DOCSLIB.ORG

Annelida: the Metameric Body Form 221

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text

Load more

12 Annelida:The Metameric Body Form The Samoan palolo worm (Eunice viridis) is one of approximately15,000 spe­ cies in the phylum Annelida. The palolo's 1·eproductive habits are unusual, but effective.As you will see in this chapter, "unusual" is not the exceptionfor this Chapter Outline taxonomicallychallenging phylum. 12.1 Evolutionary Perspective Relationships to Other Animals Metamerism and Tagmatization 12.2 Annelid Structure and Function 12. 1 EVOLUTIONARY PERSPECTIVE External Structure and Locomotion Feeding and the Digestive System Gas Exchange and Circulation LEARNING OUTCOMES Nervous and SensoryFunctions Excretion 1. Describe the relationships of members of the Annelida to other animal phyla. Regeneration, Reproduction, 2. Explain the benefits of metamerism for an annelid and Development 12.3 Clade (Class) Errantia Nereis (Neanthes, Alitta) At the time of the November full moon on islands near Samoa in the South Glycera Pacific, people rush about preparing for one of their biggest yearly feasts. In Fireworms just one week, the sea will yield a harvest that can be scooped up in nets and 12.4 Clade (Class) Sedentaria Tubeworms buckets (figure 12.1). Worms by the millions transform the ocean into what one Siboglinidae writer called "vermicelli soup!" Celebrants gorge themselves on worms that have Echiura been cooked or wrapped in breadfruit leaves.The Samoan palolo worm (Eunice Clade Clitel/ata 12.5 Basal Annelid Groups viridis, alternatively Palo/a viridis) spends its entire adult life in coral burrows Chaetopteridae at the sea bottom. Each November, one week after the full moon, this worm Sipuncula emerges from its burrow, and specialized body segments devoted to sexual 12.6 Further Phylogenetic Considerations reproduction break free and float to the surface, while the rest of the worm is safe on the ocean floor. The surface water is discolored as gonads release their countless eggs and sperm. The natives' feast is short lived, however; these reproductive swarms last only two days and do not recur for another year. The Samoan palolo worm is a member of the phylum Annelida (ah-nel' i-dah) (L. annellus, ring). Other members of this phylum include countless marine worms, the soil-building earthworms, and predatory leeches (table 12.1). Characteristics of the phylum Annelida include: 1. Body metameric, bilaterally symmetrical, and worm-like 2. Spiral cleavage, trochophore larvae (when larvae are present), and schizocoelous coelom formation 3. Paired, epidermal setae (chaetae) 4. Closed circulatory system 5. Dorsal suprapharyngeal ganglia and ventral nerve cord(s) with ganglia 6. Metanephridia (usually) or protonephridia Relationships to Other Animals It has been clear for many years that Annelida is a monophyletic assemblage of marine, freshwater, and terrestrial worms. They are lophotrochozoans and, thus, share common ancestry with Mollusca, Brachiopoda, Bryozoa, Nemertea, and others Annelida: The Metameric Body Form 221 (a) (b) FIGURE 12.1 Palolo Feasts. (a) Women from the Nggela Group (Florida Islands) of the Solomon Islands use light from a torch to attract reproductive segments of Eunice viridis (family Eunicidae) during collecting. Photoreceptors on the reproductive segments elicit the response of the worms to light. (b) A swarm of worms as viewed through the lens of a diver's camera. The photograph on page 220 shows palolo worms (referred to by islanders as "odu") ready for feasting. (Pbotograpbs courte.,y of Dr. Si111011 Foale, Principle Resectrcb Fe/10111, ARC Ce11tre of Exce/le11cefor Coral Req( Studies.James Cook University, Quee11sla11d, Austmlio.) Basal Protists Phyla FIGURE 12.2 Evolutionary Relationships of Annelids to Other Animals. This figure shows an interpretation of the relationship of the Annelida to other members of the animal kingdom. The relationships depicted here are based on evidence from developmental and molecular biology. Annelids are placed within the Lophotrochozoa along with the Mollusca, Plathyhelminthes, Rotifera, and others (see pages xvi-xvii). The phylum includes approximately 15,000 species of segmented worms. Most of these are marine and traditionally classified into the class Polychaeta. As discussed in the text, this placement is being reevaluated based on evidence from molecular biology. The Christmas-tree worm (Spirohrcmcbus gigan/e11s) (family Serpulidae) shown here is a memher of the clade Senclentaria The spiral fans of this tube-dwelling annelid are derived from prostomial palps that surround the mouth and are specialized for feeding and gas exchange. 