Botanica Marina Vol. 46,2003, pp. 215-242 © 2003 by Walter de Gruyter • Berlin ■ New York

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Annual subscription rate (Volume 46,2003) Checklist of Mediterranean Free-living Dinoflagellates Institutional rate: € 938,-/approx. SFr1 50 1 in the US and Canada US $ 938,-. Individual rate: € 118,-/approx. SFr 189,-; in the US and Canada US $ 118,-. Personal rates apply only when copies are sent to F. Gómez a private address and payment is made by a personal check or credit card. Personal subscriptions must not be donated to a library. Single issues: € 178,-/approx. SFr 285,-. All prices exclude postage. Department of Aquatic Biosciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo, Tokyo 113-8657, Japan, [email protected] Orders Institutional subscription orders should be addressed to the publishers orto your usual subscription agent. Individual subscrip­ tion orders must be sent directly to one of the addresses indicated below.

The Americas: An annotated checklist of the free-living dinoflagellates (Dinophyceae) of the Mediterranean Sea, based on Walter de Gruyter, Inc., 200 Saw Mill River Road, Hawthorne, N.Y. 10532, USA, Tel. 914-747-0110, Fax 914-747-1326, literature records, is given. The distribution of 673 species in 9 Mediterranean sub-basins is reported. The e-mail: [email protected]. number of taxa among the sub-basins was as follows: Ligurian (496 species), Balear-Provençal (360), Adri­ Australia and New Zealand: atic (322), Tyrrhenian (284), Ionian (283), Levantine (268), Aegean (182), Alborán (179) and Algerian Seas D. A. Information Services, 648 Whitehorse Road, P.O. Box 163, Mitcham, Vic. 3132, Australia, Tel. +61 3 9210 7777, (151). Fax+61 3 9210 7788

Japan: Introduction cionellum Defrandre, Pentapharsodinium Indelicato Maruzen Company Ltd., P.O. Box 5050, Tokyo International, 100-31 Japan, Tel. +3-3275-8585, Fax +3-3274-3238 et Loeblich III and Preperidinium Mangin have been Igaku-ShoinLtd., P.O. Box 5063, Tokyo International, 113-91 Japan, Tel. +3-817-5675, Fax+3-815-6776 The oligotrophic conditions in the Mediterranean added to the Peridiniaceae. Kinokuniya Company Ltd., 38-1, Sakuragoako 5-chome, Setagaya-ku, Tokyo, 156 Japan, Tel. +3-439-0161, Sea could favour the richness of dinoflagellates, Fax +3-3439-0839 Synonyms have been tracked down and relocated typical organisms of oligotrophic waters. Intensive NankodoCo. Ltd. 42-6, Hongo 3-chome, Bunkyo-ku, Tokyo, 113 Japan, Tel. +3-811-9950, Fax+3-811-5031 in order to avoid duplicate entries. Synonyms, which studies have been made by Jörgensen (1920, have not been quoted in the world literature during Europe and Rest o f World: 1923), Schiller (1931-37) (Adriatic Sea), Pavillard the last decades, are not reported. Because of space SFG - Servicecenter-Fachverlage GmbH, Postfach 43 43, 72774 Reutlingen, Tel.: +49-(0)7071 -9353-30, (1905-1937) (Gulf of Lions and Monaco), Halim Fax: +49-(0)7071-93 56-33, e-mail: [email protected] limitation, not all the references reporting each (I960) (Villefranche and Alexandria), Rampi species for each area have been included. Only when (1939-1969) (Ligurian Sea) and Margalef (1945- a taxon is reported in less than 3 of the 9 Mediter­ 1995) (Spanish coasts). However a catalogue of the ranean sub-basins considered, is the source of the dinoflagellata species recorded is not available. The record reported. Exceptionally, also in 3 of the aims of this study are to provide a checklist of Mediterranean sub-basins when the number of cita­ the species from each sub-basin and to evaluate tions was low (< 5). In some cases, these scarcely re­ Attention before copying! the species richness of dinoflagellates in the Medi­ ported taxa can be considered as misidentifications terranean Sea based on a compilation of published Do you photocopy articles from this periodical? If so, have you made certain that you are not violating the legal copyright or unreliable records, recently described species or regulations and making yourself liable to prosecution? data. rare species. The results of this study depend on the According to copyright law, it is permissible to make a few copies of individual articles for