An Introduction to Vertebrates II – the Bilateria
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Comparative Neuroanatomy of Mollusks and Nemerteans in the Context of Deep Metazoan Phylogeny
Comparative Neuroanatomy of Mollusks and Nemerteans in the Context of Deep Metazoan Phylogeny Von der Fakultät für Mathematik, Informatik und Naturwissenschaften der RWTH Aachen University zur Erlangung des akademischen Grades einer Doktorin der Naturwissenschaften genehmigte Dissertation vorgelegt von Diplom-Biologin Simone Faller aus Frankfurt am Main Berichter: Privatdozent Dr. Rudolf Loesel Universitätsprofessor Dr. Peter Bräunig Tag der mündlichen Prüfung: 09. März 2012 Diese Dissertation ist auf den Internetseiten der Hochschulbibliothek online verfügbar. Contents 1 General Introduction 1 Deep Metazoan Phylogeny 1 Neurophylogeny 2 Mollusca 5 Nemertea 6 Aim of the thesis 7 2 Neuroanatomy of Minor Mollusca 9 Introduction 9 Material and Methods 10 Results 12 Caudofoveata 12 Scutopus ventrolineatus 12 Falcidens crossotus 16 Solenogastres 16 Dorymenia sarsii 16 Polyplacophora 20 Lepidochitona cinerea 20 Acanthochitona crinita 20 Scaphopoda 22 Antalis entalis 22 Entalina quinquangularis 24 Discussion 25 Structure of the brain and nerve cords 25 Caudofoveata 25 Solenogastres 26 Polyplacophora 27 Scaphopoda 27 i CONTENTS Evolutionary considerations 28 Relationship among non-conchiferan molluscan taxa 28 Position of the Scaphopoda within Conchifera 29 Position of Mollusca within Protostomia 30 3 Neuroanatomy of Nemertea 33 Introduction 33 Material and Methods 34 Results 35 Brain 35 Cerebral organ 38 Nerve cords and peripheral nervous system 38 Discussion 38 Peripheral nervous system 40 Central nervous system 40 In search for the urbilaterian brain 42 4 General Discussion 45 Evolution of higher brain centers 46 Neuroanatomical glossary and data matrix – Essential steps toward a cladistic analysis of neuroanatomical data 49 5 Summary 53 6 Zusammenfassung 57 7 References 61 Danksagung 75 Lebenslauf 79 ii iii 1 General Introduction Deep Metazoan Phylogeny The concept of phylogeny follows directly from the theory of evolution as published by Charles Darwin in The origin of species (1859). -
Defining Phyla: Evolutionary Pathways to Metazoan Body Plans
EVOLUTION & DEVELOPMENT 3:6, 432-442 (2001) Defining phyla: evolutionary pathways to metazoan body plans Allen G. Collins^ and James W. Valentine* Museum of Paleontology and Department of Integrative Biology, University of California, Berkeley, CA 94720, USA 'Author for correspondence (email: [email protected]) 'Present address: Section of Ecology, Befiavior, and Evolution, Division of Biology, University of California, San Diego, La Jolla, CA 92093-0116, USA SUMMARY Phyla are defined by two sets of criteria, one pothesis of Nielsen; the clonal hypothesis of Dewel; the set- morphological and the other historical. Molecular evidence aside cell hypothesis of Davidson et al.; and a benthic hy- permits the grouping of animals into clades and suggests that pothesis suggested by the fossil record. It is concluded that a some groups widely recognized as phyla are paraphyletic, benthic radiation of animals could have supplied the ances- while some may be polyphyletic; the phyletic status of crown tral lineages of all but a few phyla, is consistent with molecu- phyla is tabulated. Four recent evolutionary scenarios for the lar evidence, accords well with fossil evidence, and accounts origins of metazoan phyla and of supraphyletic clades are as- for some of the difficulties in phylogenetic analyses of phyla sessed in the light of a molecular phylogeny: the trochaea hy- based on morphological criteria. INTRODUCTION Molecules have provided an important operational ad- vance to addressing questions about the origins of animal Concepts of animal phyla have changed importantly from phyla. Molecular developmental and comparative genomic their origins in the six Linnaean classis and four Cuvieran evidence offer insights into the genetic bases of body plan embranchements. -