222 CHAPTER TWELVE to differentiate groups within the polychaetes have been res­ TABLE 12.1 urrected to represent two major annelid clades: Errantia and CLASSIFICATION OF THE PHYLUM ANNELIDA Sedentaria. Sedentaria now includes some "polychaetes," leeches, earthworms, and even worms that were formerly Phylum Annelida (ah-nel'i-dah) considered separate phyla (Echiura and Pogonophora). Cli­ The phylum of triploblastic, coelomate animals whose members tellata is still a valid clade within the Sendentaria composed are metameric (segmented), elongate, and cylindrical or of earthworms, leeches, and a few smaller taxa. The leeches oval in cross section. Annelids have a complete digestive form a monophyletic clade (Hirudinea) within the Clitellata. tract; paired, epidermal setae; and a ventral nerve cord. The earthworms and their relatives, however, are not mono­ Approximately 15,000 species of annelids have been phyletic. Thus the name "Oligochaeta," which was formerly described. considered a subclass name within the Clitellata, should Clade (Class) Errantia (er-ran'tiah) be abandoned as a taxonomic designation. Two other Marine annelids; parapodia with prominent lobes supported groups lie outside of Sedentaria and Errantia. One of these, by internalized chaetae and ventral cirri; palps well Chaetopteridae, was formerly considered to be a family developed. Nereis, Eunice, Glycera. within the "Polychaeta." The other, Sipuncula, was another Clade (Class) Sedentaria (sed-en-ter'iah) phylum outside of Annelida. The hierarchical taxonomic­ Marine, freshwater, and terrestrial annelids; parapodia with categories associated with these clades have not been estab­ reduced lobes or parapodia lacking; setae associated lished. We treat Errantia and Sedentaria as clades with a with the stiff body wall to facilitate anchoring in tubes and burrows; palps reduced. Includes the clade Clitellata, parenthetical "class" designation, which seems a logical out­ echiruans, and siboglinids. Arenicola, Riftia, Lumbricus, come of ongoing and future research. Hirudo Chapter 12 is now organized to reflect this new phylo­ Other Annelid Taxa genetic work. In the next section, we describe metamerism Sipuncula (si-pun'ku-lah) and tagmatization. This discussion is followed by coverage Marine; unsegmented body; retractable anterior trunk of annelid structure and function. The term "polychaete" is called an introvelt. Formerly phylum Sipuncula. Inclusion used in a nontaxonomic sense to refer to a host of marine of this group within the Annelida remains somewhat annelids (both errantians and sedentarians) when adap­ controversial. Themiste. tations related to their common marine habitat result in Chaetopteridae (ke-top-ter' ida) similar structures and functions. After the basics of anne­ Marine; three-palt body; lives in a U-shaped tube, lid structure and function are discussed, the two major appendages for feeding and creating water currents. annelid clades and the two outlying groups are described. Chaetoptei·us. Since members of the clade Clitellata are different in many respects from other sedentarians, unique aspects of their structure and function are described in the section on Sed­ entaria. As always, this chapter ends with "Further Phyloge­ netic Considerations." (figure 12. 2 and see chapters 10 and 11). Our understanding of phylogeny within the phylum, however, has a history of contentious debate. The application of modern phylogenetic analysis is helping unravel the annelid taxonomic-tangle. Metamerism and Tagmatization Anyone who has followed the debates involved with elu­ Earthworm bodies are organized into a series of ringlike seg­ cidating annelid phylogeny will have a better appreciation ments. What is not externally obvious, however, is that the of the dynamics within the field of animal taxonomy. These body is divided internally as well. Segmental arrangement of debates underscore the importance of taxonomy in helping body parts in an animal is called metamerism (Gr. meta, zoologists understand the evolution of bilateral morphology after + mere, part). (see the discussion of metamerism and tagmatization in the Metamerism profoundly influences virtually eve1y next section). aspect of annelid structure and function, such as the anatomi­ Previous editions of this book described the Annelida cal arrangement of organs that are coincidentally associated as being composed of two classes: Polychaeta (marine with metamerism. For example, the compartmentalization worms) and Clitellata (leeches, earthworms, and others). of the body has resulted in each segment having its own While "Polychaeta" was rightly described as being paraphy­ excreto1y, nervous, and circulato1y structures. In most mod­ letic, recent molecular phylogenetic work has shown that ern annelids two related functions are probably the primary the entire annelid assemblage is encompassed by what was adaptive features of metamerism: flexible support and effi­ being called "Polychaeta." As will
Recommended publications
  • Systema Naturae∗

