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INSTITUTO POLITÉCNICO NACIONAL CENTRO INTERDISCIPLINARIO DE CIENCIAS MARINAS Departamento de Plancton y Ecología Marina TAXOCENOSIS DE MOLUSCOS HOLOPLANCTÓNICOS (MOLLUSCA:GASTROPODA) Y SU RELACIÓN BIOGEOGRÁFICA EN EL GOLFO DE CALIFORNIA TESIS Que para obtener el grado de Doctor en Ciencias Marinas P R E S E N T A M. en C. ORSO JUAN ANGULO CAMPILLO Director de Tesis Dr. Gerardo Aceves Medina La Paz, B.C.S., Junio del 2009 Dedico este trabajo a mi madre. Se lo dedico a mi familia que siempre me ha apoyado incondicionalmente. A mis profesores que gracias a sus enseñanzas he llegado hasta este punto. A mis amigos que solo ellos han podido aguantarme todo este tiempo. Y a todos aquellos que de una forma u otra han colaborado en este trabajo. AGRADECIMIENTOS Quiero agradecer a las autoridades del Instituto Politécnico Nacional, las autoridades del Centro Interdisciplinario de Ciencias Marinas y al Consejo Nacional de Ciencia y Tecnología por el apoyo otorgado para el desarrollo de este trabajo y de mi desarrollo académico. Al grupo trabajo de Plancton y Ecología Marina, en particular a la M. en C. Roxana De-Silva y M. en C. Raymundo Avedaño por su apoyo y facilitarme las muestras de los cruceros CGC, al M. en C. Marco Sanchéz Hidalgo, por sus comentarios. Al Dr. Roger Seapy por la ayuda en la identificación de los organismos. Al Dr. Jaime Gómez por sus comentarios, ayuda y permitirme participar durante las campañas CAPEGOLCA. Acrezco a los miembros del comité revisor por su paciencia y los consejos otorgados para concretar este trabajo. A los proyectos de los cuales obtuvieron las muestras de zooplancton: Detección y caracterización de las áreas de desove de peces de importancia Comercial en el Golfo de California, México (CGPI-2005053); Circulación del Golfo de California, Secretaria de Marina-Armada de México; Monitoreo de la producción secundaria, dinámica poblacional y parasitismo de especies clave del zooplancton mediante incubación in situ y métodos hidroacústicos en el Golfo de California (CONACyT-SAGARPA S007-2005-1-11717, CONACyT-FOSEMARNAT 2004-4- 01-144). ÍNDICE GENERAL ÍNDICE DE TABLAS iii ÍNDICE DE FIGURAS iv LISTADO DE ANEXOS xiii RESUMEN xiv ABSTRACT xv GLOSARIO 1 1. INTRODUCCIÓN 5 2. ANTECEDENTES 9 3. HIPÓTESIS 12 4. OBJETIVOS 12 4.1 General 12 4.2 Particulares 12 5. JUSTIFICACIÓN 13 6. ÁREA DE ESTUDIO 14 7. MATERIAL Y MÉTODO 16 7.1 Muestreos 16 7.2 Tratamiento de datos 18 7.2.1 Fichas taxonómicas 18 7.2.2 Descripción de comunidades. 19 8. RESULTADOS 22 8.1 Análisis Taxonómico. 22 8.1.1 Concha 22 8.1.2 Ojos 24 8.1.3 Opérculo 26 8.1.4 Rádula 27 8.2 Fichas Taxonómicas 29 8.2.1 Heteropoda 29 8.2.2 Cephalaspidea 79 8.2.3 Thecosomata 81 8.2.4 Gymnosomata 123 8.2.5 Nudibranchia 149 8.3 Descripción del ambiente físico (temperatura) 155 8.3.1 Temperatura superficial 155 8.3.2 Masas de agua 159 8.4 Descripción de la comunidad 161 8.4.1 Riqueza 161 8.4.2 Diversidad 167 8.4.3 Equidad 168 8.4.4 Abundancia 170 8.4.5 Distribución 173 i 8.5 Análisis de asociaciones 178 8.5.1 Grupo Cálido 182 8.5.2 Grupo Transicional 185 8.5.3 Grupo Frío 188 8.6 Bioregionalización del Golfo de California 191 8.6.1 Análisis de procedimiento de Multi-Respuesta 191 de Permutación (MRPP). 8.6.2 Análisis de Indicador de Especies (AEI). 192 9. DISCUSIÓN 194 10. CONCLUSIONES 214 11. RECOMENDACIONES 217 12. LITERATURA CITADA 218 ANEXOS 229 ii ÍNDICE DE TABLAS Tabla I. Nomenclatura, fecha y número de estaciones de los cruceros oceanográficos realizados en el Golfo de California en el periodo 2005-2007. CGC=Circulación del Golfo de California, GOLCA= Golfo de California. 16 Tabla II. Elenco taxonómico de moluscos holoplanctónicos durante el periodo de estudio, en el Golfo de California, México. 161 Tabla III. Nuevos registros de moluscos holoplanctónicos para el Golfo de California. 