Comparative Analysis of Acoel Embryonic Development

Total Page:16

File Type:pdf, Size:1020Kb

Comparative Analysis of Acoel Embryonic Development Comparative analysis of acoel embryonic development Viviana Cetrangolo Thesis for the degree of Philosophiae Doctor (PhD) University of Bergen, Norway 2020 Comparative analysis of acoel embryonic development Viviana Cetrangolo ThesisAvhandling for the for degree graden of philosophiaePhilosophiae doctorDoctor (ph.d (PhD). ) atved the Universitetet University of i BergenBergen Date of defense:2017 02.10.2020 Dato for disputas: 1111 © Copyright Viviana Cetrangolo The material in this publication is covered by the provisions of the Copyright Act. Year: 2020 Title: Comparative analysis of acoel embryonic development Name: Viviana Cetrangolo Print: Skipnes Kommunikasjon / University of Bergen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`1/-"0!&'(E+;!-$/-!2/0)!/!-$('1A$!A1-!/,.!$/C%!/!+",A2%! 3'.4! '*%,",A! -'! -$%"(! ."A%+-"C%! +4,04-"1E7! F2-$'1A$! -$%4! &%(%! ","-"/224! *2/0%.! &"-$",! +*"(/2"/,+;!3%0/1+%!'5!-$%"(!E'(*$'2'A"0/2!/55","-"%+!&"-$!-1(3%22/("/,+;!/0'%2+;!-'A%-$%(!&"-$! ,%E%(-'.%(E/-".+!/,.!5(1#)-*6(..&!Za%,/0'%2'E'(*$/\;!/(%!,'&!*2/0%.!/+!+"+-%(!A('1*!'5! /22!-$%!(%E/",",A!3"2/-%("/,+!ZD%*$('_'/!b!6('-'+-'E"/!c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de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f7;!?/(-",./2%!?7Y7;!<%/C%(!U7;!<0$,/3%2!K7;!9%:,'2!F7!1/$'2".$'3"#' ")-' %$4,+".(5$' -$5$+6#3$).' ")"+7&$&' 62' ./$' "*6$+' !"#$%&'()*&+ ,-./0*&8' 9?/,1+0("*-!",!*(%*/(/-"',\' 6/*%(!==X!>%-(/,A'2'!f7;!9%:,'2!F7!:"%+7';+"&.63$%$'(-$).(.7'%$5$"+$-';7'4$)$'$<#%$&&(6)' ()'./$'"*6$+'1#23#.-)*%.#4&+'&/*#,56&7+Z?/,1+0("*-!",!*(%*/(/-"',\+ 81(",A!E4!-(/",",A!=!0',-("31-%.!-'!-$%!*/*%(X!! ?/(-M,N81(O,!^7?7;!f%221-","!@7>7;!?/(2g-/_!G7;!>%-(/,A'2'!f7=!>C%-%+"0!D7;!#$"%2!87;! 9%,("%-!<7;!L(/1N@'Cg!a7;!>/(("22'N@/2-'./,'!F7;![%,:"/!L7;!S%(32!F7;!@%))'10$%!D;! >$'1(('1-!87;!LhE%_N<)/(E%-/!^7T7;!=("E"/!?7;!T%,$/(.-!@7;!['(+//%!S7;!9%:,'2!F7! >6)&$%5".(5$' %6,.$' .6' $<.%$3$' 4$)63(*' *63#"*.(6)' ()' "' 3()(".,%$' "))$+(-8' Z?/,1+0("*-!+13E"--%.\! ! ! ! ! ! C"""! :+&'%&'3! ! -!./$,01)#$'0/!2222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222222!-! !"!#$%&#&'()*+#,-.#/01#.&2&3+4'&-05#%+6#0+#7+)'#,#-&6#+)8,-/1'##""""""""""""""""""""""""""""""""""""""""""""""""""""""#!#
Recommended publications
  • Xenacoelomorpha's Significance for Understanding Bilaterian Evolution
    Available online at www.sciencedirect.com ScienceDirect Xenacoelomorpha’s significance for understanding bilaterian evolution Andreas Hejnol and Kevin Pang The Xenacoelomorpha, with its phylogenetic position as sister biology models are the fruitfly Drosophila melanogaster and group of the Nephrozoa (Protostomia + Deuterostomia), plays the nematode Caenorhabditis elegans, in which basic prin- a key-role in understanding the evolution of bilaterian cell types ciples of developmental processes have been studied in and organ systems. Current studies of the morphological and great detail. It might be because the field of evolutionary developmental diversity of this group allow us to trace the developmental biology — EvoDevo — has its origin in evolution of different organ systems within the group and to developmental biology and not evolutionary biology that reconstruct characters of the most recent common ancestor of species under investigation are often called ‘model spe- Xenacoelomorpha. The disparity of the clade shows that there cies’. Criteria for selected representative species are cannot be a single xenacoelomorph ‘model’ species and primarily the ease of access to collected material and strategic sampling is essential for understanding the evolution their ability to