    Systema Naturae∗

    Systema Naturae∗ c Alexey B. Shipunov v. 5.802 (June 29, 2008) 7 Regnum Monera [ Bacillus ] /Bacteria Subregnum Bacteria [ 6:8Bacillus ]1 Superphylum Posibacteria [ 6:2Bacillus ] stat.m. Phylum 1. Firmicutes [ 6Bacillus ]2 Classis 1(1). Thermotogae [ 5Thermotoga ] i.s. 2(2). Mollicutes [ 5Mycoplasma ] 3(3). Clostridia [ 5Clostridium ]3 4(4). Bacilli [ 5Bacillus ] 5(5). Symbiobacteres [ 5Symbiobacterium ] Phylum 2. Actinobacteria [ 6Actynomyces ] Classis 1(6). Actinobacteres [ 5Actinomyces ] Phylum 3. Hadobacteria [ 6Deinococcus ] sed.m. Classis 1(7). Hadobacteres [ 5Deinococcus ]4 Superphylum Negibacteria [ 6:2Rhodospirillum ] stat.m. Phylum 4. Chlorobacteria [ 6Chloroflexus ]5 Classis 1(8). Ktedonobacteres [ 5Ktedonobacter ] sed.m. 2(9). Thermomicrobia [ 5Thermomicrobium ] 3(10). Chloroflexi [ 5Chloroflexus ] ∗Only recent taxa. Viruses are not included. Abbreviations and signs: sed.m. (sedis mutabilis); stat.m. (status mutabilis): s., aut i. (superior, aut interior); i.s. (incertae sedis); sed.p. (sedis possibilis); s.str. (sensu stricto); s.l. (sensu lato); incl. (inclusum); excl. (exclusum); \quotes" for environmental groups; * (asterisk) for paraphyletic taxa; / (slash) at margins for major clades (\domains"). 1Incl. \Nanobacteria" i.s. et dubitativa, \OP11 group" i.s. 2Incl. \TM7" i.s., \OP9", \OP10". 3Incl. Dictyoglomi sed.m., Fusobacteria, Thermolithobacteria. 4= Deinococcus{Thermus. 5Incl. Thermobaculum i.s. 1 4(11). Dehalococcoidetes [ 5Dehalococcoides ] 5(12). Anaerolineae [ 5Anaerolinea ]6 Phylum 5. Cyanobacteria [ 6Nostoc ] Classis 1(13). Gloeobacteres [ 5Gloeobacter ] 2(14). Chroobacteres [ 5Chroococcus ]7 3(15). Hormogoneae [ 5Nostoc ] Phylum 6. Bacteroidobacteria [ 6Bacteroides ]8 Classis 1(16). Fibrobacteres [ 5Fibrobacter ] 2(17). Chlorobi [ 5Chlorobium ] 3(18). Salinibacteres [ 5Salinibacter ] 4(19). Bacteroidetes [ 5Bacteroides ]9 Phylum 7. Spirobacteria [ 6Spirochaeta ] Classis 1(20). Spirochaetes [ 5Spirochaeta ] s.l.10 Phylum 8. Planctobacteria [ 6Planctomyces ]11 Classis 1(21).
  • Conservation and Diversification of Appendage Identity Specification Mechanisms Along the Anteroposterior and Proximodistal Axes in Panarthropoda Frank W

    Conservation and Diversification of Appendage Identity Specification Mechanisms Along the Anteroposterior and Proximodistal Axes in Panarthropoda Frank W