personal use only. Reproduction of valid identification by the authors of each reference. articles for commercial use by an industrial enterprise is subject to charge. In m ost of th e cases, there are not photographs or fig­ Copying in the U.S.A. Material and Methods ures of the taxa and the verification of the records is Authorization to photocopy items for internal or personal use, or the internal or personal use of specific clients, is granted by This study is based on literature records of free-living difficult. Records of unarmoured cells should be con­ Walter de Gruyter GmbH & Co. KG • Berlin • New York for libraries and other users registered with the Copyright Clearance dinoflagellates (Table I), grouped in the main sub­ sidered cautiously due to the difficulties of their Center (CCC) Transactional Reporting Service, provided that the base fee of $ 20.00 per copy is paid directly to CCC 222 Rosewood Drive, Danvers, MA 01923, USA. basins of the M editerranean Sea (Fig. 1). References identification. Most of these doubtful records are in 0006-8055/03 lSSN Q006-8055 used for the elaboration of this checklist, but not cited the studies by Skolka et al. (1986) for the Libyan wa­ Copyright © 2003 by Verlag Walter de Gruyter GmbH & Co. KG Online version: ISSN 1437-4323 in the text, checklist or notes are listed in the Appen­ ters and/or Innamorati et al. (1986,1989 a,b) for the Printed in Germany dix. Species with their nomenclatural authorities are Ligurian Sea. Many species of the rare genera H is­ arranged alphabetically in each order according to tioneis and Heterodinium, mainly reported by Rampi Typesetting jürgen ullrich typosatz, Nördlingen the classification proposed by Chrétiennot-Dinet (1939-1969) and Halim (1960), were not further Printing and Binding Werner Hildebrand, Berlin et al. (1993) with the following modifications: the gen­ recorded after their first description. For recently de­ scribed taxa, the geographical distribution is still un­ All rights reserved, including those of translations into foreign languages. No part of this journal may be reproduced in any form by era Parahistioneis and Phalacrom a have been added photoprint, microfilm or any other means nor transmitted nor translated into a machine language without written permission from to the Dinophysaceae; Balechina Loeblich et Loe- known beyond the type locality (e.g., some calcare­ the publisher. blich III, Plectodinium Biecheler and the recently ous dinoflagellates). Parasitic (exceptDissodinium The quotation of registered names, trade names, trade marks, etc. in this journal does not imply, even in the absence of a specific erected genera A ka sh iw o G. H ansen et Moestrup, pseudolunula Swift ex Elbrâchter et Drebes) and statement, that such names are exempt from laws and regulations protecting trade marks, etc. and therefore free for general use. Karenia G. Hansen et Moestrup and Karlodinium symbiotic species (i.e., Symbiodinium Freudenthal) Walter de Gruyter GmbH & Co. KG, Postfach 30 34 21,10728 Berlin, Germany J. Larsen have been added to the Gymnodiniaceae; have been excluded. Freshwater species have been Tel. (030) 260 05-0, Fax (030) 260 05-251 Proterythropsis K ofoid et Swezy in Kofoid has been excluded [e.g., Peridiniella catenata (Levander) Walter de Gruyter, Inc., 200 Saw Mill River Road, Hawthorne, N.Y. 10532 U S A added to the Warnowiaceae; Pavillardinium De-Toni Balech, P. danica (Paulsen) Okolodkov et D odge, Tel. (914) 747-0110, Fax (914) 747-1326 W has been added to the Oxytoxaceae; Exuviella etc]. Sometimes these species are reported from off­ DE Internet: http://www.deGruyter.com Cienkowski has been added to the Prorocentraceae; shore waters especially in sub-basins such as the G M ysticella Carbonell-Moore has been added to the Adriatic or Aegean Seas. Taxa only reported from Walter de Gruyter • Berlin • New York Podolampadaceae; Calcigonellum Deflandre, Cal- the identification of cysts have been excluded except 216 F, Gómez Checklist of Mediterranean free-living dinoflagellates 217

Table I. References considered for each Mediterranean sub-basin (references from the Appendix are excluded). Table II. List of taxa and their distribution.

