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Platyhelminthes, Nemertea, and "Aschelminthes" - A
BIOLOGICAL SCIENCE FUNDAMENTALS AND SYSTEMATICS – Vol. III - Platyhelminthes, Nemertea, and "Aschelminthes" - A. Schmidt-Rhaesa PLATYHELMINTHES, NEMERTEA, AND “ASCHELMINTHES” A. Schmidt-Rhaesa University of Bielefeld, Germany Keywords: Platyhelminthes, Nemertea, Gnathifera, Gnathostomulida, Micrognathozoa, Rotifera, Acanthocephala, Cycliophora, Nemathelminthes, Gastrotricha, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera Contents 1. Introduction 2. General Morphology 3. Platyhelminthes, the Flatworms 4. Nemertea (Nemertini), the Ribbon Worms 5. “Aschelminthes” 5.1. Gnathifera 5.1.1. Gnathostomulida 5.1.2. Micrognathozoa (Limnognathia maerski) 5.1.3. Rotifera 5.1.4. Acanthocephala 5.1.5. Cycliophora (Symbion pandora) 5.2. Nemathelminthes 5.2.1. Gastrotricha 5.2.2. Nematoda, the Roundworms 5.2.3. Nematomorpha, the Horsehair Worms 5.2.4. Priapulida 5.2.5. Kinorhyncha 5.2.6. Loricifera Acknowledgements Glossary Bibliography Biographical Sketch Summary UNESCO – EOLSS This chapter provides information on several basal bilaterian groups: flatworms, nemerteans, Gnathifera,SAMPLE and Nemathelminthes. CHAPTERS These include species-rich taxa such as Nematoda and Platyhelminthes, and as taxa with few or even only one species, such as Micrognathozoa (Limnognathia maerski) and Cycliophora (Symbion pandora). All Acanthocephala and subgroups of Platyhelminthes and Nematoda, are parasites that often exhibit complex life cycles. Most of the taxa described are marine, but some have also invaded freshwater or the terrestrial environment. “Aschelminthes” are not a natural group, instead, two taxa have been recognized that were earlier summarized under this name. Gnathifera include taxa with a conspicuous jaw apparatus such as Gnathostomulida, Micrognathozoa, and Rotifera. Although they do not possess a jaw apparatus, Acanthocephala also belong to Gnathifera due to their epidermal structure. ©Encyclopedia of Life Support Systems (EOLSS) BIOLOGICAL SCIENCE FUNDAMENTALS AND SYSTEMATICS – Vol. -
Phylum Nemertea Or Rhynchocoela (Minor Phyla)
Animal Diversity: (Non-Chordates) Phylum Nemertea or Rhynchocoela (Minor Phyla) Hardeep Kaur Assistant Professor, Department of Zoology, Ramjas College, University of Delhi Delhi – 110 007 CONTENTs: ¾ Introduction ¾ External Structure ¾ Body Wall and Locomotion ¾ Nutrition and Digestive System ¾ Circulatory System ¾ Excretory System ¾ Nervous System and Sense Organs ¾ Regeneration ¾ Reproductive System ¾ Embryogeny ¾ Classification of Nemerteans ¾ General Characters of Nemerteans ¾ Affinities of Nemerteans ¾ Glossary ¾ References / Suggested Readings PHYLUM NEMERTEA / PHYLUM RHYNCHOCOELA INTRODUCTION: Phylum Nemertea comprises approximately 1200 species of ¾ elongated and often flattened worms, called ribbon worms (many have flattened body) or ¾ bottle worms (because of narrow anterior end) ¾ proboscis worms, (because of the presence of a remarkable proboscis apparatus used in capturing food). The Nemerteans are named for Nemertes, one of the Nereids, sea-nymph of Greek mythology. They are commonly looked upon related to the Turbellaria and were formerly included in them, but the fact that they possess a complete digestive system with anus and also a blood vascular system makes them higher in organization than the Turbellaria. However, presence of a protrusible proboscis with a separate proboscis pore, other than mouth, is the most characteristic feature of the phylum. Almost all nemerteans are free living, bottom-dwelling, marine animals. Few commensal and parasitic species have been described. Nemertopsis actinophila is a slender form living beneath the pedal disc of sea anemones. Carcinonmertes may be found on gills and egg masses of crabs. Some species of Tetrastemma live in the branchial cavity of tunicates. Only few exibit commensal mode of life eg. Gonomertes parasitica is a commensal species found on crustaceans,. -
