian to Darriwilian, then declined in the Late Ordovician, and went extinct at the end of the Ordovician. From the late Cambrian to the Early Ordovician, a large number of marine organisms switched to the planktonic mode. The emergence, rise and diversification of planktonic caryocaridids provide new evidence for the diversity of marine plankton, the complexity of the Ordovician pelagic food webs, and the occurrence of the ‘Ordovician Plankton Revolution’.

Introduction

As a representative of Malacostraca (Arthropoda), the Phyllocarida has significance in the early evolution of crustaceans due to their morphological diversity, extensive and prolonged fossil records, and extant relatives (Rolfe, 1969, Rolfe, 1981; Schram and Hof, 1998; Spears and Abele, 1998, Spears and Abele, 1999). Currently, phyllocarids are divided into two classes, i.e., the extinct Archaeostraca and the living Leptostraca. The main differences between them are that the former has a carapace with hinge line and fewer appendant limbs (Collette and Hagadorn, 2010a; Rode and Liebermann, 2002; Rolfe, 1969). The oldest known phyllocarid fossil is Arenosicaris inflataCollette and Hagadorn, 2010b from the Cambrian Furongian in Wisconsin, USA, and the youngest is Rhabdouraea bentzi (Malzahn, 1958) from the Upper Permian in western Germany (Glaessner and Malzahn, 1962; Schram and Malzahn, 1984). The Archaeostraca is currently divided into six suborders: Caryocaridina Collette and Hagadorn, 2010a, Ceratiocarina Clarke, 1900, Echinocaridina Clarke, 1900, Palaeopemphida Feldmann et al., 2004, Pephricaridina Van Straelen, 1933, and Rhinocarina Clarke, 1900, among which the Caryocaridina and Ceratiocarina are relatively ancient groups in the Archaeostraca.

Caryocaridids are bivalved arthropods distributed globally in the Ordovician, which are often preserved in graptolite shales as isolated carapaces, tail-piece, or other exoskeleton fragments (Vannier et al., 2003). They are sometimes misdescribed or mistaken for other animal fragments due to the poor preservation and incomplete original descriptions (Aceñolaza and Esteban, 1996; Gurley, 1896). The caryocaridid individuals are relatively small (length about 10-30 mm) and have a pair of slender carapaces with a dorsal hinge line, and the posterior margin of the carapace is spiny (Fig. 1A, B, E, H). Their tail-piece is composed of a conical telson and a pair of furcal rami with triangular expansions and articulated spines and/or setae (Fig. 1C, D, F, G, I). Although the caryocaridis were widespread in the Ordovician (Aceñolaza and Esteban, 1996; Bassler, 1919; Braddy et al., 2004; Bulman, 1931; Chapman, 1902, Chapman, 1934; Gurley, 1896; Hicks, 1876; Jell, 1980; Racheboeuf and Crasquin, 2010; Racheboeuf et al., 2000, Racheboeuf et al., 2009; Ruedemann, 1935; Shen, 1986; Whittle et al., 2007; Woodward, 1912), unfortunately, no complete specimens of caryocaridids have been found so far. Based on these disarticulated fragments, the paleobiogeography (Vannier et al., 2003; Whittle et al., 2007), paleoecology (Chlupáč, 1970, Chlupáč, 2003; Vannier et al., 2003; Whittle et al., 2007), and phylogeny (Collette and Hagadorn, 2010a; Rolfe, 1969) of the caryocaridids have been studied. However, with the wide application of new technologies and methods, the related contents of systematics, biostratigraphy and paleobiogeography need to be updated and improved (Collette and Hagadorn, 2010a; Racheboeuf and Crasquin, 2010; Racheboeuf and Gourvennec, 2013; Racheboeuf et al., 2009).