Alb Arg Bai Tyr Lig Ion Adr Aeg Lev Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no.

13 52 4 3 12 21 3 59 1 - 2 Actiniscales Sournia 1984 33 56 8-9 6 14-18 149 10 73-74 40 Actiniscaceae Kützing 1844 56 87-88 24 19 49 151 20 82-83 46 A ch radina Lohmann 1903 100 127 34 23 51 22 126 65-66 Achradina pulchra Lohmann + + 76,116 119 47 27-30 57-58 50 81 A ctiniscus Ehrenberg 1843 60 32 61-64 71-72 83 Actiniscus pentasterias (Ehrenberg) Ehrenberg + + + + 68-69 38-39 75-77 141 85-86 BrachydinialesLoeblich III ex Sournia 1984 96-99 54-55 84 150 106-107 Brachydiniaceae Sournia 1972 101-105 58 89-93 163-167 126 Asterodinium Sournia 1972 115-116 70 116 173-176 A stero d in iu m gracile Sournia1 + + +1,57 118 95 125 Asterodinium libanum Abboud-Abi Saab 1 + + 2,57,58 1 2 0 - 1 2 2 108-113 129-140 Brachydinium F.J.R. Taylor2 168-169 117 160-161 Brachydinium capitatum F.J.R. Taylor + 171-172 128 Brachydinium taylorii Sournia + 102 143-144 DesmomonadalesPascher 1914 146-148 Desmocapsaceae Pascher 1914 151 D esm ocapsa Pascher 1914 162 Desmocapsa gelatinosa Pascher3 + + 75,76,77,145 177-178 Haplodiniaceae Lindemann 1928 Haplodinium Klebs 1912 Haplodinium antjoliense Klebs 4 + 75 Dinococcales Pascher 1914 Gloeodiniaceae Pascher ex Schiller 1937 Gloeodinium Klebs 1912 Gloeodinium marinum Bouquaheux5 + + 12,103,160 Thoracosphaeraceae Schiller 1930 Thoracosphaera Kamptner 1927 Balea r- Thoracosphaera heimii (Lohmann) Kamptner6 + + + + + + Provengal DinophysalesLindemann 1928 Tyrrhenian Citharistaceae Kofoid et Skogsberg 1928 A lg e r ia n Citharistes Stein 1883 + 81 A lb o rán Citharistes apsteini Schütt Io n ian Citharistes regius Stein + + + Dinophysaceae Stein '1883 L evantine Amphisolenia Stein 1883 Amphisolenia bidentata Schröder + + + + + + + + + Amphisolenia bispinosa Kofoid + 29 Amphisolenia brevicauda Kofoid + 91,139 Amphisolenia clavipes Kofoid + 1 ,8 6 + 91 Fig. 1. Map of the Mediterranean sub-basins. Amphisolenia complanata Kofoid et Skogsberg Amphisolenia extensa Kofoid + + + 33,80,90 Amphisolenia globifera Stein + + + + + + + + + Amphisolenia inflata Murray et Whitting + + 91,105 Amphisolenia lemmermanni Kofoid + 40,46 when live cells have germinated from cysts (Ci- Acknowledgements Amphisolenia palaeotheroides Kofoid + 91 miniello et al. 2000, D ’Onofrio et al. 1999, Meier et al. Amphisolenia palmata Stein + + + + + I acknowledge the financial support by the Spanish 1 ,8 6 2002). Amphisolenia quadrispina Kofoid + 148,168 Ministry of Science and Technology and by the Euro­ Amphisolenia rectangulata Kofoid + + + 66 pean Commission (ICB2-CT-2001-80002). I thank Amphisolenia sigma Halim7 + + + + Results for the helpful comments and suggestions by four re­ Amphisolenia spinulosa Kofoid + + + viewers and the Editor. This checklist has been made Amphisolenia truncata Kofoid et Michener + + + + + Mediterranean free-living planktonic dinoflagellates possible with the collaboration of many colleagues Dinophysis Ehrenberg 1839 (= Phalacroma were represented by 673 species with 604 and 480 supplying less accessible literature. Stein 1883 partim.) + + + + + + species reported in the western and eastern basins Dinophysis acuminata Claparède et Lachmann8 + + + + + + + + Dinophysis acuta Ehrenberg 9 + + + respectively (Table II). The Ligurian (74 %), Balear- Accepted 22 December 2002. + + + Dinophysis alata Jörgensen 10 + + Provençal (53 %), Adriatic (48 %), Tyrrhenian = Ion­ + + + Dinophysis amandula (Balech) Sournia11 + + + ian (42%) and Levantine (40%) Seas showed the 125 Dinophysis apicata (Kofoid et Skogsberg) highest number of species whereas the Aegean Abé vel Balech (27%), Alborán (26%) and Algerian (22%) Seas 91 Dinophysis apiculata Meunier12 138,145 showed the lowest num ber of species. Dinophysis biceps Schiller 218 F. Gómez Checklist of Mediterranean free-living dinoflagellates 219

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference n Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no.