Comparative Biology of Oogenesis in Nemertean Worms Stephen A
AaaZoologka (Stockholm) 82: 213-230 (July 2001) Comparative biology of oogenesis in nemertean worms Stephen A. Strieker1, Toni L. Smythe1, Leonard Miller1 and Jon L. Norenburg2 Abstract 'Department of Biology, University of New Strieker, S. A., Smythe,T, L., Miller, L. and Norenburg, J. L. 2001. Mexico, Albuquerque, NM 87131; Comparative biology of oogenesis in nemertean worms. — Acta Zoobgica 2 Invertebrate Zoology, MRC 163,Museum (Stockholm) 82: 213-230 of Natural History, Washington, DC 20560 USA In order to supplement previous analyses of oogenesis in nemertean worms, this study uses light and electron microscopy to compare the ovaries and Keywords: oocytes in 16 species of nemerteans that represent various taxa within the ovary, ultrastructure, vitellogenesis, yolk, phylum, Nemertean ovaries comprise serially repeated sacs with an ovarian nucleoli, endoplasmic reticulum, confocal wall that characteristically includes myofilament-containing cells interspersed microscopy, oocyte maturation, serotonin among the germinal epithelium. Each oocyte can attach to the germinal epithelium by a vegetally situated stalk and resides in the ovarian lumen Accepted for publication: without being surrounded by follicle cells. In the ovary, oocytes arrest at 26 September 2000 prophase I of meiosis and contain a hypertrophied nucleus ('germinal vesicle') that often possesses multiple nucleoli. Intraovarian growth apparently involves an autosynthetic mode of yolk formation in most nemerteans and generates oocytes that measure ~60 Jim to 1 mm. When fully developed, oocytes can be discharged through a short gonoduct and are either spawned freely or deposited within egg cases. In most species, oocytes released from the ovary possess extracellular coats and resume maturation by undergoing germinal vesicle breakdown (GVBD). -
Resolving a 200-Year-Old Taxonomic Conundrum: Neotype Designation for Cephalothrix Linearis (Nemertea: Palaeonemertea) Based on a Topotype from Bergen, Norway
Fauna norvegica 2019 Vol. 39: 39–76. Resolving a 200-year-old taxonomic conundrum: neotype designation for Cephalothrix linearis (Nemertea: Palaeonemertea) based on a topotype from Bergen, Norway Hiroshi Kajihara1 Kajihara H. 2019. Resolving a 200-year-old taxonomic conundrum: neotype designation for Cephalothrix linearis (Nemertea: Palaeonemertea) based on a topotype from Bergen, Norway. Fauna norvegica 39: 39–76. The taxonomic identity of the palaeonemertean Cephalothrix linearis (Rathke, 1799) has been obscure for nearly two centuries, because its original description applies to almost any congeners, including Cephalothrix filiformis (Johnston 1828) and Cephalothrix rufifrons (Johnston, 1837), which occur commonly in the North Sea and adjacent waters. In this paper, I redescribe C. linearis based on two topotypes from Bergen, one herein designated as the neotype for C. linearis, because Rathke’s original material is not extant; I invoke Article 70.3.2 of the International Code of Zoological Nomenclature to fix Planaria linearis Rathke, 1799 as the type species of Cephalothrix Örsted, 1843 for the sake of stability. From the neotype, I determined sequences of the 28S rRNA, 16S rRNA, and cytochrome c oxidase subunit I (COI) genes. Using the COI sequence, I inferred the phylogenetic position of C. linearis along with 316 cephalotrichid sequences currently available in public databases. A tree-based species delimitation analysis detected 43 entities among them, with 34 in Cephalothrix and nine in either Balionemertes or Cephalotrichella. -
Phylum Nemertea