Living phyllocarids are mainly benthos in shallow water environments (Haney and Martin, 2016), but a few species can reach deeper water or even deep-sea environments (Haney et al., 2001; Hessler, 1984; Mauchline and Gage, 1983; Petryashov, 2016). The earliest phyllocarids lived in the shallow-water shelf to intertidal environments at tropical low latitudes in the late Cambrian (Collette and Hagadorn, 2010b), and then they migrated to the outer shelf and slope environments during the Ordovician (Vannier et al., 2003). The phyllocarids have an extremely low diversity in the late Cambrian, and began to rapidly diversify in the Tremadocian. Among them, the caryocaridids, which mainly lived in the epipelagic and/or mesopelagic environment, peaked in diversity in the Floian–Darriwilian (Chlupáč, 2003; Vannier et al., 2003; Whittle et al., 2007), declined sharply during the Late Ordovician (Sandbian–Hirnantian), and eventually died out in the mass extinction at the end of the Ordovician. Other phyllocarids continued to flourish after the Ordovician (Collette and Hagadorn, 2010a; Rode and Liebermann, 2002; Vannier et al., 2003). In the Ordovician, the global biodiversity, the complexity of the global ecosystem, and the utilization of the ecological space increased significantly (Bambach, 1983; Bambach et al., 2007; Deng et al., 2021; Harper, 2006; Servais and Harper, 2018; Servais et al., 2008, Servais et al., 2010, Servais et al., 2016). However, the expansion of the living environment of caryocaridids during this special interval and its response to the ‘Ordovician Plankton Revolution’ in the ‘Great Ordovician Biodiversification Event’ (GOBE), and the evolutional trend of its diversity have not been systematically studied. In this paper, we collected all published literature on caryocaridids, systematically reviewed their species classification, stratigraphic distribution, and paleobiogeography. In addition, the response patterns of caryocaridids and other plankton to the ‘Ordovician Plankton Revolution’ in the background of GOBE are discussed, and the pelagic food webs of planktonic and free-swimming animals including caryocaridids are established.

Section snippets

Phylogenetic analyses

Initially, we intended to use the characters encoded by Collette and Hagadorn (2010a) for phylogenetic analysis of Caryocaridina, but some of their characters, such as the carapace ornamentation, the ridges of the carapace, and the rostrum are not relevant for such analysis. Therefore, a new analysis (Supplementary material, File A) was performed using 43 characters, partly building on those of Collette and Hagadorn (2010a) and Bergmann and Rust (2014). Some new characters have been added in

Research status of systematic classification

The systematic classification of fossil phyllocarids is different from that of living phyllocarids. Because the soft-body parts (e.g., abdomen and appendages) are rarely preserved, the classification of fossil phyllocarids is mainly based on the morphology and ornamentation of carapace, telson, and furcal rami. This has also resulted in a small number of species being misclassified due to the transitional morphology or poor preservation. For example, Caryocaris acuta (Bulman, 1931) has been

British Isles

Lake District, Northern England The type species Caryocaris wrightiiSalter, 1863 was originally described by Salter from the Skiddaw Group (= the Skiddaw Slates) in the Lake District of northern England (loc. 1) (Jones and Woodward, 1892). According to the classification of lithostratigraphy and biozones given by Cooper et al. (1995), they were mainly distributed in the Kirkstile Formation (gibberulus and hirundo graptolite biozones), the Loweswater and Hope Beck formations (the upper

Paleogeographical distribution of caryocaridids

Except for isolated plates such as the Siberia, Tarim and the North China Block, caryocaridids are widely distributed around the margins of continental blocks on both sides of the Panthalassic, Iapetus, and Rheic oceans (Fig. 6). According to the morphological characteristics and ultramicrostructure of carapaces, abdominal morphology, and other co-preserved fossils, caryocaridids have long been regarded as planktonic animals (Chlupáč, 1970, Chlupáč, 2003; Ruedemann, 1934; Størmer, 1937; Vannier

Discussion

The GOBE is a unique and major event in the evolutionary history of the marine biosphere and corresponds to an exceptionally rapid increase in the diversity of marine organisms at lower taxonomic levels (e.g., genus and family) (Servais and Harper, 2018; Webby, 2004). It records an explosion of adaptive radiations in all the components of the marine ecosystem, from plankton to benthos, as well as reefs (Servais and Harper, 2018). Servais et al. (2008) pointed out that the GOBE represented an

Conclusions

(1)

The fossil record shows that caryocaridids first appeared in the middle Tremadocian, and their diversity and abundance increased rapidly in the Floian to Darriwilian, with three small peaks. In the Late Ordovician, the diversity of caryocaridids began to decline rapidly, and they were all extinct by the end of the Ordovician.