be cultivated in the lab [1]. In some cases, of major traits. With this strategy, fundamental insights into the a supposedly larger number of ancestral characters or a evolution of molecular mechanisms and their role in shaping dominant role in ecosystems have played an additional animal organ systems can be expected in the near future. role in selecting model species. These arguments were Address used to attract sufficient funding for genome sequencing Sars International Centre for Marine Molecular Biology, University of and developmental studies that are cost-intensive inves- Bergen, Thormøhlensgate 55, 5008 Bergen, Norway tigations.
    [Show full text]
  • Platyhelminthes) at the Queensland Museum B.M
    VOLUME 53 ME M OIRS OF THE QUEENSLAND MUSEU M BRIS B ANE 30 NOVE mb ER 2007 © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email [email protected] Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 Volume 53 is complete in one part. NOTE Papers published in this volume and in all previous volumes of the Memoirs of the Queensland Museum may be reproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site www.qm.qld.gov.au/organisation/publications/memoirs/guidetoauthors.pdf A Queensland Government Project Typeset at the Queensland Museum THE STUDY OF TURBELLARIANS (PLATYHELMINTHES) AT THE QUEENSLAND MUSEUM B.M. ANGUS Angus, B.M. 2007 11 30: The study of turbellarians (Platyhelminthes) at the Queensland Museum. Memoirs of the Queensland Museum 53(1): 157-185. Brisbane. ISSN 0079-8835. Turbellarian research was largely ignored in Australia, apart from some early interest at the turn of the 19th century. The modern study of this mostly free-living branch of the phylum Platyhelminthes was led by Lester R.G. Cannon of the Queensland Museum. A background to the study of turbellarians is given particularly as it relates to the efforts of Cannon on symbiotic fauna, and his encouragement of visiting specialists and students.
    [Show full text]
  • Zootaxa, Platyhelminthes, Acoela, Acoelomorpha, Convolutidae
    Zootaxa 1008: 1–11 (2005) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 1008 Copyright © 2005 Magnolia Press ISSN 1175-5334 (online edition) Waminoa brickneri n. sp. (Acoela: Acoelomorpha) associated with corals in the Red Sea MAXINA V. OGUNLANA1, MATTHEW D. HOOGE1,4, YONAS I. TEKLE3, YEHUDA BENAYAHU2, ORIT BARNEAH2 & SETH TYLER1 1Department of Biological sciences, The University of Maine, 5751 Murray Hall, Orono, ME 04469-5751, USA., e-mail: [email protected], [email protected], [email protected] 2 Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Ramat Aviv, Tel Aviv, 69978, Israel., e-mail: [email protected] 3 Department of Systematic Zoology, Evolutionary Biology Centre, Upsala University, Norbyvägen 18D, SE- 752 36 Uppsala, Sweden., e-Mail: [email protected] 4 Author of correspondence Abstract While the majority of acoels live in marine sediments, some, usually identified as Waminoa sp., have been found associated with corals, living closely appressed to their external surfaces. We describe a new species collected from the stony coral Plesiastrea laxa in the Red Sea. Waminoa brickneri n. sp. can infest corals in high numbers, often forming clusters in non-overlapping arrays. It is bronze-colored, owing to the presence of two types of dinoflagellate endosymbionts, and speckled white with small scattered pigment spots. Its body is disc-shaped, highly flattened and cir- cular in profile except for a small notch at the posterior margin where the reproductive organs lie. The male copulatory organ is poorly differentiated, but comprises a seminal vesicle weakly walled by concentrically layered muscles, and a small penis papilla with serous glands at its juncture with the male pore.