Biol. Lett. (2007) 3, 570–573 annelids, molluscs and entoprocts (plus some other doi:10.1098/rsbl.2007.0306 phyla varying between analyses). Published online 7 August 2007 Other Annulonemertes (supposedly different) species Phylogeny are reported in the literature (Norenburg 1988; Chernyshev & Minichev 2004) but A. minusculus is the only named species in the genus. There are other Annulonemertes (phylum nemertean species described with similar constrictions (notably Arenonemertes minutus, Friedrich 1949; Nemertea): when segments Nemertellina yamaokai, Kajihara et al. 2000), but neither species have these repeated constrictions do not count observed in Annulonemertes. The species is referred to as ‘segmented’ in zoological textbooks and thus enig- Per Sundberg* and Malin Strand matic from a phylogenetic point of view (e.g. Brusca & Department of Zoology, Go¨teborg University, PO Box 463, Brusca 2003). Its segmentation has also been 405 30 Go¨teborg, Sweden *Author for correspondence ([email protected]). discussed in the context of whether the last common ancestor of bilaterian animals was segmented We estimated the phylogenetic position of the (Balavoine & Adouette 2003). Giribet (2003) dis- pseudosegmented ribbon worm Annulonemertes minusculus to test proposed evolutionary hypo- cussed the morphological characters supporting the theses to explain these body constrictions. The Articulata versus Ecdyozoa metazoan phylogeny analysis is based on 18S rDNA sequences and hypotheses and refers to Annulonemertes as segmented shows that the species belongs to an apomorphic but at the same time conclude that this kind of serial clade of hoplonemertean species. The segmenta- repetition of structures is found in many phyla and tion has no phylogenetic bearing as previously do not really count as true segmentation (see also discussed, but is a derived character probably Scmidt-Rasea et al. -
Nemertea: Hoplonemertea: Polystilifera
Zootaxa 2429: 43–51 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition) Dinonemertes shinkaii sp. nov., (Nemertea: Hoplonemertea: Polystilifera: Pelagica) a new species of bathypelagic nemertean HIROSHI KAJIHARA1 & DHUGAL J. LINDSAY2 1Faculty of Science, Hokkaido University, Sapporo 060-0810, Japan. E-mail: [email protected] 2Japan Agency for Marine-Earth Science and Technology, Natsushima-cho 2-15, Yokosuka 237-0061, Japan. E-mail: [email protected] Abstract A new species of bathypelagic polystiliferous nemertean Dinonemertes shinkaii is described based on the holotype obtained by the manned submersible Shinkai 6500 from a depth of 2343 m in Japan Trench, Northwest Pacific. Dinonemertes shinkaii can be distinguished from its congeners in having a translucent body, 24 proboscis nerves, two pairs of intestinal caecal diverticula, and about 25 pairs of intestinal lateral diverticula. This species represents the first dinonemertid to have pseudostriated muscle fibres in the rhynchocoel circular muscle layers. Key words: Pacific, Japan Trench, plankton, manned submersible Shinkai 6500, taxonomy, striated muscle Introduction Since the description of Pelagonemertes rollestoni—the first pelagic nemertean species discovered during the Challenger expedition (Moseley 1875a)—so far about 100 species of pelagic nemerteans have been described/reported from all the oceans in the epipelagic, mesopelagic, and bathypelagic zones, although they seem to be most abundant at 625–2500 m (Roe & Norenburg 1999). All the pelagic forms belong to the Hoplonemertea, of which only two named and two undescribed species represent the Monostilifera (Wheeler 1934; Korotkevich 1961; Crandall & Gibson 1998; Chernyshev 2005; Crandall 2006), while the rest of the 98 species constitute the Pelagica within the Polystilifera (Maslakova & Norenburg 2001). -
A Taxonomic Catalogue of Japanese Nemerteans (Phylum Nemertea)