A Phylum-Wide Survey Reveals Multiple Independent Gains of Head Regeneration Ability in Nemertea
bioRxiv preprint doi: https://doi.org/10.1101/439497; this version posted October 11, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. A phylum-wide survey reveals multiple independent gains of head regeneration ability in Nemertea Eduardo E. Zattara1,2,5, Fernando A. Fernández-Álvarez3, Terra C. Hiebert4, Alexandra E. Bely2 and Jon L. Norenburg1 1 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA 2 Department of Biology, University of Maryland, College Park, MD, USA 3 Institut de Ciències del Mar, Consejo Superior de Investigaciones Científicas, Barcelona, Spain 4 Institute of Ecology and Evolution, University of Oregon, Eugene, OR, USA 5 INIBIOMA, Consejo Nacional de Investigaciones Científicas y Tecnológicas, Bariloche, RN, Argentina Corresponding author: E.E. Zattara, [email protected] Abstract Animals vary widely in their ability to regenerate, suggesting that regenerative abilities have a rich evolutionary history. However, our understanding of this history remains limited because regeneration ability has only been evaluated in a tiny fraction of species. Available comparative regeneration studies have identified losses of regenerative ability, yet clear documentation of gains is lacking. We surveyed regenerative ability in 34 species spanning the phylum Nemertea, assessing the ability to regenerate heads and tails either through our own experiments or from literature reports. Our sampling included representatives of the 10 most diverse families and all three orders comprising this phylum. -
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. -
Invertebrates Invertebrates: • Are Animals Without Backbones • Represent 95% of the Animal Kingdom Animal Diversity Morphological Vs
Invertebrates Invertebrates: • Are animals without backbones • Represent 95% of the animal kingdom Animal Diversity Morphological vs. Molecular Character Phylogeny? A tree is a hypothesis supported or not supported by evidence. Groupings change as new evidence become available. Sponges - Porifera Natural Bath Sponges – over-collected, now uncommon Sponges • Perhaps oldest animal phylum (Ctenphora possibly older) • may represent several old phyla, some now extinct ----------------Ctenophora? Sponges - Porifera • Mostly marine • Sessile animals • Lack true tissues; • Have only a few cell types, cells kind of independent • Most have no symmetry • Body resembles a sac perforated with holes, system of canals. • Strengthened by fibers of spongin, spicules Sponges have a variety of shapes Sponges Pores Choanocyte Amoebocyte (feeding cell) Skeletal Water fiber flow Central cavity Flagella Choanocyte in contact with an amoebocyte Sponges - Porifera • Sessile filter feeder • No mouth • Sac-like body, perforated by pores. • Interior lined by flagellated cells (choanocytes). Flagellated collar cells generate a current, draw water through the walls of the sponge where food is collected. • Amoeboid cells move around in the mesophyll and distribute food. Sponges - Porifera Grantia x.s. Sponge Reproduction Asexual reproduction • Fragmentation or by budding. • Sponges are capable of regeneration, growth of a whole from a small part. Sexual reproduction • Hermaphrodites, produce both eggs and sperm • Eggs and sperm released into the central cavity • Produces -
Carbonic Anhydrases: an Ancient Tool in Calcareous Sponge Biomineralization