    [Show full text]
  • Frontiers in Zoology Biomed Central
    Frontiers in Zoology BioMed Central Research Open Access Myogenesis in the basal bilaterian Symsagittifera roscoffensis (Acoela) Henrike Semmler*1, Xavier Bailly2 and Andreas Wanninger1 Address: 1University of Copenhagen, Department of Biology, Research Group for Comparative Zoology, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark and 2Station Station Biologique de Roscoff, Place Georges Teissier BP74, F-29682 Roscoff Cedex, France Email: Henrike Semmler* - [email protected]; Xavier Bailly - [email protected]; Andreas Wanninger - [email protected] * Corresponding author Published: 19 September 2008 Received: 5 May 2008 Accepted: 19 September 2008 Frontiers in Zoology 2008, 5:14 doi:10.1186/1742-9994-5-14 This article is available from: http://www.frontiersinzoology.com/content/5/1/14 © 2008 Semmler et al; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Abstract Background: In order to increase the weak database concerning the organogenesis of Acoela – a clade regarded by many as the earliest extant offshoot of Bilateria and thus of particular interest for studies concerning the evolution of animal bodyplans – we analyzed the development of the musculature of Symsagittifera roscoffensis using F-actin labelling, confocal laserscanning microscopy, and 3D reconstruction software. Results: At 40% of development between egg deposition and hatching short subepidermal fibres form. Muscle fibre development in the anterior body half precedes myogenesis in the posterior half. At 42% of development a grid of outer circular and inner longitudinal muscles is present in the bodywall.
    [Show full text]
  • Species Composition of the Free Living Multicellular Invertebrate Animals
    Historia naturalis bulgarica, 21: 49-168, 2015 Species composition of the free living multicellular invertebrate animals (Metazoa: Invertebrata) from the Bulgarian sector of the Black Sea and the coastal brackish basins Zdravko Hubenov Abstract: A total of 19 types, 39 classes, 123 orders, 470 families and 1537 species are known from the Bulgarian Black Sea. They include 1054 species (68.6%) of marine and marine-brackish forms and 508 species (33.0%) of freshwater-brackish, freshwater and terrestrial forms, connected with water. Five types (Nematoda, Rotifera, Annelida, Arthropoda and Mollusca) have a high species richness (over 100 species). Of these, the richest in species are Arthropoda (802 species – 52.2%), Annelida (173 species – 11.2%) and Mollusca (152 species – 9.9%). The remaining 14 types include from 1 to 38 species. There are some well-studied regions (over 200 species recorded): first, the vicinity of Varna (601 spe- cies), where investigations continue for more than 100 years. The aquatory of the towns Nesebar, Pomorie, Burgas and Sozopol (220 to 274 species) and the region of Cape Kaliakra (230 species) are well-studied. Of the coastal basins most studied are the lakes Durankulak, Ezerets-Shabla, Beloslav, Varna, Pomorie, Atanasovsko, Burgas, Mandra and the firth of Ropotamo River (up to 100 species known). The vertical distribution has been analyzed for 800 species (75.9%) – marine and marine-brackish forms. The great number of species is found from 0 to 25 m on sand (396 species) and rocky (257 species) bottom. The groups of stenohypo- (52 species – 6.5%), stenoepi- (465 species – 58.1%), meso- (115 species – 14.4%) and eurybathic forms (168 species – 21.0%) are represented.