Title A Taxonomic Catalogue of Japanese Nemerteans (Phylum Nemertea) Author(s) Kajihara, Hiroshi Zoological Science, 24(4), 287-326 Citation https://doi.org/10.2108/zsj.24.287 Issue Date 2007-04 Doc URL http://hdl.handle.net/2115/39621 Rights (c) Zoological Society of Japan / 本文献の公開は著者の意思に基づくものである Type article Note REVIEW File Information zsj24p287.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP ZOOLOGICAL SCIENCE 24: 287–326 (2007) © 2007 Zoological Society of Japan [REVIEW] A Taxonomic Catalogue of Japanese Nemerteans (Phylum Nemertea) Hiroshi Kajihara* Department of Natural History Sciences, Faculty of Science, Hokkaido University, Sapporo 060-0810, Japan A literature-based taxonomic catalogue of the nemertean species (Phylum Nemertea) reported from Japanese waters is provided, listing 19 families, 45 genera, and 120 species as valid. Applications of the following species names to forms previously recorded from Japanese waters are regarded as uncertain: Amphiporus cervicalis, Amphiporus depressus, Amphiporus lactifloreus, Cephalothrix filiformis, Cephalothrix linearis, Cerebratulus fuscus, Lineus vegetus, Lineus bilineatus, Lineus gesserensis, Lineus grubei, Lineus longifissus, Lineus mcintoshii, Nipponnemertes pulchra, Oerstedia venusta, Prostoma graecense, and Prostoma grande. The identities of the taxa referred to by the fol- lowing four nominal species require clarification through future investigations: Cosmocephala japonica, Dicelis rubra, Dichilus obscurus, and Nareda serpentina. The nominal species established from Japanese waters are tabulated. In addition, a brief history of taxonomic research on Japanese nemerteans is reviewed. Key words: checklist, Pacific, classification, ribbon worm, Nemertinea 2001). The only recent listing of previously described Japa- INTRODUCTION nese species is the checklist of Crandall et al. (2002), but The phylum Nemertea comprises about 1,200 species the relevant literature is scattered. -
(Marlin) Review of Biodiversity for Marine Spatial Planning Within
The Marine Life Information Network® for Britain and Ireland (MarLIN) Review of Biodiversity for Marine Spatial Planning within the Firth of Clyde Report to: The SSMEI Clyde Pilot from the Marine Life Information Network (MarLIN). Contract no. R70073PUR Olivia Langmead Emma Jackson Dan Lear Jayne Evans Becky Seeley Rob Ellis Nova Mieszkowska Harvey Tyler-Walters FINAL REPORT October 2008 Reference: Langmead, O., Jackson, E., Lear, D., Evans, J., Seeley, B. Ellis, R., Mieszkowska, N. and Tyler-Walters, H. (2008). The Review of Biodiversity for Marine Spatial Planning within the Firth of Clyde. Report to the SSMEI Clyde Pilot from the Marine Life Information Network (MarLIN). Plymouth: Marine Biological Association of the United Kingdom. [Contract number R70073PUR] 1 Firth of Clyde Biodiversity Review 2 Firth of Clyde Biodiversity Review Contents Executive summary................................................................................11 1. Introduction...................................................................................15 1.1 Marine Spatial Planning................................................................15 1.1.1 Ecosystem Approach..............................................................15 1.1.2 Recording the Current Situation ................................................16 1.1.3 National and International obligations and policy drivers..................16 1.2 Scottish Sustainable Marine Environment Initiative...............................17 1.2.1 SSMEI Clyde Pilot ..................................................................17 -
Paranemertes Peregrina Class: Enopla Order: Hoplonemertea, Monostylifera "The Wanderer" Family: Emplectomenatidae
