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Tabelliscolex (Cricocosmiidae: Palaeoscolecidomorpha) from the Early Cambrian Chengjiang Biota, and the Evolution of Seriation in Ecdysozoa
Accepted Manuscript Journal of the Geological Society Tabelliscolex (Cricocosmiidae: Palaeoscolecidomorpha) from the early Cambrian Chengjiang Biota, and the evolution of seriation in Ecdysozoa Xiaomei Shi, Richard J. Howard, Gregory D. Edgecombe, Xianguang Hou & Xiaoya Ma DOI: https://doi.org/10.1144/jgs2021-060 To access the most recent version of this article, please click the DOI URL in the line above. When citing this article please include the above DOI. This article is part of the Advances in the Cambrian Explosion collection available at: https://www.lyellcollection.org/cc/advances-cambrian-explosion Received 26 May 2021 Revised 2 August 2021 Accepted 7 August 2021 © 2021 The Author(s). This is an Open Access article distributed under the terms of the Creative Commons Attribution 4.0 License (http://creativecommons.org/licenses/by/4.0/). Published by The Geological Society of London. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics Supplementary material at https://doi.org/10.6084/m9.figshare.c.5551565 Manuscript version: Accepted Manuscript This is a PDF of an unedited manuscript that has been accepted for publication. The manuscript will undergo copyediting, typesetting and correction before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. Although reasonable efforts have been made to obtain all necessary permissions from third parties to include their copyrighted content within this article, their full citation and copyright line may not be present in this Accepted Manuscript version. Before using any content from this article, please refer to the Version of Record once published for full citation and copyright details, as permissions may be required. -
Palaeoecology of the Early Cambrian Sinsk Biota from the Siberian Platform
Palaeogeography, Palaeoclimatology, Palaeoecology 220 (2005) 69–88 www.elsevier.com/locate/palaeo Palaeoecology of the Early Cambrian Sinsk biota from the Siberian Platform Andrey Yu. Ivantsova, Andrey Yu. Zhuravlevb,T, Anton V. Legutaa, Valentin A. Krassilova, Lyudmila M. Melnikovaa, Galina T. Ushatinskayaa aPalaeontological Institute, Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow 117997, Russia bA´rea y Museo de Paleontologı´a, faculdad de Ciences, Universidad de Zaragoza, C/ Pedro Cerbuna, 12, E-50009, Zaragoza, Spain Received 1 February 2002; accepted 15 January 2004 Abstract The Sinsk biota (Early Cambrian, Botoman Stage, Siberian Platform) inhabited an open-marine basin within the photic zone, but in oxygen-depleted bottom waters. Its rapid burial in a fine-grained sediment under anoxic conditions led to the formation of one of the earliest Cambrian Lagerst7tte. All the organisms of the biota were adapted to a life under dysaerobic conditions. It seems possible that the adaptations of many Cambrian organisms, which composed the trophic nucleus of the Sinsk Algal Lens palaeocommunity to low oxygen tensions allowed them to diversify in the earliest Palaeozoic, especially during the Cambrian. Nowadays these groups comprise only a negligible part of communities and usually survive in settings with low levels of competition. Nonetheless, the organization of the Algal Lens palaeocommunity was not simple, it consisted of diverse trophic guilds. The tiering among sessile filter-feeders was well developed with the upper tier at the 50 cm level. In terms of individuals, the community was dominated by sessile filter-feeders, vagrant detritophages, and diverse carnivores/scavengers. The same groups, but in slightly different order, comprised the bulk of the biovolume: vagrant epifaunal and nektobenthic carnivores/ scavengers, sessile filter-feeders, and vagrant detritophages. -
New Evolutionary and Ecological Advances in Deciphering the Cambrian Explosion of Animal Life