    University of Connecticut OpenCommons@UConn Doctoral Dissertations University of Connecticut Graduate School 8-23-2013 Conservation and Diversification of Appendage Identity Specification Mechanisms Along the Anteroposterior and Proximodistal Axes in Panarthropoda Frank W. Smith III University of Connecticut, [email protected] Follow this and additional works at: https://opencommons.uconn.edu/dissertations Recommended Citation Smith, Frank W. III, "Conservation and Diversification of Appendage Identity Specification Mechanisms Along the Anteroposterior and Proximodistal Axes in Panarthropoda" (2013). Doctoral Dissertations. 161. https://opencommons.uconn.edu/dissertations/161 Conservation and Diversification of Appendage Identity Specification Mechanisms Along the Anteroposterior and Proximodistal Axes in Panarthropoda Frank Wesley Smith III, PhD University of Connecticut, 2013 A key characteristic of arthropods is their diverse serially homologous segmented appendages. This dissertation explores diversification of these appendages, and the developmental mechanisms producing them. In Chapter 1, the roles of genes that specify antennal identity in Drosophila melanogaster were investigated in the flour beetle Tribolium castaneum. Antenna-to-leg transformations occurred in response to RNA interference (RNAi) against homothorax, extradenticle, spineless and Distal-less. However, for homothorax/extradenticle RNAi, the extent of transformation along the proximodistal axis differed between embryogenesis and metamorphosis. This suggests that distinct mechanisms specify antennal identity during flour beetle embryogenesis and metamorphosis and leads to a model for the evolution of the Drosophila antennal identity mechanism. Homothorax/Extradenticle acquire many of their identity specification roles by acting as Hox cofactors. In chapter 2, the metamorphic roles of the Hox genes, extradenticle, and homothorax were compared in T. castaneum. homothorax/extradenticle RNAi and Hox RNAi produced similar body wall phenotypes but different appendage phenotypes.
  • RECORDS of the HAWAII BIOLOGICAL SURVEY for 1995 Part 2: Notes1

    RECORDS of the HAWAII BIOLOGICAL SURVEY for 1995 Part 2: Notes1

    RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1995 Part 2: Notes1 This is the second of two parts to the Records of the Hawaii Biological Survey for 1995 and contains the notes on Hawaiian species of plants and animals including new state and island records, range extensions, and other information. Larger, more compre- hensive treatments and papers describing new taxa are treated in the first part of this Records [Bishop Museum Occasional Papers 45]. New Hawaiian Pest Plant Records for 1995 PATRICK CONANT (Hawaii Dept. of Agriculture, Plant Pest Control Branch, 1428 S King St, Honolulu, HI 96814) Fabaceae Ulex europaeus L. New island record On 6 October 1995, Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife employee C. Joao submitted an unusual plant he found while work- ing in the Molokai Forest Reserve. The plant was identified as U. europaeus and con- firmed by a Hawaii Department of Agriculture (HDOA) nox-A survey of the site on 9 October revealed an infestation of ca. 19 m2 at about 457 m elevation in the Kamiloa Distr., ca. 6.2 km above Kamehameha Highway. Distribution in Wagner et al. (1990, Manual of the flowering plants of Hawai‘i, p. 716) listed as Maui and Hawaii. Material examined: MOLOKAI: Molokai Forest Reserve, 4 Dec 1995, Guy Nagai s.n. (BISH). Melastomataceae Miconia calvescens DC. New island record, range extensions On 11 October, a student submitted a leaf specimen from the Wailua Houselots area on Kauai to PPC technician A. Bell, who had the specimen confirmed by David Lorence of the National Tropical Botanical Garden as being M.
  • Systematics , Zoogeography , Andecologyofthepriapu