Phylum: Nemertea Paranemertes peregrina Class: Enopla Order: Hoplonemertea, Monostylifera "The wanderer" Family: Emplectomenatidae Taxonomy: Coe (1905) found two Proboscis: Eversible (phylum Nemertea) morphotypes within Paranemertes peregrina and, when not everted, coiled inside (var. alaskensis, var. californiensis) that rhynchocoel (cavity). Rhynchocoel half to differed in size, color and stylet morphology three-quarters body length (genus (Roe et al. 2007). Whether these Paranemertes). Proboscis whitish with one morphotypes represent two difference (suborder Monostilifera) short, stylet (order species or intraspecific divergence is currently Hoplonemertea) of lengths 85–90 µm (Coe unknown. 1905; Stricker and Cloney 1981). Stylet sculpture is variable and is either with (var. Description californiensis) (Fig. 4) or without spiral Size: Individuals vary in size from 2–40 cm grooves (var. alaskensis) (Roe et al. 2007). with average size range 15–25 cm (Coe Two (var. californiensis) to four (var. 1901; Roe et al. 2007). Northern specimens alaskensis) pouches of accessory stylets are (var. alaskensis, 40 cm) larger than southern present, each pouch with 6–10 stylets (Fig. 3) ones (var. californiensis, 10 cm) (Coe 1905). (Roe et al. 2007). Proboscis eversion can be Color: Dark dorsally, purple or olive green induced with fresh water or dilute acetic acid with head brown. Lighter ventrally, white or (Haderlie 1980). pale yellow with mid-ventral section sometimes lighter than the rest. No lines or Possible Misidentifications other patterns, except V-shape behind head. There are five genera of the family General Morphology: Long and slender Emplectonematidae on the Pacific coast, all Body: Elongate, contractile and non- of which have a short proboscis, numerous segmented (phylum Nemertea). -
Nectonemertes Mirabilis (Nemertea: Hoplonemertea: Pelagica) from Deep Waters of the Gulf of Mexico, first Sighting of a Pelagic Nemertean in the Gulf
Plankton Benthos Res 16(2): 139–144, 2021 Plankton & Benthos Research © The Plankton Society of Japan Nectonemertes mirabilis (Nemertea: Hoplonemertea: Pelagica) from deep waters of the Gulf of Mexico, first sighting of a pelagic nemertean in the gulf Marco Violante-Huerta1,* & Laura Sanvicente-Añorve2 1 Universidad Nacional Autónoma de México, Posgrado en Ciencias del Mar y Limnología, Ciudad Universitaria, C.P. 04510, Mexico City, Mexico 2 Universidad Nacional Autónoma de México, Departamento de Ecología y Biodiversidad Acuática, Instituto de Ciencias del Mar y Limnología, Ciudad Universitaria, C.P. 04510, Mexico City, Mexico Received 25 May 2020; Accepted 14 March 2021 Responsible Editor: Hiroomi Miyamoto doi: 10.3800/pbr.16.139 Abstract: The deep-water bathypelagic nemertean Nectonemertes mirabilis, previously found in the Atlantic and Pa- cific oceans, was confirmed for the first time in the Gulf of Mexico. This new record is represented by a juvenile male collected by a stratified plankton sampling at 800–1,000 m depth in the southern gulf (20.5°N, 94.5°W), during summer 2014. This is the first record of a pelagic nemertean in the gulf; extending our knowledge about the species’ global dis- tribution, the number of nemertean species present in Mexican Atlantic waters, and the need to explore the mesopelagic zone of the oceans. Key words: mesopelagic zone, ribbon worms, stratified sampling, western Atlantic, zooplankton The phylum Nemertea Schultze, 1851 is scarcely range of pelagic nemerteans encompasses deep waters. studied due to the difficulties in correctly identifying Among the pelagic nemerteans, the genus Nectonemertes species. Worldwide, the number of nemertean species Verril, 1892 is recognized as the most common and abun- currently recognized as valid is approximately 1,275 dant taxon, particularly the species Nectonemertes mirabilis (Kajihara et al. -
Los Nemertinos De España Y Portugal
