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Priapulida Priapulida Aysheaia A Paucipodia B Siberion Microdictyon Tardigrada Diania Antennacanthopodia Antennacanthopodia Onychophora Aysheaia Diania Xenusion Paucipodia Hadranax Microdictyon 100 Orstenotubulus Hadranax Siberion Orstenotubulus Onychophora Xenusion Tardigrada Cardiodictyon Cardiodictyon Hallucigenia Onychodictyon Onychodictyon 55 Hallucigenia Collins’ monster 88 Luolishania Luolishania Collins’ monster Megadictyon Megadictyon Jianshanopodia Jianshanopodia Pambdelurion 65 Kerygmachela Kerygmachela Pambdelurion Anomalocaris 62 62 Opabinia Opabinia Anomalocaris Schinderhannes 83 Schinderhannes Fuxianhuia Fuxianhuia 88 Eoredlichia Eoredlichia 74 Leanchoilia 0.2 changes / character 74 Leanchoilia Tubiluchus (Priapulida) Tubiluchus (Priapulida) Cricocosmia C Cricocosmia D Aysheaia Aysheaia Siberion Onychodictyon gracilis Onychodictyon gracilis Diania Hadranax Xenusion 100 61 Megadictyon Paucipodia Jianshanopodia Microdictyon Onychodictyon ferox Cardiodictyon Siberian Orsten tardigrade Hallucigenia fortis 95 60 93 64 Actinarctus (Heterotardigrada) Hallucigenia sparsa Halobiotus (Eutardigrada) Carbotubulus Kerygmachela Hallucigenia hongmeia Pambdelurion Collins’ monster Emu Bay 64 Opabinia Luolishania 90 Anomalocaris Antennacanthopodia Peytoia Ilyodes 96 69 Hurdia Euperipatoides (Onychophora) 83 Fuxianhuia 50 Chengjiangocaris Onychodictyon ferox 67 Misszhouia Siberian Orsten tardigrade 100 Kuamaia Halobiotus (Eutardigrada) Supella (Hexapoda) Actinarctus (Heterotardigrada) 81 58 Leanchoilia Hadranax 89 Alalcomenaeus Siberion Paucipodia Megadictyon 55 Diania Jianshanopodia Xenusion Kerygmachela Antennacanthopodia 50 Ilyodes Pambdelurion 92 Hurdia Euperipatoides (Onychophora) Anomalocaris Microdictyon Peytoia Cardiodictyon Opabinia 73 Hallucigenia fortis Orstenotubulus Fuxianhuia 75 74 Hallucigenia sparsa Chengjiangocaris 80 Carbotubulus Kuamaia Hallucigenia hongmeia Misszhouia 54 Luolishania Alalcomenaeus 0.1 changes / character Collins’ monster Emu Bay Leanchoilia 98 Supella (Hexapoda) 100 Caenorhabditis Caenorhabditis Priapulus Priapulus E F 95 Hypsibius Hypsibius Milnesium Milnesium 100 Diania Diania Paucipodia Paucipodia 97 Microdictyon Microdictyon 60 Cardiodictyon Cardiodictyon 76 Hallucigenia Hallucigenia 86 Onychodictyon 72 Antennacanthopodia 99 Collins’ monster Helenodora Luolishania 55 Antennacanthopodia Succinipatopsis 91 Helenodora Tertiapatus 96 Succinipatopsis Peripatus 96 Tertiapatus Euperipatoides 97 Euperipatoides Onychodictyon 100 Peripatus Collins’ monster Aysheaia Luolishania Siberion Aysheaia 85 Hadranax Xenusion Siberion 76 Xenusion 96 Jianshanopodia Hadranax Megadictyon 72 Kerygmachela Jianshanopodia 99 Pambdelurion Megadictyon 100 Opabinia Kerygmachela 0.2 changes / character 93 Anomalocaris Pambdelurion 100 Hurdia Opabinia Peytoia 87 56 ARTHROPODA ARTHROPODA.