    [Show full text]
  • Marine Flora and Fauna of the Northeastern United States
    NOAA Technical Report NMFS Circular 440 Marine Flora and Fauna of the Northeastern United States. Turbellaria: Acoela and Nemertodermatida Louise F. Bush July 1981 u.s. DEPARTMENT OF COMMERCE Malcolm Baldrige, Secretary National Oceanic and Atmospheric Administration National Marine Fisheries Service Terry L. Leitzell, Assistant Administrator for FisherIes FOREWORD This NMFS Circular is part of the subseries "Marine Flora and Fauna of the Northeastern United States;' which consists of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and, animals of the northeastern United States. The manuals are published at irregular intervals on as many taxa of the region as there are specialists available to collaborate in their preparation. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward to approximately the 200 m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of collaborators. Whenever possible representative specimens dealt with in the manuals are deposited in the reference collections of major museums of the region. The "Marine Flora and Fauna of the Northeastern United States" is being prepared in col­ laboration with systematic specialists in the United States and abroad. Each manual is based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals, Each major taxon, treated in a separate manual, includes an introduction, illustrated glossary, uniform originally illustrated keys, annotated checklist with information \vhen available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic jnde:\.
    [Show full text]
  • The Evolution of Bilaterian Body‐Plan: Perspectives from the Developmental Genetics of the Acoela (Acoelomorpha)
    The evolution of bilaterian body‐plan: perspectives from the developmental genetics of the Acoela (Acoelomorpha) Marta Chiodin ADVERTIMENT. La consulta d’aquesta tesi queda condicionada a l’acceptació de les següents condicions d'ús: La difusió d’aquesta tesi per mitjà del servei TDX (www.tdx.cat) i a través del Dipòsit Digital de la UB (diposit.ub.edu) ha estat autoritzada pels titulars dels drets de propietat intel·lectual únicament per a usos privats emmarcats en activitats d’investigació i docència. No s’autoritza la seva reproducció amb finalitats de lucre ni la seva difusió i posada a disposició des d’un lloc aliè al servei TDX ni al Dipòsit Digital de la UB. No s’autoritza la presentació del seu contingut en una finestra o marc aliè a TDX o al Dipòsit Digital de la UB (framing). Aquesta reserva de drets afecta tant al resum de presentació de la tesi com als seus continguts. En la utilització o cita de parts de la tesi és obligat indicar el nom de la persona autora. ADVERTENCIA. La consulta de esta tesis queda condicionada a la aceptación de las siguientes condiciones de uso: La difusión de esta tesis por medio del servicio TDR (www.tdx.cat) y a través del Repositorio Digital de la UB (diposit.ub.edu) ha sido autorizada por los titulares de los derechos de propiedad intelectual únicamente para usos privados enmarcados en actividades de investigación y docencia. No se autoriza su reproducción con finalidades de lucro ni su difusión y puesta a disposición desde un sitio ajeno al servicio TDR o al Repositorio Digital de la UB.
    [Show full text]
  • Xenacoelomorpha Is the Sister Group to Nephrozoa
    LETTER doi:10.1038/nature16520 Xenacoelomorpha is the sister group to Nephrozoa Johanna Taylor Cannon1, Bruno Cossermelli Vellutini2, Julian Smith III3, Fredrik Ronquist1, Ulf Jondelius1 & Andreas Hejnol2 The position of Xenacoelomorpha in the tree of life remains a major as sister taxon to remaining Bilateria19 (Fig. 1b–e). The deuterostome unresolved question in the study of deep animal relationships1. affiliation derives support from three lines of evidence4: an analysis Xenacoelomorpha, comprising Acoela, Nemertodermatida, and of mitochondrial gene sequences, microRNA complements, and a Xenoturbella, are bilaterally symmetrical marine worms that lack phylogenomic data set. Analyses of mitochondrial genes recovered several features common to most other bilaterians, for example Xenoturbella within deuterostomes18. However, limited mitochon- an anus, nephridia, and a circulatory system. Two conflicting drial data (typically ~16 kilobase total nucleotides, 13 protein-coding hypotheses are under debate: Xenacoelomorpha is the sister group genes) are less efficient in recovering higher-level animal relationships to all remaining Bilateria (= Nephrozoa, namely protostomes than phylogenomic approaches, especially in long-branching taxa1. and deuterostomes)2,3 or is a clade inside Deuterostomia4. Thus, The one complete and few partial mitochondrial genomes for acoelo- determining the phylogenetic position of this clade is pivotal for morphs are highly divergent in terms of both gene order and nucleotide understanding the early evolution of bilaterian features, or as a sequence19,20. Analyses of new complete mitochondrial genomes of case of drastic secondary loss of complexity. Here we show robust Xenoturbella spp. do not support any phylogenetic hypothesis for this phylogenomic support for Xenacoelomorpha as the sister taxon taxon21. Ref. 4 proposes that microRNA data support Xenacoelomorpha of Nephrozoa.