LOS NEMERTINOS DE ESPAÑA Y PORTUGAL FACULTAD DE BIOLOGÍA, CIENCIAS AMBIENTALES Y QUÍMICA DEPARTAMENTO DE CIENCIAS DE LA VIDA Programa de Doctorado en Biología Evolutiva y Biodiversidad LOS NEMERTINOS DE ESPAÑA Y PORTUGAL Tesis Doctoral presentada por ALFONSO HERRERA BACHILLER Director Dr. Juan Junoy Alcalá de Henares, 2016 DEPARTAMENTO DE CIENCIAS DE LA VIDA Edificio de Ciencias. Campus Universitario 28805 Alcalá de Henares (Madrid) Tel.: 91 885 49927/4965 Fax: 91 8854929 / 5066 [email protected] El Dr. D. Juan Mª Junoy Pintos, Profesor Titular del Departamento de Ciencias de la Vida de la Universidad de Alcalá INFORMA Que D. Alfonso Herrera Bachiller, licenciado en Biología, ha realizado bajo su dirección y asesoramiento el presente trabajo titulado “LOS NEMER- TINOS DE ESPAÑA Y PORTUGAL”, que considera reúne las condiciones de calidad científica necesarias para optar al grado de Doctor por la Univer- sidad de Alcalá. De lo que informo en Alcalá de Henares, a de de 2016. UNIVERSIDAD DE ALCALÁ, PATRIMONIO DE LA HUMANIDAD DE LA ALCALÁ, PATRIMONIO UNIVERSIDAD DE Dr. Juan Junoy Pintos DEPARTAMENTO DE CIENCIAS DE LA VIDA Edificio de Ciencias. Campus Universitario 28805 Alcalá de Henares (Madrid) Tel.: 91 885 49927/4965 Fax: 91 8854929 / 5066 [email protected] D. Miguel Ángel Rodríguez Fernández, director en funciones del Depar- tamento de Ciencias de la Vida de la Universidad de Alcalá, Hace constar: Que el trabajo descrito en la presente memoria, titulado “ Los Nemer- tinos de España y Portugal”, ha sido realizado bajo la dirección de D. Juan Junoy Pintos en el Departamento de Ciencias de la Vida de la Universidad de Alcalá, dentro del Programa de Doctorado “Biología Evolutiva y Biodiversi- dad”, y reúne todos los requisitos necesarios para su aprobación como Tesis Doctoral. -
Multi-Level Convergence of Complex Traits and the Evolution of Bioluminescence Emily S
Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 15 December 2020 Multi-level convergence of complex traits and the evolution of bioluminescence Emily S. Lau* and Todd H. Oakley* Department of Ecology, Evolution, and Marine Biology, University of California Santa Barbara, Santa Barbara CA 93106 Corresponding authors information: Emily S. Lau: [email protected]; Todd H. Oakley: [email protected] Keywords multi‐level convergence | evolution | bioluminescence | biological organization | complex trait ABSTRACT Evolutionary convergence provides natural opportunities to investigate how, when, and why novel traits evolve. Many convergent traits are complex, highlighting the importance of explicitly considering convergence at different levels of biological organization, or ‘multi‐level convergent evolution’. To investigate multi‐level convergent evolution, we propose a holistic and hierarchical framework that emphasizes breaking down traits into several functional modules. We begin by identifying long‐standing questions on the origins of complexity and the diverse evolutionary processes underlying phenotypic convergence to discuss how they can be addressed by examining convergent systems. We argue that bioluminescence, a complex trait that evolved dozens of times through either novel mechanisms or conserved toolkits, is particularly well suited for these studies. We present an updated estimate of at least 94 independent origins of bioluminescence across the tree of life, which we calculated by reviewing and summarizing all estimates of independent origins. Then, we use our framework to review the biology, chemistry, and evolution of bioluminescence, and for each biological level identify questions that arise from our systematic review. We focus on luminous organisms that use the shared luciferin substrates coelenterazine or vargulin to produce light because these organisms convergently evolved bioluminescent proteins that use the same luciferins to produce bioluminescence.