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Supporting Information Yang et al. 10.1073/pnas.1522434113 SI Text (2) three Character Coding. The dataset used for the phylogenetic analysis has (3) four been updated from that presented by Yang et al. (36), including the (4) six formulation of new neurological characters to resolve large-scale (5) seven relationships within Panarthropoda. The crown-group euarthropods (−) inapplicable: terminal claws (character 64) present. Limulus polyphemus (Chelicerata, Xiphosura) and Triops cancri- State 4 is introduced to reflect the presence of six toe-like formis (Mandibulata, Notostraca) were included as their neuro- claws in the heterotardigrade Batillipes pennaki (28). logical organization has been extensively studied (1–3, 37) and to Cardiovascular and neurological organization. appropriately polarize the characters in the analysis. Additional 81. Dorsal heart. extant taxa were included based on recent studies describing aspects of the brain and VNC organization; these include the eutardigrades (0) absent Macrobiotus cf harmsworthi and Hypsibius dujardini (16, 18), the (1) present heterotardigrades Echiniscus testudo, Actinarctus doryphorus,and Batillipes pennaki (28, 38), the peripatopsid Metaperipatus blainvillei, See character 86 in Yang et al. (36). and the peripatid Epiperipatus biolleyi (12, 13, 15). Fossil and extant 82. Dorsal condensed brain. taxa are scored according to a single model of head segmental (0) absent organization that is informed by developmental studies on extant (1) present Onychophora, Tardigrada, and Euarthropoda (36). Characters 1–80 largely follow those of Yang et al. (36); only See character 82 in Yang et al. (36). those with minor modifications are outlined here. Characters 83. Number of neuromeres integrated into the dorsal condensed brain.
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The Tiger Beetles (Coleoptera, Cicindelidae) of the Southern Levant and Adjacent Territories: from Cybertaxonomy to Conservation Biology
A peer-reviewed open-access journal ZooKeys 734: 43–103 The(2018) tiger beetles( Coleoptera, Cicindelidae) of the southern Levant... 43 doi: 10.3897/zookeys.734.21989 MONOGRAPH http://zookeys.pensoft.net Launched to accelerate biodiversity research The tiger beetles (Coleoptera, Cicindelidae) of the southern Levant and adjacent territories: from cybertaxonomy to conservation biology Thorsten Assmann1, Estève Boutaud1, Jörn Buse2, Jörg Gebert3, Claudia Drees4,5, Ariel-Leib-Leonid Friedman4, Fares Khoury6, Tamar Marcus1, Eylon Orbach7, Ittai Renan4, Constantin Schmidt8, Pascale Zumstein1 1 Institute of Ecology, Leuphana University Lüneburg, Universitätsallee 1, D-21335 Lüneburg, Germany 2 Ecosystem Monitoring, Research and Wildlife Conservation (SB 23 Invertebrates and Biodiversity), Black Forest National Park, Kniebisstraße 67, D-72250 Freudenstadt, Germany 3 Karl-Liebknecht-Straße 73, D-01109 Dresden. Germany 4 Steinhardt Museum of Natural History, Tel Aviv University, Ramat-Aviv, Tel Aviv, IL-69978, Israel 5 Biocentre Grindel, Universität Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany 6 Department of Biology and Biotechnology, American University of Madaba, P.O.Box 2882, Amman, JO-11821, Jordan 7 Remez St. 49, IL-36044 Qiryat Tiv’on, Israel 8 Deichstr. 13, D-21354 Bleckede, Germany Corresponding author: Thorsten Assmann ([email protected]) Academic editor: B. Guéorguiev | Received 1 November 2017 | Accepted 15 January 2018 | Published 5 February 2018 http://zoobank.org/7C3C687B-64BB-42A5-B9E4-EC588BCD52D5 Citation: Assmann T, Boutaud E, Buse J, Gebert J, Drees C, Friedman A-L-L, Khoury F, Marcus T, Orbach E, Renan I, Schmidt C, Zumstein P (2018) The tiger beetles (Coleoptera, Cicindelidae) of the southern Levant and adjacent territories: from cybertaxonomy to conservation biology.