    Systematics , Zoogeography , Andecologyofthepriapu

    SYSTEMATICS, ZOOGEOGRAPHY, AND ECOLOGY OF THE PRIAPULIDA by J. VAN DER LAND (Rijksmuseum van Natuurlijke Historie, Leiden) With 89 text-figures and five plates CONTENTS Ι Introduction 4 1.1 Plan 4 1.2 Material and methods 5 1.3 Acknowledgements 6 2 Systematics 8 2.1 Historical introduction 8 2.2 Phylum Priapulida 9 2.2.1 Diagnosis 10 2.2.2 Classification 10 2.2.3 Definition of terms 12 2.2.4 Basic adaptive features 13 2.2.5 General features 13 2.2.6 Systematic notes 2 5 2.3 Key to the taxa 20 2.4 Survey of the taxa 29 Priapulidae 2 9 Priapulopsis 30 Priapulus 51 Acanthopriapulus 60 Halicryptus 02 Tubiluchidae 7° Tubiluchus 73 3 Zoogeography and ecology 84 3.1 Distribution 84 3.1.1 Faunistics 90 3.1.2 Expeditions 93 3.2 Ecology 96 3.2.1 Substratum 96 3.2.2 Depth 97 3.2.3 Temperature 97 3.24 Salinity 98 3.2.5 Oxygen 98 3.2.6 Food 99 3.2.7 Predators 100 3.2.8 Communities 102 3.3 Theoretical zoogeography 102 3.3.1 Bipolarity 102 3.3.2 Relict distribution 103 3.3.3 Concluding remarks 104 References 105 4 ZOOLOGISCHE VERHANDELINGEN 112 (1970) 1 INTRODUCTION The phylum Priapulida is only a very small group of marine worms but, since these animals apparently represent the last remnants of a once un- doubtedly much more important animal type, they are certainly of great scientific interest. Therefore, it is to be regretted that a comprehensive review of our present knowledge of the group does not exist, which does not only cause the faulty way in which the Priapulida are treated usually in textbooks and works on phylogeny, but also hampers further studies.
  • New Zealand's Genetic Diversity

    New Zealand's Genetic Diversity

    1.13 NEW ZEALAND’S GENETIC DIVERSITY NEW ZEALAND’S GENETIC DIVERSITY Dennis P. Gordon National Institute of Water and Atmospheric Research, Private Bag 14901, Kilbirnie, Wellington 6022, New Zealand ABSTRACT: The known genetic diversity represented by the New Zealand biota is reviewed and summarised, largely based on a recently published New Zealand inventory of biodiversity. All kingdoms and eukaryote phyla are covered, updated to refl ect the latest phylogenetic view of Eukaryota. The total known biota comprises a nominal 57 406 species (c. 48 640 described). Subtraction of the 4889 naturalised-alien species gives a biota of 52 517 native species. A minimum (the status of a number of the unnamed species is uncertain) of 27 380 (52%) of these species are endemic (cf. 26% for Fungi, 38% for all marine species, 46% for marine Animalia, 68% for all Animalia, 78% for vascular plants and 91% for terrestrial Animalia). In passing, examples are given both of the roles of the major taxa in providing ecosystem services and of the use of genetic resources in the New Zealand economy. Key words: Animalia, Chromista, freshwater, Fungi, genetic diversity, marine, New Zealand, Prokaryota, Protozoa, terrestrial. INTRODUCTION Article 10b of the CBD calls for signatories to ‘Adopt The original brief for this chapter was to review New Zealand’s measures relating to the use of biological resources [i.e. genetic genetic resources. The OECD defi nition of genetic resources resources] to avoid or minimize adverse impacts on biological is ‘genetic material of plants, animals or micro-organisms of diversity [e.g. genetic diversity]’ (my parentheses).
  • A Phylogenetic Analysis of the Genus Eunice (Eunicidae, Polychaete, Annelida)

    A Phylogenetic Analysis of the Genus Eunice (Eunicidae, Polychaete, Annelida)