    [Show full text]
  • Introduction to the Bilateria and the Phylum Xenacoelomorpha Triploblasty and Bilateral Symmetry Provide New Avenues for Animal Radiation
    CHAPTER 9 Introduction to the Bilateria and the Phylum Xenacoelomorpha Triploblasty and Bilateral Symmetry Provide New Avenues for Animal Radiation long the evolutionary path from prokaryotes to modern animals, three key innovations led to greatly expanded biological diversification: (1) the evolution of the eukaryote condition, (2) the emergence of the A Metazoa, and (3) the evolution of a third germ layer (triploblasty) and, perhaps simultaneously, bilateral symmetry. We have already discussed the origins of the Eukaryota and the Metazoa, in Chapters 1 and 6, and elsewhere. The invention of a third (middle) germ layer, the true mesoderm, and evolution of a bilateral body plan, opened up vast new avenues for evolutionary expan- sion among animals. We discussed the embryological nature of true mesoderm in Chapter 5, where we learned that the evolution of this inner body layer fa- cilitated greater specialization in tissue formation, including highly specialized organ systems and condensed nervous systems (e.g., central nervous systems). In addition to derivatives of ectoderm (skin and nervous system) and endoderm (gut and its de- Classification of The Animal rivatives), triploblastic animals have mesoder- Kingdom (Metazoa) mal derivatives—which include musculature, the circulatory system, the excretory system, Non-Bilateria* Lophophorata and the somatic portions of the gonads. Bilater- (a.k.a. the diploblasts) PHYLUM PHORONIDA al symmetry gives these animals two axes of po- PHYLUM PORIFERA PHYLUM BRYOZOA larity (anteroposterior and dorsoventral) along PHYLUM PLACOZOA PHYLUM BRACHIOPODA a single body plane that divides the body into PHYLUM CNIDARIA ECDYSOZOA two symmetrically opposed parts—the left and PHYLUM CTENOPHORA Nematoida PHYLUM NEMATODA right sides.
    [Show full text]
  • Oceanic Mixotrophic Flatworms*
    MARINE ECOLOGY PROGRESS SERIES Vol. 58: 41-51, 1989 Published December 15 1 Mar. Ecol. Prog. Ser. Oceanic mixotrophic flatworms* ' Biology Department, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, USA Department of Marine Biology, University of Bergen, N-5065 Blomsterdalen, Norway Department of Zoology, University of Maine, Orono, Maine 04469, USA ABSTRACT Most reports of photosynthetic flatworms are from benthic or littoral habitats, but small (< 1 mm) acoel flatworms with algal endosymbionts are a widespread, though sporadic component of the open-ocean plankton in warm waters Among oceanic flatworms are specimens harbor~ng prasinophyte or less commonly, dinophyte endosymbionts Photosynthesis was measured by I4C uptake in flatworms from shelf/slope waters in the western north Atlantlc and from the Sargasso Sea Rates were as high as 27 ng C fixed ind -' h-' Assimilation ratios ranged from 0 9 to 1 3 ng C fixed (ng Chlorophyll a)-' h-' Although these acoels were photosynthetic, they were also predatory on other plankton Remains of crustaceans and radiolanan central capsules were observed in the guts or fecal material of some specimens These acoel-algal associations apparently depend on both autotrophic and heterotrophic nutrition and are thus mixotrophic Among the planktonic protozoa, mixotrophy is a common nutnbonal strategy, it also appears to be common strategy among certain taxa of open-ocean metazoa INTRODUCTION pelagica, C, schultzei, and Adenopea illardatus (Lohner & Micoletzky 1911, Dorjes 1970). Acoel
    [Show full text]
  • Xenacoelomorpha's Significance for Understanding Bilaterian