Journal of Paleontology, 92(1), 2018, p. 1–2 Copyright © 2018, The Paleontological Society 0022-3360/18/0088-0906 doi: 10.1017/jpa.2017.140 New evolutionary and ecological advances in deciphering the Cambrian explosion of animal life Zhifei Zhang1 and Glenn A. Brock2 1Shaanxi Key Laboratory of Early Life and Environments, State Key Laboratory of Continental Dynamics and Department of Geology, Northwest University, Xi’an, 710069, China 〈[email protected]〉 2Department of Biological Sciences and Marine Research Centre, Macquarie University, Sydney, NSW, 2109, Australia 〈[email protected]〉 The Cambrian explosion represents the most profound animal the body fossil record of ecdysozoans and deuterostomes is very diversification event in Earth history. This astonishing evolu- poorly known during this time, potentially the result of a distinct tionary milieu produced arthropods with complex compound lack of exceptionally preserved faunas in the Terreneuvian eyes (Paterson et al., 2011), burrowing worms (Mángano and (Fortunian and the unnamed Stage 2). However, this taxonomic Buatois, 2017), and a variety of swift predators that could cap- ‘gap’ has been partially filled with the discovery of exceptionally ture and crush prey with tooth-rimmed jaws (Bicknell and well-preserved stem group organisms in the Kuanchuanpu Paterson, 2017). The origin and evolutionary diversification of Formation (Fortunian Stage, ca. 535 Ma) from Ningqiang County, novel animal body plans led directly to increased ecological southern Shaanxi Province of central China. High diversity and complexity, and the roots of present-day biodiversity can be disparity of soft-bodied cnidarians (see Han et al., 2017b) and traced back to this half-billion-year-old evolutionary crucible. -
Ecdysis in a Stem-Group Euarthropod from the Early Cambrian of China Received: 2 November 2018 Jie Yang1,2, Javier Ortega-Hernández 3,4, Harriet B
www.nature.com/scientificreports OPEN Ecdysis in a stem-group euarthropod from the early Cambrian of China Received: 2 November 2018 Jie Yang1,2, Javier Ortega-Hernández 3,4, Harriet B. Drage5,6, Kun-sheng Du1,2 & Accepted: 20 March 2019 Xi-guang Zhang1,2 Published: xx xx xxxx Moulting is a fundamental component of the ecdysozoan life cycle, but the fossil record of this strategy is susceptible to preservation biases, making evidence of ecdysis in soft-bodied organisms extremely rare. Here, we report an exceptional specimen of the fuxianhuiid Alacaris mirabilis preserved in the act of moulting from the Cambrian (Stage 3) Xiaoshiba Lagerstätte, South China. The specimen displays a fattened and wrinkled head shield, inverted overlap of the trunk tergites over the head shield, and duplication of exoskeletal elements including the posterior body margins and telson. We interpret this fossil as a discarded exoskeleton overlying the carcass of an emerging individual. The moulting behaviour of A. mirabilis evokes that of decapods, in which the carapace is separated posteriorly and rotated forward from the body, forming a wide gape for the emerging individual. A. mirabilis illuminates the moult strategy of stem-group Euarthropoda, ofers the stratigraphically and phylogenetically earliest direct evidence of ecdysis within total-group Euarthropoda, and represents one of the oldest examples of this growth strategy in the evolution of Ecdysozoa. Te process of moulting consists of the periodical shedding (i.e. ecdysis) of the cuticular exoskeleton during growth that defnes members of Ecdysozoa1, a megadiverse animal group that includes worm-like organisms with radial mouthparts (Priapulida, Loricifera, Nematoida, Kinorhyncha), as well as more familiar forms with clawed paired appendages (Euarthropoda, Tardigrada, Onychophora). -
An Ordovician Lobopodian from the Soom Shale Lagerstätte, South Africa
[Palaeontology, Vol. 52, Part 3, 2009, pp. 561–567] AN ORDOVICIAN LOBOPODIAN FROM THE SOOM SHALE LAGERSTA¨ TTE, SOUTH AFRICA by ROWAN J. WHITTLE*,à, SARAH E. GABBOTT*, RICHARD J. ALDRIDGE* and JOHANNES THERON *Department of Geology, University of Leicester, Leicester LE1 7RH, UK; e-mails: [email protected]; [email protected] Department of Geology, University of Stellenbosch, Private Bag XI, Stellenbosch 7602, South Africa; e-mail: [email protected] àPresent address: British Antarctic Survey, Madingley Road, Cambridge, CB3 0ET, UK; e-mail: [email protected] Typescript received 18 July 2008; accepted in revised form 27 January 2009 Abstract: The first lobopodian known from the Ordovician and Carboniferous. The new fossil preserves an annulated is described from the Soom Shale Lagersta¨tte, South Africa. trunk, lobopods with clear annulations, and curved claws. It The organism shows features homologous to Palaeozoic mar- represents a rare record of a benthic organism from the ine lobopodians described from the Middle Cambrian Bur- Soom Shale, and demonstrates intermittent water oxygena- gess