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The Cambrian Explosion: a Big Bang in the Evolution of Animals
The Cambrian Explosion A Big Bang in the Evolution of Animals Very suddenly, and at about the same horizon the world over, life showed up in the rocks with a bang. For most of Earth’s early history, there simply was no fossil record. Only recently have we come to discover otherwise: Life is virtually as old as the planet itself, and even the most ancient sedimentary rocks have yielded fossilized remains of primitive forms of life. NILES ELDREDGE, LIFE PULSE, EPISODES FROM THE STORY OF THE FOSSIL RECORD The Cambrian Explosion: A Big Bang in the Evolution of Animals Our home planet coalesced into a sphere about four-and-a-half-billion years ago, acquired water and carbon about four billion years ago, and less than a billion years later, according to microscopic fossils, organic cells began to show up in that inert matter. Single-celled life had begun. Single cells dominated life on the planet for billions of years before multicellular animals appeared. Fossils from 635,000 million years ago reveal fats that today are only produced by sponges. These biomarkers may be the earliest evidence of multi-cellular animals. Soon after we can see the shadowy impressions of more complex fans and jellies and things with no names that show that animal life was in an experimental phase (called the Ediacran period). Then suddenly, in the relatively short span of about twenty million years (given the usual pace of geologic time), life exploded in a radiation of abundance and diversity that contained the body plans of almost all the animals we know today.
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The Origin and Evolution of Arthropods Graham E
INSIGHT REVIEW NATURE|Vol 457|12 February 2009|doi:10.1038/nature07890 The origin and evolution of arthropods Graham E. Budd1 & Maximilian J. Telford2 The past two decades have witnessed profound changes in our understanding of the evolution of arthropods. Many of these insights derive from the adoption of molecular methods by systematists and developmental biologists, prompting a radical reordering of the relationships among extant arthropod classes and their closest non-arthropod relatives, and shedding light on the developmental basis for the origins of key characteristics. A complementary source of data is the discovery of fossils from several spectacular Cambrian faunas. These fossils form well-characterized groupings, making the broad pattern of Cambrian arthropod systematics increasingly consensual. The arthropods are one of the most familiar and ubiquitous of all ani- Arthropods are monophyletic mal groups. They have far more species than any other phylum, yet Arthropods encompass a great diversity of animal taxa known from the living species are merely the surviving branches of a much greater the Cambrian to the present day. The four living groups — myriapods, diversity of extinct forms. One group of crustacean arthropods, the chelicerates, insects and crustaceans — are known collectively as the barnacles, was studied extensively by Charles Darwin. But the origins Euarthropoda. They are united by a set of distinctive features, most and the evolution of arthropods in general, embedded in what is now notably the clear segmentation of their bodies, a sclerotized cuticle and known as the Cambrian explosion, were a source of considerable con- jointed appendages. Even so, their great diversity has led to consider- cern to him, and he devoted a substantial and anxious section of On able debate over whether they had single (monophyletic) or multiple the Origin of Species1 to discussing this subject: “For instance, I cannot (polyphyletic) origins from a soft-bodied, legless ancestor.