fgene-12-624533 March 30, 2021 Time: 13:30 # 1 BRIEF RESEARCH REPORT published: 07 April 2021 doi: 10.3389/fgene.2021.624533 Carbonic Anhydrases: An Ancient Tool in Calcareous Sponge Biomineralization Oliver Voigt1*, Benedetta Fradusco1, Carolin Gut1, Charalampos Kevrekidis1, Sergio Vargas1 and Gert Wörheide1,2,3 1 Department of Earth and Environmental Sciences, Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany, 2 GeoBio-Center, Ludwig-Maximilians-Universität München, Munich, Germany, 3 SNSB-Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany Enzymes of the a-carbonic anhydrase gene family (CAs) are essential for the deposition of calcium carbonate biominerals. In calcareous sponges (phylum Porifera, class Calcarea), specific CAs are involved in the formation of calcite spicules, a unique trait and synapomorphy of this class. However, detailed studies on the CA repertoire of calcareous sponges exist for only two species of one of the two Calcarea subclasses, the Calcaronea. The CA repertoire of the second subclass, the Calcinea, has not been investigated so far, leaving a considerable gap in our knowledge about this Edited by: Melanie Debiais-Thibaud, gene family in Calcarea. Here, using transcriptomic analysis, phylogenetics, and in situ Université de Montpellier, France hybridization, we study the CA repertoire of four additional species of calcareous Reviewed by: sponges, including three from the previously unsampled subclass Calcinea. Our data Helena Cetkovi´ c,´ Rudjer Boskovic Institute, Croatia indicate that the last common ancestor of Calcarea had four ancestral CAs with defined Ana Riesgo, subcellular localizations and functions (mitochondrial/cytosolic, membrane-bound, and Natural History Museum, secreted non-catalytic). The evolution of membrane-bound and secreted CAs involved United Kingdom gene duplications and losses, whereas mitochondrial/cytosolic and non-catalytic CAs *Correspondence: Oliver Voigt are evidently orthologous genes. -
S I Section 4
3/31/2011 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Porifera Ecdysozoa Deuterostomia Lophotrochozoa Cnidaria and Ctenophora Cnidaria and Protostomia SSiection 4 Radiata Bilateria Professor Donald McFarlane Parazoa Eumetazoa Lecture 13 Invertebrates: Parazoa, Radiata, and Lophotrochozoa Ancestral colonial choanoflagellate 2 Traditional classification based on Parazoa – Phylum Porifera body plans Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Parazoa 4 main morphological and developmental Sponges features used Loosely organized and lack Porifera Ecdysozoa Cnidaria and Ctenophora tissues euterostomia photrochozoa D 1. o Presence or absence of different tissue L Multicellular with several types of types Protostomia cells 2. Type of body symmetry Radiata Bilateria 8,000 species, mostly marine Parazoa Eumetazoa 3. Presence or absence of a true body No apparent symmetry cavity Ancestral colonial Adults sessile, larvae free- choanoflagellate 4. Patterns of embryonic development swimming 3 4 1 3/31/2011 Water drawn through pores (ostia) into spongocoel Flows out through osculum Reproduce Choanocytes line spongocoel Sexually Most hermapppgggphrodites producing eggs and sperm Trap and eat small particles and plankton Gametes are derived from amoebocytes or Mesohyl between choanocytes and choanocytes epithelial cells Asexually Amoebocytes absorb food from choanocytes, Small fragment or bud may detach and form a new digest it, and carry -
Analysis of the Complete Mitochondrial DNA Sequence of the Brachiopod Terebratulina Retusa Places Brachiopoda Within the Protostomes
See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/12415870 Analysis of the complete mitochondrial DNA sequence of the brachiopod Terebratulina retusa places Brachiopoda within the protostomes Article in Proceedings of the Royal Society B: Biological Sciences · November 1999 DOI: 10.1098/rspb.1999.0885 · Source: PubMed CITATIONS READS 83 50 2 authors, including: Martin Schlegel University of Leipzig 151 PUBLICATIONS 2,931 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Rare for a reason? Scale-dependence of factors influencing rarity and diversity of xylobiont beetles View project Bat diversity and vertical niche activity in the fluvial flood forest Leipzig View project All content following this page was uploaded by Martin Schlegel on 22 May 2014. The user has requested enhancement of the downloaded file. Analysis of the complete mitochondrial DNA sequence of the brachiopod Terebratulina retusa places Brachiopoda within the protostomes Alexandra Stechmann* and Martin Schlegel UniversitÌt Leipzig, Institut fÏr Zoologie/Spezielle Zoologie,Talstr. 