    Blackwell Publishing LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082© 2007 The Linnean Society of London? 2007 1502 413434 Original Article PHYLOGENY OF EUNICEJ. ZANOL ET AL. Zoological Journal of the Linnean Society, 2007, 150, 413–434. With 12 figures A phylogenetic analysis of the genus Eunice (Eunicidae, polychaete, Annelida) JOANA ZANOL1*, KRISTIAN FAUCHALD2 and PAULO C. PAIVA3 1Pós-Graduação em Zoologia, Museu Nacional/UFRJ, Quinta da Boa Vista s/n°, São Cristovão, Rio de Janeiro, RJ 20940–040, Brazil 2Department of Invertebrate Zoology, NMNH, Smithsonian Institution, PO Box 37012, NHB MRC 0163, Washington, DC 20013–7012, USA 3Departamento de Zoologia, Insituto de Biologia, Universidade Federal do Rio de Janeiro, CCS, Bloco A, Sala A0-104, Ilha do Fundão, Rio de Janeiro, RJ 2240–590, Brazil Received April 2006; accepted for publication December 2006 Species of Eunice are distributed worldwide, inhabiting soft and hard marine bottoms. Some of these species play sig- nificant roles in coral reef communities and others are commercially important. Eunice is the largest and most poorly defined genus in Eunicidae. It has traditionally been subdivided in taxonomically informal groups based on the colour and dentition of subacicular hooks, and branchial distribution. The monophyly of Eunice and of its informal subgroups is tested here using cladistic analyses of 24 ingroup species based on morphological data. In the phylo- genetic hypothesis resulting from the present analyses Eunice and its subgroups are paraphyletic; the genus may be divided in at least two monophyletic groups, Eunice s.s. and Leodice, but several species do not fall inside these two groups.
  • Number of Living Species in Australia and the World

    Number of Living Species in Australia and the World

    Numbers of Living Species in Australia and the World 2nd edition Arthur D. Chapman Australian Biodiversity Information Services australia’s nature Toowoomba, Australia there is more still to be discovered… Report for the Australian Biological Resources Study Canberra, Australia September 2009 CONTENTS Foreword 1 Insecta (insects) 23 Plants 43 Viruses 59 Arachnida Magnoliophyta (flowering plants) 43 Protoctista (mainly Introduction 2 (spiders, scorpions, etc) 26 Gymnosperms (Coniferophyta, Protozoa—others included Executive Summary 6 Pycnogonida (sea spiders) 28 Cycadophyta, Gnetophyta under fungi, algae, Myriapoda and Ginkgophyta) 45 Chromista, etc) 60 Detailed discussion by Group 12 (millipedes, centipedes) 29 Ferns and Allies 46 Chordates 13 Acknowledgements 63 Crustacea (crabs, lobsters, etc) 31 Bryophyta Mammalia (mammals) 13 Onychophora (velvet worms) 32 (mosses, liverworts, hornworts) 47 References 66 Aves (birds) 14 Hexapoda (proturans, springtails) 33 Plant Algae (including green Reptilia (reptiles) 15 Mollusca (molluscs, shellfish) 34 algae, red algae, glaucophytes) 49 Amphibia (frogs, etc) 16 Annelida (segmented worms) 35 Fungi 51 Pisces (fishes including Nematoda Fungi (excluding taxa Chondrichthyes and (nematodes, roundworms) 36 treated under Chromista Osteichthyes) 17 and Protoctista) 51 Acanthocephala Agnatha (hagfish, (thorny-headed worms) 37 Lichen-forming fungi 53 lampreys, slime eels) 18 Platyhelminthes (flat worms) 38 Others 54 Cephalochordata (lancelets) 19 Cnidaria (jellyfish, Prokaryota (Bacteria Tunicata or Urochordata sea anenomes, corals) 39 [Monera] of previous report) 54 (sea squirts, doliolids, salps) 20 Porifera (sponges) 40 Cyanophyta (Cyanobacteria) 55 Invertebrates 21 Other Invertebrates 41 Chromista (including some Hemichordata (hemichordates) 21 species previously included Echinodermata (starfish, under either algae or fungi) 56 sea cucumbers, etc) 22 FOREWORD In Australia and around the world, biodiversity is under huge Harnessing core science and knowledge bases, like and growing pressure.
  • Annelida, Polychaeta, Chaetopteridae), with Re- Chaetopteridae), with Re-Description of M

    Annelida, Polychaeta, Chaetopteridae), with Re- Chaetopteridae), with Re-Description of M