(2)

Based on the morphological characteristics of phyllocarids in the Paleozoic, the phylogeny of caryocaridids was analyzed and reviewed. The phylogenetic tree showed that

Credit author statement

Yilong Liu: Conceptualization, methodology, software and original draft preparation. Ruoying Fan: Writing-review and editing. Ruiwen Zong: Writing-review and editing, Conceptualization, Project administration, Resources. Yiming Gong: Writing-review and editing. All authors discussed the content, reviewed, edited the manuscript and ultimately agreed to the submitted version of the manuscript.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgments

Thanks go to Jiayi Yin (China University of Geosciences, Wuhan) for her assistance in the discussion part. This work was supported by the National Natural Science Foundation of China [grant numbers 42072041, 41902002, 41872034] and the Independent Innovation Funding Program For Undergraduate of China University of Geosciences (Wuhan). We would also like to thank Thomas Servais and one anonymous reviewer for their constructive comments on the manuscript.

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      The southeast Brazilian rifted continental margin is not a single, continuous upwarp: Variations in morphology and denudation patterns along the continental drainage divide

      Earth-Science Reviews, Volume 231, 2022, Article 104091

      Rifted continental margins (RCM) are large-scale features of Earth's surface that show substantial morphological variations. Classical escarpment features are the subject of many studies in these settings while other morphologies that characterize this tectonic environment receive less attention. The case of the Brazilian South Atlantic margin, a continental-scale topographically pronounced terrain covering >1000km of the western South Atlantic rifted margin, is not an exception. Most landscape evolution studies concentrate on the Serra do Mar escarpment system, while other segments with no escarpments have received less attention. Various authors assume the Brazilian elevated continental margin as a well-defined and continuous marginal upwarp instead of a diversified and more complex landscape. Here we debate this issue and explore how the first-order topographic forms and time-space denudation patterns differ along the Brazilian South Atlantic margin. We focus on the continental margin of southeast Brazil (CMSEB) that includes a southern segment featured by a prominent escarpment system and a northern segment where the seaward-facing steep escarpment is absent. We show that, similar to other RCMs, the CMSEB presents a continental drainage divide separating two distinct regions with contrasting denudation patterns, (i) an inland continental interior, characterized by high elevation and relatively low relief with a predominance of apatite fission track (AFT) ages significantly older than South Atlantic rifting event, and (ii) a coastal region characterized by low elevations and high relief with the predominance AFT ages younger than the rifting. However, besides the differential denudation associated with the South Atlantic opening, the margin has experienced substantial post-rift exhumation attributed to rock uplift triggered by the reactivation of inherited basement structures. The morphological differences between CMSEB's southern and northern extensions reflect sectors with contrasting geomorphic evolution supporting the idea that the rifted Brazilian RCM is not a single and continuous tectonic setting. Finally, our findings indicate that tectonic inheritance strongly impacts the denudation pattern, which contributes to the geomorphic diversification along the Brazilian RCM.

    • Research article

      Links of high velocity anomalies in the mantle to the Proto-South China Sea slabs: Tomography-based review and perspective