33, 04103 Leipzig, Germany Brachiopod phylogeny is still a controversial subject. Analyses using nuclear 18SrRNA and mitochondrial 12SrDNA sequences place them within the protostomes but some recent interpretations of morphological data support a relationship with deuterostomes. In order to investigate brachiopod a¤nities within the metazoa further,we compared the gene arrangement on the brachiopod mitochondrial genome with several metazoan taxa. The complete (15 451bp) mitochondrial DNA (mtDNA) sequence of the articulate brachiopod Terebratulina retusa was determined from two overlapping long polymerase chain reaction products. All the genes are encoded on the same strand and gene order comparisons showed that only one major rearrangement is required to interconvert the T.retusa and Katharina tunicata (Mollusca: Polyplaco- phora) mitochondrial genomes. -
The Genome of the Poecilogonous Annelid Streblospio Benedicti Christina Zakas1, Nathan D
bioRxiv preprint doi: https://doi.org/10.1101/2021.04.15.440069; this version posted April 16, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. The genome of the poecilogonous annelid Streblospio benedicti Christina Zakas1, Nathan D. Harry1, Elizabeth H. Scholl2 and Matthew V. Rockman3 1Department of Genetics, North Carolina State University, Raleigh, NC, USA 2Bioinformatics Research Center, North Carolina State University, Raleigh, NC, USA 3Department of Biology and Center for Genomics & Systems Biology, New York University, New York, NY, USA [email protected] [email protected] Abstract Streblospio benedicti is a common marine annelid that has become an important model for developmental evolution. It is the only known example of poecilogony, where two distinct developmental modes occur within a single species, that is due to a heritable difference in egg size. The dimorphic developmental programs and life-histories exhibited in this species depend on differences within the genome, making it an optimal model for understanding the genomic basis of developmental divergence. Studies using S. benedicti have begun to uncover the genetic and genomic principles that underlie developmental uncoupling, but until now they have been limited by the lack of availability of genomic tools. Here we present an annotated chromosomal-level genome assembly of S. benedicti generated from a combination of Illumina reads, Nanopore long reads, Chicago and Hi-C chromatin interaction sequencing, and a genetic map from experimental crosses. At 701.4 Mb, the S. benedicti genome is the largest annelid genome to date that has been assembled to chromosomal scaffolds, yet it does not show evidence of extensive gene family expansion, but rather longer intergenic regions. -
Phylogenomics of Tubeworms (Siboglinidae, Annelida) and Comparative Performance of Different Reconstruction Methods
Zoologica Scripta Phylogenomics of tubeworms (Siboglinidae, Annelida) and comparative performance of different reconstruction methods YUANNING LI,KEVIN M. KOCOT,NATHAN V. WHELAN,SCOTT R. SANTOS,DAMIEN S. WAITS, DANIEL J. THORNHILL &KENNETH M. HALANYCH Submitted: 28 January 2016 Li, Y., Kocot, K.M., Whelan, N.V., Santos, S.R., Waits, D.S., Thornhill, D.J. & Halanych, Accepted: 18 June 2016 K.M. (2016). Phylogenomics of tubeworms (Siboglinidae, Annelida) and comparative perfor- doi:10.1111/zsc.12201 mance of different reconstruction methods. —Zoologica Scripta, 00: 000–000. Deep-sea tubeworms (Annelida, Siboglinidae) represent dominant species in deep-sea chemosynthetic communities (e.g. hydrothermal vents and cold methane seeps) and occur in muddy sediments and organic falls. Siboglinids lack a functional digestive tract as adults, and they rely on endosymbiotic bacteria for energy, making them of evolutionary and physi- ological interest. Despite their importance, inferred evolutionary history of this group has been inconsistent among studies based on different molecular markers. In particular, place- ment of