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Fossils from South China Redefine the Ancestral Euarthropod Body Plan Cédric Aria1 , Fangchen Zhao1, Han Zeng1, Jin Guo2 and Maoyan Zhu1,3*
Aria et al. BMC Evolutionary Biology (2020) 20:4 https://doi.org/10.1186/s12862-019-1560-7 RESEARCH ARTICLE Open Access Fossils from South China redefine the ancestral euarthropod body plan Cédric Aria1 , Fangchen Zhao1, Han Zeng1, Jin Guo2 and Maoyan Zhu1,3* Abstract Background: Early Cambrian Lagerstätten from China have greatly enriched our perspective on the early evolution of animals, particularly arthropods. However, recent studies have shown that many of these early fossil arthropods were more derived than previously thought, casting uncertainty on the ancestral euarthropod body plan. In addition, evidence from fossilized neural tissues conflicts with external morphology, in particular regarding the homology of the frontalmost appendage. Results: Here we redescribe the multisegmented megacheirans Fortiforceps and Jianfengia and describe Sklerolibyon maomima gen. et sp. nov., which we place in Jianfengiidae, fam. nov. (in Megacheira, emended). We find that jianfengiids show high morphological diversity among megacheirans, both in trunk ornamentation and head anatomy, which encompasses from 2 to 4 post-frontal appendage pairs. These taxa are also characterized by elongate podomeres likely forming seven-segmented endopods, which were misinterpreted in their original descriptions. Plesiomorphic traits also clarify their connection with more ancestral taxa. The structure and position of the “great appendages” relative to likely sensory antero-medial protrusions, as well as the presence of optic peduncles and sclerites, point to an overall
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The Extent of the Sirius Passet Lagerstätte (Early Cambrian) of North Greenland
The extent of the Sirius Passet Lagerstätte (early Cambrian) of North Greenland JOHN S. PEEL & JON R. INESON Ancillary localities for the Sirius Passet biota (early Cambrian; Cambrian Series 2, Stage 3) are described from the im- mediate vicinity of the main locality on the southern side of Sirius Passet, north-western Peary Land, central North Greenland, where slope mudstones of the Transitional Buen Formation abut against the margin of the Portfjeld Forma- tion carbonate platform. Whilst this geological relationship may extend over more than 500 km east–west across North Greenland, known exposures of the sediments yielding the lagerstätte are restricted to a 1 km long window at the south-western end of Sirius Passet. • Keywords: Early Cambrian, Greenland, lagerstätte. PEEL, J.S. & INESON, J.R. The extent of the Sirius Passet Lagerstätte (early Cambrian) of North Greenland. Bulletin of Geosciences 86(3), 535–543 (4 figures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received March 24, 2011; accepted in revised form July 8, 2011; published online July 28, 2011; issued September 30, 2011. John S. Peel, Department of Earth Sciences (Palaeobiology), Uppsala University, Villavägen 16, SE-75 236 Uppsala, Sweden; [email protected] • Jon R. Ineson, Geological Survey of Denmark and Greenland, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark; [email protected] Almost all of the fossils described from the early Cambrian The first fragmentary fossils from the Sirius Passet Sirius Passet Lagerstätte of northern Peary Land, North Lagerstätte (GGU collection 313035) were collected by Greenland, were collected from a single, west-facing talus A.K.
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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.
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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.
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The Snodgrass Tapes Evolution of the Arthropods Robert Evans Snodgrass Page 1 Figure 1
The Snodgrass Tapes Evolution of the Arthropods The third of three lectures by the insect morphologist Robert Evans Snodgrass delivered to the Department of Entomology at the University of Maryland in 1960. Transcribed, assembled and annotated by Jeffrey W. Shultz Robert Evans Snodgrass Well, the subject today will be the evolution of the arthropods. But, of course, I'll have to admit to begin with that I don't really know the truth of the matter. So, judging from what facts you can get to together... I suppose at the present time that all .... evolution is accepted as a fact by all zoologists. And apparently the fundamentalists have given up trying to do anything about it. Yet it is a theory. And ... But it seems the idea of natural selection well- enough accounts for the physical evolution of animals; that is, certain genes produce the proper variations. But what bothers me about the ... about the evolution of the animals is how did the animal ever become such a com- plex assemblage of chemical substances. I've had a cold, but I guess I can talk through it. Every cell in the body, for example, has to have its own enzymes to do its work it's supposed to do. And all these activities have to be correlated and regulated by hormones, and hormones, again, are just chemical compounds. And, so, it seems to me that that's one of the problems of evolution yet is to find out how all of these chemical substances ever got together in the animal in the proper amount, in the proper places and [how they came] to do the things that they do do...
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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.