164 Tabla IV. Nuevos registros de Moluscos Holoplanctónicos para el Pacifico Americano. 164 Tabla V. Especies de Moluscos Holoplanctónicos exclusivas para cada una de las regiones. 166 Tabla VI. Abundancia de las especies que contribuyeron con más del 2% de la abundancia relativa, durante el periodo de estudio, por crucero y total 172 Tabla VII. Especies correspondientes a cada uno de los Grupos. 179 Tabla VIII. Resultados del análisis de Procedimiento Permutacional de Multirespuesta (MRPP), para cada uno de los grupos. 191 Tabla IX. Resultados del análisis AIE. Sólo se muestran aquellas especies que presentaron fidelidad a las regiones establecidas para el Grupo Calido (>25% y p=<0.05) 193 Tabla X. Resultados del análisis AIE. Sólo se muestran aquellas especies que presentaron fidelidad a las regiones establecidas para el Grupo Transicional (>25% y p=<0.05). 193 Tabla XI. Resultados del análisis AIE. Sólo se muestran aquellas especies que presentaron fidelidad a las regiones establecidas para el Grupo Frío (>25% y p=<0.05). 194 Tabla XII. Número de especies de moluscos holoplanctónicos registrados en: Mundo= alrededor del mundo, Pacífico= Océano pacifico Ecuatorial, GC= Golfo de California y Estudio: presente trabajo. 194 iii INDICE DE FIGURAS Figura. 1. Moluscos Holoplanctónicos. A)Atlantidae; B) Pterotraquidae;C) Pteropoda (Thecosomata); D) Gymnostomida y E) Nudibranchia 7 Figura. 2. Modelos de bioregionalización del Golfo de California. 11 Figura.3. Área de estudio. 14 Figura.4. Localización de las estaciones de muestreo. A) Cruceros CGC correspondientes a marzo, mayo y septiembre de 2005 y marzo de 2006, (CGC0503, CGC0505, CGC0509 y CGC0603 respectivamente; B) Cruceros GOLCA correspondientes a noviembre de 2005, enero y julio de 2007 (GOLCA0511, GOLCA0707 y GOLCA0707 respectivamente). 17 Figura. 5. Caracteres taxonómicos del Grupo de los Atlantidos. A) Posición y forma generalizada de la quilla y de la ultima espiral. B) Posición del plano de la espiral. C) Determinación del número de espirales, a partir de un ángulo de 90°. Esquemas modificados a partir de Tokioka, 1955 y Seapy, 1990. 22 Figura.6. Caracteres taxonómicos del Grupo de los Atlantidos. A) Sutura. B) Tipos de ornamentaciones. C) Inclinación de la espira. Esquemas tomados de Spoel et al., 1997. 23 Figura.7. Morfología de la concha dentro de los pteropodos. A) Enrollada; B) semi-rectas o clavo; C) Triangular y D) Lobulada. Modificado de Spoel & Boltovskoy, 1999. Abreviaturas: cl=columnela; cor=cornisa; csd= costilla dorsal; csl=costilla lateral; cr=cresta; est= estrias; spc=espina caudal; spl= espina lateral. 24 Figura.8 Tipos de morfología ocular de los atlantidos. Tomado de Seapy 1990b. 25 Figura.9. Tipos de morfología ocular de los pterotraquidos. Tomado de Richter & Seapy, 1999. 26 Figura. 10. Morfología opercular de los atlántidos. Tomado de Richter & Seapy (1999). 27 Figura.11. Forma de la rádula y ejemplos de los tipos de dientes. A) Dientes marginales; B) Dientes laterales y C) Diente raquídeo. 28 iv Figura.12. Caracteres taxonómicos adicionales empleados en la determinación de especies A) Carinoideos; B) Gymnostomidos; C) Nudibranquio. Abreviaturas: al=aletas; br=branquias; brl=branquia lateral; bri= branquia inferior; cl= cola; cn=concha; cr=cromatoforos; lbc=lóbulo central; lbl= lóbulo lateral; tc=tentáculos cefálicos; vn= ventosa. 28 Figura.13. Atlanta californiensis. a) organismo preservado, b) organismo trasparentado, donde se observa la posición de la rádula, c) Acercamiento de la espira (SEM), d) rádula, e) distribución. Fotografías c y d, tomadas de Seapy & Richter, 1993 30 Figura.14. Atlanta echinogyra. a) concha, b) acercamiento de la espira, c) rádula, d) distribución. Fotografías: (b) Seapy 1990, (c) Richter, 1986 32 Figura.15. Atlanta fragilis. a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución. 