Shale, the Lower Cambrian Chengjiang biota, the Lower tion during the deposition of the unit. Cambrian Sirius Passet Lagersta¨tte and the Lower Cambrian of the Baltic. The discovery provides a link between marine Key words: Lagersta¨tte, lobopodian, Ordovician, Soom Cambrian lobopodians and younger forms from the Silurian Shale. Cambrian lobopodians are a diverse group showing a in an intracratonic basin with water depths of approxi- great variety of body shape, size and ornamentation. mately 100 m (Gabbott 1999). Dominantly quiet water However, they share a segmented onychophoran-like conditions are indicated by a lack of flow-induced sedi- body, paired soft-skinned annulated lobopods, and in mentary structures and the taphonomy of the fossils. -
Lobopodian Phylogeny Reanalysed
BRIEF COMMUNICATIONS ARISING Phylogenetic position of Diania challenged ARISING FROM J. Liu et al. Nature 470, 526–530 (2011) Liu et al.1 describe a new and remarkable fossil, Diania cactiformis. absent. For example, character 6 (position of frontal appendage) can This animal apparently combined the soft trunk of lobopodians (a only be coded in taxa that possess a frontal appendage (character 5) in group including the extant velvet worms in addition to many the first instance (such that a ‘‘0’’ for character 5 necessitates a ‘‘-’’ for Palaeozoic genera) with the jointed limbs that typify arthropods. character 6). In morphological analyses such as this, inapplicable They go on to promote Diania as the immediate sister group to the states are usually assumed to have no bearing on the analysis, being arthropods, and conjecture that sclerotized and jointed limbs may reconstructed passively in the light of known states. In analyses of therefore have evolved before articulated trunk tergites in the imme- nucleotide data, by contrast, gaps may alternatively be construed as a diate arthropod stem. The data published by Liu et al.1 do not un- fifth and novel state, because shared deletions from some ancestral ambiguously support these conclusions; rather, we believe that Diania sequence may actually be informative. If this assumption is made with probably belongs within an unresolved clade or paraphyletic grade of morphological data, however, all the logically uncodable states in a lobopodians. character are initially assumed to be homologous, and a legitimate Without taking issue with the interpretation of Diania offered by basis for recognizing clades. -
Hallucigenia's Onychophoran-Like Claws
LETTER doi:10.1038/nature13576 Hallucigenia’s onychophoran-like claws and the case for Tactopoda Martin R. Smith1 & Javier Ortega-Herna´ndez1 The Palaeozoic form-taxon Lobopodia encompasses a diverse range of Onychophorans lack armature sclerites, but possess two types of ap- soft-bodied‘leggedworms’ known from exceptionalfossil deposits1–9. pendicular sclerite: paired terminal claws in the walking legs, and den- Although lobopodians occupy a deep phylogenetic position within ticulate jaws within the mouth cavity9,23.AsinH. sparsa, claws in E. Panarthropoda, a shortage of derived characters obscures their evo- kanangrensis exhibit a broad base that narrows to a smooth conical point lutionary relationships with extant phyla (Onychophora, Tardigrada (Fig. 1e–h). Each terminal clawsubtends anangle of130u and comprises and Euarthropoda)2,3,5,10–15. Here we describe a complex feature in two to three constituent elements (Fig. 1e–h). Each smaller element pre- the terminal claws of the mid-Cambrian lobopodian Hallucigenia cisely fills the basal fossa of its container, from which it can be extracted sparsa—their construction from a stack of constituent elements— with careful manipulation (Fig. 1e, g, h and Extended Data Fig. 3a–g). and demonstrate that equivalent elements make up the jaws and claws Each constituent element has a similar morphology and surface orna- of extant Onychophora. A cladistic analysis, informed by develop- ment (Extended Data Fig. 3a–d), even in an abnormal claw where mental data on panarthropod head segmentation, indicates that the element tips are flat instead of pointed (Extended Data Fig. 3h). The stacked sclerite components in these two taxa are homologous— proximal bases of the innermost constituent elements are associated with resolving hallucigeniid lobopodians as stem-group onychophorans. -
Evolutionary History of Life