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Aysheaia Prolata from the Utah Wheeler Formation (Drumian, Cambrian) Is a Frontal Appendage of the Radiodontan Stanleycaris
Aysheaia prolata from the Utah Wheeler Formation (Drumian, Cambrian) is a frontal appendage of the radiodontan Stanleycaris STEPHEN PATES, ALLISON C. DALEY, and JAVIER ORTEGA-HERNÁNDEZ Pates, S., Daley, A.C., and J. Ortega-Hernández, J. 2017. Aysheaia prolata from the Utah Wheeler Formation (Drumian, Cambrian) is a frontal appendage of the radiodontan Stanleycaris. Acta Palaeontologica Polonica 62 (3): 619–625. Aysheaia prolata, was described as the only lobopodian from the Drumian (Cambrian) Wheeler Formation in Utah, USA, and the sole representative of this genus besides the type species Aysheaia pedunculata, from the Cambrian (Stage 5) Stephen Formation, British Columbia. A redescription of Aysheaia prolata reveals previously overlooked morphological features, including segmental boundaries between putative lobopods, and curved terminal spines on the putative anterior end. These observations undermine lobopodian affinities of Aysheaia prolata, and instead we interpret this specimen as an isolated radiodontan frontal appendage. The presence of 11 podomeres, five of which possess elongate and anteri- orly recurved ventral blades with auxiliary spines, together with shorter robust dorsal spines, identify the specimen as Stanleycaris. This represents the first report of Stanelycaris outside of the Cambrian Stage 5 thin Stephen Formation in British Columbia, expanding its palaeobiogeographic and stratigraphic range. Aysheaia is left as a monotypic genus endemic to the Burgess Shale. The Spence Shale luolishaniid Acinocrinus stichus is currently the only lobopodian known from the Cambrian of Utah. Key words: Euarthropoda, Radiodonta, Hurdiidae, Cambrian, United States. Stephen Pates [[email protected]], Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK. Allison C. Daley [[email protected]], Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015, Lausanne, Switzerland.
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The Palaeoscolecida and the Evolution of the Ecdysozoa Andrey Yu
The Palaeoscolecida and the evolution of the Ecdysozoa Andrey Yu. Zhuravlev1, José Antonio Gámez Vintaned2 and Eladio Liñán1 1Área y Museo de Paleontología, Departamento de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, C/ Pedro Cerbuna, 12, E-50009 Zaragoza, Spain 2Área de Paleontología, Departamento de Geologica, Facultad de Ciencias Biológicas, Univeristat de València, C/ Doctor Moliner, 50, E-46100 Burjassot, Spain Email: [email protected] AbstrAct rÉsUMÉ Palaeoscolecidans are a key group for understanding the ear- Les Paléoscolécides sont un groupe clé pour la compréhen- ly evolution of the Ecdysozoa. The Palaeoscolecida possess sion des débuts de l’évolution des Ecdysozoa. Les Pal- a terminal mouth and an anus, an invertible proboscis with aeoscolecida possèdent une bouche terminale et un anus, pointed scalids, a thick integument of diverse plates, sensory un proboscis inversible aux scalides pointues, un tégument papillae and caudal hooks. These are features that draw a épais de plaques diverses, des papilles sensorielles et des secret out of these worms, indicating palaeoscolecidan af- crochets caudaux. Ceux-ci sont des traits qui tirent un secret finities with the phylum Cephalorhyncha, which embraces de ces vers, ce qui indique des affinités paléoscolecides avec priapulids, kinorhynchs, loriciferans and nematomorphs. At le phylum des Cephalorhyncha qui inclut les priapulides, les the same time, the Palaeoscolecida share a number of char- kinorhynches, les loricifères et les nématomorphes. Cepen- acters with the lobopod-bearing Cambrian ecdysozoans, the dant, les Palaeoscolecida ont aussi quelques-uns des mêmes Xenusia. Xenusians commonly possess a terminal mouth, caractères que les Xenusia, ces écdysozaires cambriens qui a proboscis (although not retractable), and a thick integu- portaient des lobopodes.
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