34 Figura.16. Atlanta frontieri. a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución. 36 Figura.17. Atlanta fusca. a) concha, b) acercamiento de la espiras vista superior, c) vista lateral de la concha d) espiras vista lateral, e) distribución. Fotografías tomadas de Seapy, 1990 38 Figura.18. Atlanta gaudichaudi a) concha b) acercamiento de la espira, c) vista lateral de la concha d) dientes, e) distribución. Fotografías: tomadas de Seapy (1990). 40 Figura.19. Atlanta helicinoides. a) concha, b) acercamiento de la espira, c) dientes, d) rádula, e) distribución. 42 Figura.20. Atlanta inclinata. a) concha, b) acercamiento de la espira, c) dientes, d) rádula, e) distribución. Fotografías: a) y b) Seapy et al., (2003), d) Spoel et al., (1997). 44 Figura.21. Atlanta inflata. a) concha, b) acercamiento de la espira, c) distribución. Fotografías: a) y b) Spoel et al., 1997. 46 Figura.22. Atlanta lesueuri. a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución. Fotografías: a) Seapy, (1990), d) Spoel et al.,(1997). 48 v Figura.23. Atlanta meteori. a) concha, b) acercamiento de la espira, c) juvenil, d) distribución. Fotografías: a y b) Seapy (1990). 50 Figura.24. Atlanta oligogyra. a) concha, b) acercamiento de la espira, c) distribución. 52 Figura.25. Atlanta peroni. a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución. 54 Figura.26. Atlanta plana a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución 56 Figura.27. Atlanta tokiokai a) concha, b) acercamiento de la espira, c) rádula, d) dientes, e) distribución. 58 Figura.28. Atlanta turriculata a) concha, b) acercamiento de la espira, d) distribución. 60 Figura.29. Protoatlanta souleyeti a) concha, b) acercamiento de la espira, c) Vista apical de la espiral, d) dientes, e) distribución. Fotografías: a-c) Seapy (1990), d) Spoel et al.,(1997). 62 Figura.30. Oxygyrus keraudreni a) organismo preservado, b) Acercamiento de la espira, c) rádula, d) dientes, e) distribución. Fotografías: a-d Spoel et al.,(1997).
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  • Atlanta Ariejansseni, a New Species of Shelled Heteropod

    Atlanta Ariejansseni, a New Species of Shelled Heteropod

    A peer-reviewed open-access journal ZooKeys 604: 13–30 (2016) Atlanta ariejansseni, a new species of shelled heteropod.... 13 doi: 10.3897/zookeys.604.8976 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Atlanta ariejansseni, a new species of shelled heteropod from the Southern Subtropical Convergence Zone (Gastropoda, Pterotracheoidea) Deborah Wall-Palmer1,2, Alice K. Burridge2,3, Katja T.C.A. Peijnenburg2,3 1 School of Geography, Earth and Environmental Sciences, Plymouth University, Drake Circus, Plymouth, PL4 8AA, UK 2 Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands3 Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P. O. Box 94248, 1090 GE Amster- dam, The Netherlands Corresponding author: Deborah Wall-Palmer ([email protected]) Academic editor: N. Yonow | Received 21 April 2016 | Accepted 22 June 2016 | Published 11 July 2016 http://zoobank.org/09E534C5-589D-409E-836B-CF64A069939D Citation: Wall-Palmer D, Burridge AK, Peijnenburg KTCA (2016) Atlanta ariejansseni, a new species of shelled heteropod from the Southern Subtropical Convergence Zone (Gastropoda, Pterotracheoidea). ZooKeys 604: 13–30. doi: 10.3897/zookeys.604.8976 Abstract The Atlantidae (shelled heteropods) is a family of microscopic aragonite shelled holoplanktonic gastro- pods with a wide biogeographical distribution in tropical, sub-tropical and temperate waters. The arago- nite shell and surface ocean habitat of the atlantids makes them particularly susceptible to ocean acidifica- tion and ocean warming, and atlantids are likely to be useful indicators of these changes. However, we still lack fundamental information on their taxonomy and biogeography, which is essential for monitoring the effects of a changing ocean.