Evolutionary history of life The evolutionary history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to the present. Earth formed about 4.5 billion years (Ga) ago and evidence suggests life emerged prior to 3.7 Ga.[1][2][3] (Although there is some evidence of life as early as 4.1 to 4.28 Ga, it remains controversial due to the possible non- biological formation of the purported fossils.[1][4][5][6][7]) The similarities among all known present-day species indicate that they have diverged through the process of evolution from a common ancestor.[8] Approximately 1 trillion species currently live on Earth[9] of which only 1.75–1.8 million have been named[10][11] and 1.6 million documented in a central database.[12] These currently living species represent less than one percent of all species that have ever lived on earth.[13][14] The earliest evidence of life comes from biogenic carbon signatures[2][3] and stromatolite fossils[15] discovered in 3.7 billion- Life timeline Ice Ages year-old metasedimentary rocks from western Greenland. In 2015, 0 — Primates Quater nary Flowers ←Earliest apes possible "remains of biotic life" were found in 4.1 billion-year-old P Birds h Mammals [16][17] – Plants Dinosaurs rocks in Western Australia. In March 2017, putative evidence of Karo o a n ← Andean Tetrapoda possibly the oldest forms of life on Earth was reported in the form of -50 0 — e Arthropods Molluscs r ←Cambrian explosion fossilized microorganisms discovered in hydrothermal -
An Armoured Cambrian Lobopodian from China with Arthropod-Like Appendages
LETTER doi:10.1038/nature09704 An armoured Cambrian lobopodian from China with arthropod-like appendages Jianni Liu1,2, Michael Steiner2, Jason A. Dunlop3, Helmut Keupp2, Degan Shu1,4, Qiang Ou4, Jian Han1, Zhifei Zhang1 & Xingliang Zhang1 Cambrian fossil Lagersta¨tten preserving soft-bodied organisms expanded compared to the trunk and shows no sign of becoming have contributed much towards our understanding of metazoan thinner towards the end. No mouth can be observed (Figs 2a–e and origins1–3. Lobopodians are a particularly interesting group that 3). A small projection occurs at the posterior end of the body (Figs 2a–c diversified and flourished in the Cambrian seas. Resembling and 3). The main body trunk is sub-circular in outline and composed ‘worms with legs’, they have long attracted much attention in that of nine segments, each comprising five rows of sub-parallel transverse they may have given rise to both Onychophora (velvet worms)4–6 annulations alternating with five rings of tubercles with tiny spines and Tardigrada (water bears)7,8,aswellastoarthropodsingeneral9–12. (Fig. 1a, b); each with 10–12 tubercles. The tiny spines are not well- Here we describe Diania cactiformis gen. et sp. nov. as an ‘armoured’ preserved, possibly as an artefact of taphonomy, and in most cases only lobopodian from the Chengjiang fossil Lagersta¨tte (Cambrian Stage the tubercles of these spines are observable (Figs 1a, b and 2b–c). 3), Yunnan, southwestern China. Although sharing features with A pair of strongly armoured appendages protrudes from the ven- other typical lobopodians, it is remarkable for possessing robust trolateral region of each segment, each limb attached robustly at its and probably sclerotized appendages, with what appear to be articu- base to the otherwise slender trunk. -
A Cambrian Unarmoured Lobopodian, †Lenisambulatrix Humboldti Gen. Et Sp
www.nature.com/scientificreports OPEN A Cambrian unarmoured lobopodian, †Lenisambulatrix humboldti gen. et sp. nov., Received: 29 January 2018 Accepted: 14 August 2018 compared with new material of Published: xx xx xxxx †Diania cactiformis Qiang Ou 1,2 & Georg Mayer2 Cambrian marine lobopodians are generally considered as predecessors of modern panarthropods (onychophorans, tardigrades, and arthropods). Hence, further study of their morphological diversity and early radiation may enhance our understanding of the ground pattern and evolutionary history of panarthropods. Here, we report a rare lobopodian species, †Lenisambulatrix humboldti gen. et sp. nov. (“Humboldt lobopodian”), from the early Cambrian Chengjiang Lagerstätte and describe new morphological features of †Diania cactiformis, a coeval armoured lobopodian nicknamed “walking cactus”. Both lobopodian species were similar in possessing rather thick, elongate lobopods without terminal claws. However, in contrast to †Diania cactiformis, the body of which was heavily armored with spines, the trunk and limbs of the Humboldt