    2 We would like to thank the Zoological Journal of the Linnean Society, The Linnean Society of London and Blackwell Publishing for accepting our manuscript entitled “Description of a Description of a new species of Mesochaetopterus (Annelida, Polychaeta, new species of Mesochaetopterus (Annelida, Polychaeta, Chaetopteridae), with re- Chaetopteridae), with re-description of M. xerecus and an approach to the description of M. xerecus and an approach to the phylogeny of the family”, which has phylogeny of the family been published in the Journal issue Zool. J. Linnean Soc. 2008, 152: 201–225. D. MARTIN1,* J. GIL1, J. CARRERAS-CARBONELL1 and M. BHAUD2 By posting this version of the manuscript (i.e. pre-printed), we agree not to sell or reproduce the Article or any part of it for commercial purposes (i.e. for monetary gain on your own 1Centre d'Estudis Avançats de Blanes (CSIC), Carrer d’accés a la Cala Sant Francesc 14, account or on that of a third party, or for indirect financial gain by a commercial entity), and 17300 Blanes (Girona), Catalunya (Spain). we expect the same from the users. 2 Observatoire Océanologique de Banyuls, Université P. et M. Curie - CNRS, BP 44, 66650 As soon as possible, we will add a link to the published version of the Article at the editors Banyuls-sur-Mer, Cedex, France. web site. * Correspondence author: Daniel Martin. Centre d'Estudis Avançats de Blanes (CSIC), Carrer Daniel Martin, Joao Gil, Michel Bhaud & Josep Carreras-Carbonell d’accés a la Cala Sant Francesc 14, 17300 Blanes (Girona), Catalunya (Spain). Tel. +34972336101; Fax: +34 972337806; E-mail: [email protected].
  • DEEP SEA LEBANON RESULTS of the 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project

    DEEP SEA LEBANON RESULTS of the 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project

    DEEP SEA LEBANON RESULTS OF THE 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project March 2018 DEEP SEA LEBANON RESULTS OF THE 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project Citation: Aguilar, R., García, S., Perry, A.L., Alvarez, H., Blanco, J., Bitar, G. 2018. 2016 Deep-sea Lebanon Expedition: Exploring Submarine Canyons. Oceana, Madrid. 94 p. DOI: 10.31230/osf.io/34cb9 Based on an official request from Lebanon’s Ministry of Environment back in 2013, Oceana has planned and carried out an expedition to survey Lebanese deep-sea canyons and escarpments. Cover: Cerianthus membranaceus © OCEANA All photos are © OCEANA Index 06 Introduction 11 Methods 16 Results 44 Areas 12 Rov surveys 16 Habitat types 44 Tarablus/Batroun 14 Infaunal surveys 16 Coralligenous habitat 44 Jounieh 14 Oceanographic and rhodolith/maërl 45 St. George beds measurements 46 Beirut 19 Sandy bottoms 15 Data analyses 46 Sayniq 15 Collaborations 20 Sandy-muddy bottoms 20 Rocky bottoms 22 Canyon heads 22 Bathyal muds 24 Species 27 Fishes 29 Crustaceans 30 Echinoderms 31 Cnidarians 36 Sponges 38 Molluscs 40 Bryozoans 40 Brachiopods 42 Tunicates 42 Annelids 42 Foraminifera 42 Algae | Deep sea Lebanon OCEANA 47 Human 50 Discussion and 68 Annex 1 85 Annex 2 impacts conclusions 68 Table A1. List of 85 Methodology for 47 Marine litter 51 Main expedition species identified assesing relative 49 Fisheries findings 84 Table A2. List conservation interest of 49 Other observations 52 Key community of threatened types and their species identified survey areas ecological importanc 84 Figure A1.
  • Phylum Nemertea)

    Phylum Nemertea)

    THE BIOLOGY AND SYSTEMATICS OF A NEW SPECIES OF RIBBON WORM, GENUS TUBULANUS (PHYLUM NEMERTEA) By Rebecca Kirk Ritger Submitted to the Faculty of the College of Arts and Sciences of American University in Partial Fulfillment of the Requirements for the Degree of Master of Science In Biology Chair: Dr. Qiristopher'Tudge m Dr.David C r. Jon L. Norenburg Dean of the College of Arts and Sciences JuK4£ __________ Date 2004 American University Washington, D.C. 20016 AMERICAN UNIVERSITY LIBRARY 1 1 0 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number: 1421360 INFORMATION TO USERS The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleed-through, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. ® UMI UMI Microform 1421360 Copyright 2004 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE BIOLOGY AND SYSTEMATICS OF A NEW SPECIES OF RIBBON WORM, GENUS TUBULANUS (PHYLUM NEMERTEA) By Rebecca Kirk Ritger ABSTRACT Most nemerteans are studied from poorly preserved museum specimens.