      Earth-Science Reviews, Volume 231, 2022, Article 104074

      The high velocity anomalies in global tomographic images indicate clearly the present large-scale subducting slabs and already-subducted old oceanic lithosphere in the deeper mantle. The subducted lithosphere can be identified by tracking the high velocity anomalies in the upper and lower mantle. We review several previous kinematic plate reconstruction models involved in the evolution of the Proto-South China Sea and point out challenges of the plate reconstruction for the plate-mantle system of the Proto-South China Sea. We used four global P-wave tomographic models and one global S-wave tomographic model in our analyses and have summarized model parameters and methodologies. We used the average P-wave velocity perturbation variations from four global P-wave tomographic models to constrain the distribution of the Proto-South China Sea lithosphere in the different horizontal slice depths. The north and south slabs of the Proto-South China Sea in the mantle were identified and interpreted from the different latitudinal and longitudinal direction vertical cross-sections from the MIT-P08 tomography model. We then discuss the repeatability of the Proto-South China Sea slabs in different global tomographic models and construct one slab subduction and evolution model of the Proto-South China Sea. The north slab of the Proto-South China Sea is one flat-lying slab at depths of ~400–700km in the mantle transition zone and the south slab of the Proto-South China Sea is one detached slab at depths of ~800–1600km same as the Kalimantan anomaly identified by the previous tomographic images. Tearing and detachment of the Proto-South China Sea subducted oceanic lithosphere, could explain the distribution of the north slab and south slab in the mantle by either a double-sided subduction or single-sided subduction model. The plate-mantle system and the slab configuration involved in the Proto-South China Sea are complex and detached slabs of the Proto-South China Sea in the lower mantle requires the further high resolution local tomographic observations beneath Borneo.

    • Research article

      Tectono-sedimentary evolution of Southern Mexico. Implications for Cretaceous and younger source-to-sink systems in the Mexican foreland basins and the Gulf of Mexico

      Earth-Science Reviews, Volume 231, 2022, Article 104066

      An extensive dataset of existing and new geo/thermochronological data from several areas in Southern Mexico constrains the tectonic history of the region, as well as various source-to-sink relationships and local burial histories. Our interpretation acknowledges that not all cooling/heating observed in the source areas is due to erosional exhumation/burial but, in some cases, due to advective heat transfer from magmatic sources, which potentially overprinted earlier events. In this work, we identified several areas that have been exhumed since the Early Cretaceous and potentially provided clastic material to the southern Gulf of Mexico area.

      We help to document how the Mexican (Laramide) Orogeny propagated eastwards and southwards from the Late Cretaceous through the early Oligocene. The first sediments reaching the Tampico–Misantla and Veracruz basins derived mostly from eroded Cretaceous carbonate material that covered the Sierra Madre Oriental, the Sierra de Juárez Complex and the Cuicateco belts, as well as foredeep/intra-orogenic basin deposits formerly covering them. Possibly by the end of the Mexican Orogeny, the clastic Jurassic and older crystalline basement rocks became exposed and became the main sources of quartz-rich clastic material to the most easterly foreland basins and Gulf of Mexico. Exposure was probably assisted by higher angle basement thrusts such as the Vista Hermosa/Valle Nacional faults. The Mixtequita and Guichicovi blocks have also provided an important source of quartz-rich and metamorphic lithic-rich material to the southern Veracruz Basin possibly since the Eocene.

      For most of the Cenozoic, the Chiapas and the Sureste basins were sourced from areas south of the Chiapas Massif, i.e., the North America–Caribbean plate boundary zone along today's Chiapas coastal plain. This plate boundary zone accommodated relative displacement between Mexico and the Chortis Block of the Caribbean Plate. Paleocene–middle Miocene sediments within the Chiapas Basin were at least partially sourced from i) metamorphic complexes in the northern Chortis Block; ii) the parautochthonous Chontal Complex, an oceanic-like basin sandwiched between Chortis and southern Mexico; iii) the elongating volcanic arc along southern Mexico and western Chortis; and iv) the Cretaceous and Jurassic sedimentary cover of the southern flank of the Chiapas Massif,

      The westward telescoping of southern Mexico onto the Cocos Plate in the wake of Chortis has produced flat slab subduction geometry and eastwardly-younging uplift of the Xolapa Belt (Oligo–Miocene) and the Chiapas Massif (late Miocene). It also caused reorganization of the drainage systems providing material to the Chiapas and Sureste basins.

      Our results highlight the importance of understanding relative block and plate boundary displacements in a dynamic hinterland and consider the role of major faults when interpreting source-to-sink relationships in the area. We describe the latter relationships for several geologic time intervals in which reservoir-prone sediments were delivered to the southern Gulf of Mexico. Finally, we integrate the source-to-sink history to provide an assessment of reservoir quality and hydrocarbon prospectivity in the region.

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