Dinophysis caudata Saville-Kent + + + + + + + + 4- Ornithocercus francescae (Murray et Whitting) + + + Dinophysis circumsuta (Karsten) Balech + + + + + + Balech26 Dinophysis dentata Schiller + 4- 77,145,175 Ornithocercus geniculatus Dangeard + + + Dinophysis diegensis Kofoid13 + + + 76,168,175 Ornithocercus heteroporus Kofoid + + + + + + + Dinophysis exigua Kofoid et Skogsberg + 81 Ornithocercus magnificus Stein + + + + + + + + + Dinophysis fortii Pavillard14 + + + + + + + + Ornithocercus quadratus Schütt27 + + + + + + + Dinophysis hastata Stein15 + + + + + + + + Ornithocercus splendidus Schütt28 + -t- + + Dinophysis irregularis (Lebour) Balech + 175 Ornithocercus steinii Schüttem end. Kofoid et + + + Dinophysis minuta (Cleve) Balech 150 Skogsberg 29 Dinophysis mitra (Schütt) Abé vel Balech16 + + + + + + + 4* Ornithocercus thumii (Schmidt) Kofoid et Skogsberg + + 33,115 Dinophysis monacantha Kofoid et Skogsberg + 95 Parahistioneis Kofoid et Skogsberg 1928 Dinophysis ovum Schütt17 + + + + + + + (= H istio n e is Stein 1883 pa rtim .) Dinophysis parva Schiller18 h- + + + Parahistioneis acutiformis Rampi + 136 Dinophysis punctata Jörgensen + + + + Parahistioneis karstenii (Kofoid et Michener) + 129 Dinophysis pusilla Jörgensen + + 76,115 Kofoid et Skogsberg 30 Dinophysis recurva Kofoid et Skogsberg 19 + + + + + + -h Parahistioneis mediterranea Schiller + + + Dinophysis rete Sournia20 + 173,175 Parahistioneis paraformis Kofoid et Skogsberg + + 81,136 Dinophysis rotundatum Claparède et Lachmann + + + + + 4- + + + Parahistioneis sphaeroidea Rampi + + 73,136 Dinophysis sacculus Stein21 + + + + + + + + + Parahistioneis varians Böhm in Schiller + 10 Dinophysis schilleri Sournia22 + + + + + Phalacroma Stein 1883 (= Dinophysis Ehrenberg Dinophysis schroederi Pavillard + 4- + + + + + 1839 p a rtim .) Dinophysis schuettii Murray et Whitting23 + + + + + + + 4~ Phalacroma acutum (Schütt) Pavillard31 + + + + + Dinophysis similis Kofoid et Skogsberg 24 + + 81,122,168 Phalacroma argus Stein + + + + + + + + Dinophysis sphaerica Stein + + + + + + + + 4- Phalacroma bipartitum Kofoid et Skogsberg 99 Dinophysis spinosa Rampi + + + + + Phalacroma cuneus Schütt + + + + + + + Dinophysis tripos Gourret + + + + + + + 4- + Phalacroma doryphorum Stein + + + + + + + + Dinophysis uracantha Stein 4" + + + +■ + Phalacroma expulsum (Kofoid et Michener) Kofoid + + 64,69,99 Histioneis Stein 1883 (= Parahistioneis Kofoid et et Skogsberg 32 Skogsberg 1928 partim.) Phalacroma favus Kofoid et Michener + + + + + + Histioneis alata Rampi + 136 Phalacroma nasutum Stein33 + + + + + + Histioneis bernhardii Rampi + 140 Phalacroma operculatum Stein + + + + Histioneis cerasus Böhm + 10 Phalacroma ovatum (Claparède et Lachmann) + + + + + + + Histioneis depressa Schiller + + + + Jörgensen Histioneis detonii Rampi25 + 136 Phalacroma parvulum (Schütt) Jörgensen + + + + + + + + + Histioneis elegans Halim + 64 Phalacroma porodictyum Stein + + + + + Histioneis expansa Rampi + 136 Phalacroma praetextum Kofoid et Michener + 95 Histioneis faouzii Halim + 64,140 Phalacroma pulchellum Lebour + + + + + + + Histioneis fragilis Böhm in Schiller + 149 Phalacroma striatum Kofoid + + + 80,125,173, Histioneis gubernans Schütt + + 130,140,176 175 Histioneis hippoperoides Kofoid et Michener + 81 Triposolenia Kofoid 1906 Histioneis hyalina Kofoid et Michener + + 81,149 Triposolenia bicornis Kofoid + + + + + + Histioneis imbricata Halim + 64 Triposolenia longicornis Kofoid + 76 Histioneis inclinata Kofoid et Michener 4- + + 47,136,149 Triposolenia truncata Kofoid + + + + + + Histioneis isselii Forti + + 51,141 Oxyphysaceae Sournia 1984 Histioneis joergensenii Schiller + + + + + Oxyphysis Kofoid 1926 Histioneis kofoidii Forti et Issel + + 50,95,141 Oxyphysis oxytoxoides Kofoid + + + + Histioneis ligustica Rampi + 133,136 GymnodinialesLemmermanii 1910 Histioneis longicollis Kofoid 4- + + 4- Gymnodiniaceae Lankester 1885 Histioneis marchesonii Rampi 4- + 34,133 Akashiwo G. Hansen et Moestrup 2000 Histioneis oxypteris Schiller + + 140,145 Akashiwo sanguinea (Hirasaka) G. Hansen et + + + + -f + + + Histioneis pavillardii Rampi + 129 Moestrup34 Histioneis rampii Halim + 64 Amphidinium Claparède et Lachmann 1885 Histioneis remora Stein 4- 41 81,122 Amphidinium acutissimum Schiller + + + + + Histioneis robusta Rampi + 140 Amphidinium acutum Lohmann + + + 1,168 Histioneis speciosa Rampi + 140 Amphidinium carterae Hulburt + + Histioneis subcarinata Rampi 4- + 99,136 Amphidinium conus Schiller + + 77,145 Histioneis sublongicollis Halim + + + 64 Amphidinium crassum Lohmann35 + + F + 149 Histioneis variabilis Schiller + + + Amphidinium cucurbitella Kofoid et Swezy + Histioneis villafranca Halim + 64 Amphidinium curvatum Schiller + + + Histioneis vouckii Schiller 75,76,149 + + + Amphidinium extensum Wulff + + Amphidinium flagellans Schiller + + + 220 F. Gómez Checklist of Mediterranean free-living dinoflagellates 221

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion A dr Aeg Lev Reference no. Dinophyceae West et Fritsch 1927 Bai Tyr Lig Ion Adr Aeg Lev Reference no.