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Elsevier - Publisher Connector Available online at www.sciencedirect.com ScienceDirect Xenacoelomorpha’s significance for understanding bilaterian evolution Andreas Hejnol and Kevin Pang The Xenacoelomorpha, with its phylogenetic position as sister biology models are the fruitfly Drosophila melanogaster and group of the Nephrozoa (Protostomia + Deuterostomia), plays the nematode Caenorhabditis elegans, in which basic prin- a key-role in understanding the evolution of bilaterian cell types ciples of developmental processes have been studied in and organ systems. Current studies of the morphological and great detail. It might be because the field of evolutionary developmental diversity of this group allow us to trace the developmental biology — EvoDevo — has its origin in evolution of different organ systems within the group and to developmental biology and not evolutionary biology that reconstruct characters of the most recent common ancestor of species under investigation are often called ‘model spe- Xenacoelomorpha. The disparity of the clade shows that there cies’. Criteria for selected representative species are cannot be a single xenacoelomorph ‘model’ species and primarily the ease of access to collected material and strategic sampling is essential for understanding the evolution their ability to be cultivated in the lab [1]. In some cases, of major traits. With this strategy, fundamental insights into the a supposedly larger number of ancestral characters or a evolution of molecular mechanisms and their role in shaping dominant role in ecosystems have played an additional animal organ systems can be expected in the near future. role in selecting model species.
    [Show full text]
  • Hidden Diversity of Acoelomorpha Revealed Through
    View metadata, citation and similar papers atDownloaded core.ac.uk from http://rsbl.royalsocietypublishing.org/ on June 20, 2017 brought to you by CORE provided by Digital.CSIC Evolutionary biology Hidden diversity of Acoelomorpha rsbl.royalsocietypublishing.org revealed through metabarcoding Alicia S. Arroyo1, David Lo´pez-Escardo´1, Colomban de Vargas2,3 and In˜aki Ruiz-Trillo1,4,5 Research 1Institut de Biologia Evolutiva (CSIC-Universitat Pompeu Fabra), Passeig Marı´tim de la Barceloneta, 37-49, Cite this article: Arroyo AS, Lo´pez-Escardo´ D, 08003 Barcelona, Spain 2CNRS, UMR 7144, Station Biologique de Roscoff, Place Georges Teissier, 29680 Roscoff, France de Vargas C, Ruiz-Trillo I. 2016 Hidden 3Sorbonne Universite´s, Universite´ Pierre et Marie Curie (UPMC) Paris 06, UMR 7144, Station Biologique de diversity of Acoelomorpha revealed through Roscoff, Place Georges Teissier, 29680 Roscoff, France metabarcoding. Biol. Lett. 12: 20160674. 4Departament de Gene`tica, Microbiologia i Estadı´stica, Universitat de Barcelona, Avinguda Diagonal 643, 08028 http://dx.doi.org/10.1098/rsbl.2016.0674 Barcelona, Spain 5ICREA, Pg. Lluı´s Companys 23, 08010 Barcelona, Spain IR-T, 0000-0001-6547-5304 Received: 16 August 2016 Animals with bilateral symmetry comprise the majority of the described Accepted: 6 September 2016 species within Metazoa. However, the nature of the first bilaterian animal remains unknown. As most recent molecular phylogenies point to Xenacoe- lomorpha as the sister group to the rest of Bilateria, understanding their biology, ecology and diversity is key to reconstructing the nature of the last common bilaterian ancestor (Urbilateria). To date, sampling efforts Subject Areas: have focused mainly on coastal areas, leaving potential gaps in our under- evolution, ecology standing of the full diversity of xenacoelomorphs.
    [Show full text]