  • Annelida, Amphinomidae) in the Mediterranean Sea with an Updated Revision of the Alien Mediterranean Amphinomids

    A peer-reviewed open-access journal ZooKeys 337: 19–33 (2013)On the occurrence of the firewormEurythoe complanata complex... 19 doi: 10.3897/zookeys.337.5811 RESEARCH ARTICLE www.zookeys.org Launched to accelerate biodiversity research On the occurrence of the fireworm Eurythoe complanata complex (Annelida, Amphinomidae) in the Mediterranean Sea with an updated revision of the alien Mediterranean amphinomids Andrés Arias1, Rômulo Barroso2,3, Nuria Anadón1, Paulo C. Paiva4 1 Departamento de Biología de Organismos y Sistemas (Zoología), Universidad de Oviedo, Oviedo 33071, Spain 2 Pontifícia Universidade Católica do Rio de Janeiro , Rio de Janeiro, Brazil 3 Museu de Zoologia da Unicamp, Campinas, SP, Brazil 4 Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro (UFRJ) , Rio de Janeiro, RJ, Brasil Corresponding author: Andrés Arias ([email protected]) Academic editor: C. Glasby | Received 17 June 2013 | Accepted 19 September 2013 | Published 30 September 2013 Citation: Arias A, Barroso R, Anadón N, Paiva PC (2013) On the occurrence of the fireworm Eurythoe complanata complex (Annelida, Amphinomidae) in the Mediterranean Sea with an updated revision of the alien Mediterranean amphinomids. ZooKeys 337: 19–33. doi: 10.3897/zookeys.337.5811 Abstract The presence of two species within the Eurythoe complanata complex in the Mediterranean Sea is reported, as well as their geographical distributions. One species, Eurythoe laevisetis, occurs in the eastern and cen- tral Mediterranean, likely constituting the first historical introduction to the Mediterranean Sea and the other, Eurythoe complanata, in both eastern and Levantine basins. Brief notes on their taxonomy are also provided and their potential pathways for introduction to the Mediterranean are discussed.
  • In the Atlantic-Mediterranean Biogeographic Area

    In the Atlantic-Mediterranean Biogeographic Area

    SPECIES OF SPIOCHAETOPTERUS (POLYCHAETA, CHAETOPTERIDAE) IN THE ATLANTIC-MEDITERRANEAN BIOGEOGRAPHIC AREA MICHEL R. BHAUD BHAUD, MICHEL R. 1998 08 28. Species of Spiochaetopterus (Polychaeta, Chaetopteridae) in the Atlantic- SARSIA Mediterranean biogeographic area. – Sarsia 83:243-263. Bergen. ISSN 0036-4827. Five species of the genus Spiochaetopterus: S. typicus SARS, S. bergensis GITAY, S. costarum (CLAPARÈDE), S. oculatus WEBSTER, and S. solitarius (RIOJA) have been compared. These species can be divided into two groups: group A, with boreal biogeographic affinity, consisting of S. typicus and S. bergensis; group B, with temperate biogeographic affinity, consisting of S. costarum, S. solitarius and S. oculatus. The two groups differ in the shape of the specialized setae of segment A4, the number of segments in region B, and the structure of the tube. Representatives of group A have a middle region of two seg- ments, modified setae of segment A4 are distally rhomboid and the tubes are without articulations. Representatives of group B have a middle region of many, always more than 2 segments; modified setae of A4 are distally cordate, and the tubes are with articulations. In each geographic unit of the Atlantic-Mediterranean area, the species are divided into two groups of different sizes. The usefulness of such systematic characters as the number of segments in region B, ornamentation of the tubes and relative size of the pro- and peristomium is reexamined. New characters include the detailed morphol- ogy of the A4 setae, location of the secretory part of the ventral shield, and the structure of the secretory pores. Two diagnostic keys to species, one based on the tubes and the other on the A4 setae, are included.