lobopodian were entirely unarmored. Our study augments the morphological diversity of Cambrian lobopodians and presents two evolutionary extremes of cuticular ornamentation: one represented by the Humboldt lobopodian, which was most likely entirely “naked”, the other epitomized by †D. cactiformis, which was highly “armoured”. Lobopodians were marine, caterpillar-like Palaeozoic animals characterized by non-segmented limbs called lobo- pods or lobopodia (singular: lobopodium; from Greek λοβός [lobos], rounded projection or protuberance; and ποδός [podos], foot). Lobopodians originated and rapidly diversifed1 during the Cambrian radiation of meta- zoan body plans and their marine representatives survived at least until the end of the Carboniferous Period2. Lobopodians are generally considered as a paraphyletic assemblage because some were most likely the forerun- ners of modern lobopod-bearing animals3–5, the onychophorans and tardigrades, and their closest relatives, the arthropods. -
Lecture 15: Evolution of the Atmosphere and Biosphere III: the Phanerozoic and The
Lecture 15: Evolution of the Atmosphere and Biosphere III: The Phanerozoic and the EAS 2200 Introduction to the Earth System Cambrian Explosion: 543 Ma. Almost all animal phyla now K-T present extinction 65 Ma Ediacaran and the rise Permo-Triassic of extinction 248 metazoans Ma Ithaca Formation deposited ~370Ma Darwin’s Dilemma “There is another…difculty, which is much more serious. I allude to the manner in which species belonging to several of the main divisions of the animal kingdom suddenly appear in the lowest known fossiliferous rocks. If the theory be true, it is indisputable that before the lowest Cambrian stratum was deposited, long periods elapsed…and that during these vast periods, the world swarmed with living creatures. But to the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods Darwin, Origin of Species, 1859 Rise of Metazoans As the late Proterozoic glaciations end around 630 Ma, nearly 4 billion years of Earth history have passed. Life has been around for the last 3 to 3.5 Ga, and has managed a few tricks. Complex cells (eukaryotes) have evolved, and perhaps a few colonial algae. The Sun has become nearly as bright as it is today and oxygen is reasonably abundant in atmosphere. Still, to an alien spacecraft parked in orbit, the Earth may not have looked much diferent than it did in the Archean - life is dominantly, if not exclusively, still single-celled and microscopic. But that is already beginning to change – and the pace of change will soon become breathtaking. -
“Explosion”: Comparison of Early Cambrian and Present Faunal Communities with Emphasis on Velvet Worms (Onychophora)
Rev. Biol. Trop., 50(2): 823-841, 2002 www.ucr.ac.cr www.ots.ac.cr www.ots.duke.edu Reimpreso de: / Reprinted from: Rev. Biol. Trop., 48(2/3): 333-351, 2000 Disparity, decimation and the Cambrian “explosion”: comparison of early Cambrian and Present faunal communities with emphasis on velvet worms (Onychophora) Julián Monge-Nájera1 and Xianguang Hou2 1 Biología Tropical, Universidad de Costa Rica, San José, Costa Rica. Fax (506) 2075550; [email protected] 2 Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing 210008, China; [email protected] Abstract: The controversy about a Cambrian “explosion” of morphological disparity (followed by decima- tion), cladogenesis and fossilization is of central importance for the history of life. This paper revisits the con- troversy (with emphasis in onychophorans, which include emblematic organisms such as Hallucigenia), pre- sents new data about the Chengjiang (Cambrian of China) faunal community and compares it and the Burgess Shale (Cambrian of Canada) with an ecologically similar but modern tropical marine site where onychopho- rans are absent, and with a modern neotropical terrestrial onychophoran community. Biovolume was estimated from material collected in Costa Rica and morphometric measurements were made on enlarged images of fos- sils. Cambrian tropical mudflats were characterized by the adaptive radiation of two contrasting groups: the va- gile arthropods and the sessile poriferans. Arthropods were later replaced as the dominant benthic taxon by polychaetes. Vagility and the exoskeleton may explain the success of arthropods from the Cambrian to the mo- dern marine and terrestrial communities, both in population and biovolume. Food ecological displacement was apparent in the B.