bone-eating Osedax worms has been unclear in part because of their distinctive biol- ogy, including harbouring heterotrophic bacteria as endosymbionts, displaying extreme sexual dimorphism and exhibiting a distinct body plan. Here, we reconstructed siboglinid evolutionary history using 12 newly sequenced transcriptomes. We parsed data into three data sets that accommodated varying levels of missing data, and we evaluate effects of miss- ing data on phylogenomic inference. Additionally, several multispecies-coalescent approaches and Bayesian concordance analysis (BCA) were employed to allow for a compar- ison of results to a supermatrix approach. Every analysis conducted herein strongly sup- ported Osedax being most closely related to the Vestimentifera and Sclerolinum clade, rather than Frenulata, as previously reported. -
Animal Diversity Part 2
Textbook resources • pp. 517-522 • pp. 527-8 Animal Diversity • p. 530 part 2 • pp. 531-2 Clicker question In protostomes A. The blastopore becomes the mouth. B. The blastopore becomes the anus. C. Development involves indeterminate cleavage. D. B and C Fig. 25.2 Phylogeny to know (1). Symmetry Critical innovations to insert: Oral bilateral symmetry ecdysis mouth develops after anus multicellularity Aboral tissues 1 Animal diversity, part 2 Parazoa Diversity 2 I. Parazoa • Porifera: Sponges II. Cnidaria & Ctenophora • Tissues • Symmetry I. Outline the • Germ Layers III. Lophotrochozoa unique • Embryonic characteristics Development of sponges IV. Ecdysozoa • Body Cavities • Segmentation Parazoa Parazoa • Porifera: Sponges • Porifera: Sponges – Multicellular without – Hermaphrodites tissues – Sexual and asexual reproduction – Choanocytes (collar cells) use flagella to move water and nutrients into pores – Intracellular digestion Fig. 25.11 Animal diversity, part 2 Clicker Question Diversity 2 I. Parazoa In diploblastic animals, the inner lining of the digestive cavity or tract is derived from II. Cnidaria & Ctenophora A. Endoderm. II. Outline the B. Ectoderm. unique III. Lophotrochozoa C. Mesoderm. characteristics D. Coelom. of cnidarians and IV. Ecdysozoa ctenophores 2 Coral Box jelly Cnidaria and Ctenophora • Cnidarians – Coral; sea anemone; jellyfish; hydra; box jellies • Ctenophores – Comb jellies Sea anemone Jellyfish Hydra Comb jelly Cnidaria and Ctenophora Fig. 25.12 Coral Box jelly Cnidaria and Ctenophora • Tissues Fig. 25.12 – -
Tropical Marine Invertebrates CAS BI 569 Phylum ANNELIDA by J
Tropical Marine Invertebrates CAS BI 569 Phylum ANNELIDA by J. R. Finnerty Phylum ANNELIDA Porifera Ctenophora Cnidaria Deuterostomia Ecdysozoa Lophotrochozoa Chordata Arthropoda Annelida Hemichordata Onychophora Mollusca Echinodermata Nematoda Platyhelminthes Acoelomorpha Silicispongiae Calcispongia PROTOSTOMIA “BILATERIA” (=TRIPLOBLASTICA) Bilateral symmetry (?) Mesoderm (triploblasty) Phylum ANNELIDA Porifera Ctenophora Cnidaria Deuterostomia Ecdysozoa Lophotrochozoa Chordata Arthropoda Annelida Hemichordata Onychophora Mollusca Echinodermata Nematoda Platyhelminthes Acoelomorpha Silicispongiae Calcispongia PROTOSTOMIA “COELOMATA” True coelom Coelomata gut cavity endoderm mesoderm coelom ectoderm [note: dorso-ventral inversion] Phylum ANNELIDA Porifera Ctenophora Cnidaria Deuterostomia Ecdysozoa Lophotrochozoa Chordata Arthropoda Annelida Hemichordata Onychophora Mollusca Echinodermata Nematoda Platyhelminthes Acoelomorpha Silicispongiae Calcispongia PROTOSTOMIA PROTOSTOMIA “first mouth” blastopore contributes to mouth ventral nerve cord The Blastopore ! Forms during gastrulation ectoderm blastocoel blastocoel endoderm gut blastoderm BLASTULA blastopore The Gut “internal, epithelium-lined cavity for the digestion and absorption of food sponges lack a gut simplest gut = blind sac (Cnidaria) blastopore gives rise to dual- function mouth/anus through-guts evolve later Protostome = blastopore contributes to the mouth Deuterostome = blastopore becomes the anus; mouth is a second opening Protostomy blastopore mouth anus Deuterostomy blastopore