  • Time-Calibrated Molecular Phylogeny of Pteropods

    Time-Calibrated Molecular Phylogeny of Pteropods

    RESEARCH ARTICLE Time-calibrated molecular phylogeny of pteropods Alice K. Burridge1,2☯, Christine HoÈ rnlein2,3, Arie W. Janssen1, Martin Hughes2,4, Stephanie L. Bush5,6, Ferdinand MarleÂtaz7, Rebeca Gasca8, Annelies C. Pierrot-Bults1,2, Ellinor Michel4, Jonathan A. Todd4, Jeremy R. Young9, Karen J. Osborn5,6, Steph B. J. Menken2, Katja T. C. A. Peijnenburg1,2☯* 1 Naturalis Biodiversity Center, Leiden, The Netherlands, 2 Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, Amsterdam, The Netherlands, 3 Koninklijk Nederlands Instituut voor Onderzoek der Zee (NIOZ), Yerseke, The Netherlands, 4 Natural History Museum (NHM), Cromwell a1111111111 Road, London, United Kingdom, 5 Smithsonian Institution National Museum of Natural History, Washington a1111111111 DC, United States of America, 6 Monterey Bay Aquarium Research Institute (MBARI), Moss Landing, a1111111111 California, United States of America, 7 Molecular Genetics Unit, Okinawa Institute of Science and a1111111111 Technology, Onna-son, Japan, 8 El Colegio de la Frontera Sur (ECOSUR), Unidad Chetumal, Quintana Roo, a1111111111 Chetumal, Mexico, 9 Department of Earth Sciences, University College London, London, United Kingdom ☯ These authors contributed equally to this work. * [email protected], [email protected] OPEN ACCESS Abstract Citation: Burridge AK, HoÈrnlein C, Janssen AW, Hughes M, Bush SL, MarleÂtaz F, et al. (2017) Time- Pteropods are a widespread group of holoplanktonic gastropod molluscs and are uniquely calibrated molecular phylogeny of pteropods. PLoS suitable for study of long-term evolutionary processes in the open ocean because they are the ONE 12(6): e0177325. https://doi.org/10.1371/ journal.pone.0177325 only living metazoan plankton with a good fossil record.
  • Describing Species

    Describing Species

    DESCRIBING SPECIES Practical Taxonomic Procedure for Biologists Judith E. Winston COLUMBIA UNIVERSITY PRESS NEW YORK Columbia University Press Publishers Since 1893 New York Chichester, West Sussex Copyright © 1999 Columbia University Press All rights reserved Library of Congress Cataloging-in-Publication Data © Winston, Judith E. Describing species : practical taxonomic procedure for biologists / Judith E. Winston, p. cm. Includes bibliographical references and index. ISBN 0-231-06824-7 (alk. paper)—0-231-06825-5 (pbk.: alk. paper) 1. Biology—Classification. 2. Species. I. Title. QH83.W57 1999 570'.1'2—dc21 99-14019 Casebound editions of Columbia University Press books are printed on permanent and durable acid-free paper. Printed in the United States of America c 10 98765432 p 10 98765432 The Far Side by Gary Larson "I'm one of those species they describe as 'awkward on land." Gary Larson cartoon celebrates species description, an important and still unfinished aspect of taxonomy. THE FAR SIDE © 1988 FARWORKS, INC. Used by permission. All rights reserved. Universal Press Syndicate DESCRIBING SPECIES For my daughter, Eliza, who has grown up (andput up) with this book Contents List of Illustrations xiii List of Tables xvii Preface xix Part One: Introduction 1 CHAPTER 1. INTRODUCTION 3 Describing the Living World 3 Why Is Species Description Necessary? 4 How New Species Are Described 8 Scope and Organization of This Book 12 The Pleasures of Systematics 14 Sources CHAPTER 2. BIOLOGICAL NOMENCLATURE 19 Humans as Taxonomists 19 Biological Nomenclature 21 Folk Taxonomy 23 Binomial Nomenclature 25 Development of Codes of Nomenclature 26 The Current Codes of Nomenclature 50 Future of the Codes 36 Sources 39 Part Two: Recognizing Species 41 CHAPTER 3.