Amphidinium glaucum Conrad + 76 Gymnodinium cucumis Schütt + + + + Amphidinium globosum Schröder + + + + + Gymnodinium diploconus Schütt + + + Amphidinium hyalinum Entz + + 77,149 Gymnodinium dissimile Kofoid et Swezy + + 76,149 Amphidinium inflatum Kofoid + 127 Gymnodinium elongatum Hope + + 144,149 Amphidinium kesslitzi Schiller + H- + Gymnodinium flavum Kofoid ei Swezy + + 76,81 Amphidinium lacustriforme Schiller36 + ■i- + Gymnodinium fulvum Kofoid et Swezy + 149 Amphidinium lanceolatum Schröder + + + Gymnodinium fuscum (Ehrenberg) Stein + + Amphidinium latum Lebour + 4- + Gymnodinium galeaeforme Matzenauer + 81 Amphidinium lissae Schiller + + 76,77,175 Gymnodinium gelbum Kofoid + 81 Amphidinium oceanicum Lohmann + 4- 75,149 Gymnodinium. gibberum Schiller + + + + Amphidinium operculatum Claparède et Lachmanii37 + + + 145,177 Gymnodinium gleba Schütt + + 76,147 Amphidinium ovoideum (Lemmermann) + 76 G ym n o d in iu m gracile Bergh + 149 Lemmermann Gymnodinium grammaticum (Pouchet) Kofoid et + + + + Amphidinium pellucidum Herdman + 76 Swezy48 Amphidinium roseolum (Schmarda) Schiller + 149 Gymnodinium heterostriatum Kofoid et Swezy40 + + + + Amphidinium schroederi Schiller38 + + 4- + Gymnodinium impudicum (Fraga et Bravo) + + + + + Amphidinium sphenoides Wulff39 + F 76,149 G. Hansen et Moestrup46 Amphidinium stigmatum Schiller 4- 4- + Gymnodinium incertum Herdman + 76 Amphidinium turbo Kofoid et Swezy + + 77,81 Gymnodinium incisum Kofoid et Swezy + 76 - Amphidinium vasculum Kofoid et Swezy 4- 149 Gymnodinium lachmannii Saville-Kent H" 75 Amphidinium vigrense Woloszynska + 76 Gymnodinium lineatum Kofoid et Swezy + 149 Balechina Loeblich et Loeblich III 1966 Gymnodinium lira Kofoid et Swezy + 76 Balechina coerulea (Dogiel) F.J.R. Taylor + 4- 76,149 Gymnodinium lohmannii Paulsen + + 40,149 Balechina marianae F.J.R. Taylor40 4- 160 Gymnodinium maguelonnense Biecheler50 + + 9,149 Cochlodinium Schütt 1896 Gymnodinium marinum Saville-Kent + + + Cochlodinium achromaticum Lebour + + 76,102 Gymnodinium minor Lebour + + + 21,22,76 Cochlodinium adriaticum Schiller + + 77,145 Gymnodinium mitratum Schiller + 76 Cochlodinium brandtii Wulff + + + + + Gymnodinium multilineatum Kofoid et Swezy + 76 Cochlodinium citron Kofoid et Swezy 4- 149 Gymnodinium multistriatum Kofoid et Swezy + + 81,168 Cochlodinium constrictum (Schütt) Lemmermann + + 81,147 Gymnodinium najadeum Schiller + + + H- Cochlodinium faurei Kofoid et Swezy + 81 Gymnodinium nanum Schiller + + 75,76,77,102 Cochlodinium geminatum (Schütt) Schütt + 147 Gymnodinium neapolitanum Schiller + + + + + Cochlodinium helix (Pouchet) Lemmermann41 + 4~ H- Gymnodinium opressum Conrad + + 75,76,102 Cochlodinium polykrikoides Margalef42 + 143 Gymnodinium ostenfeldii Schiller + + 75,76,145 Cochlodinium pulchellum Lebour + + + Gymnodinium ovulum Kofoid et Swezy + 75,76 Cochlodinium pupa Lebour + 1 0 1 ,1 0 2 Gymnodinium paulsenii Schiller + + + Cochlodinium strangulatum (Schütt) Schütt + + 4- Gymnodinium pulchellum J. Larsensl + + 23,171 Cochlodinium turbineum Kofoid et Swezy 4- 149 Gymnodinium pulchrum Schiller + Cochodinium schuettii Kofoid et Swezy + 141,145 Gymnodinium punctatum Pouchet + 75,76 Gymnodinium Stein 1878 emend. G. Hansen et Gymnodinium, pygmaeum Lebour + 76 Moestrup Gymnodinium ravenescens Kofoid et Swezy + 76 Gymnodinium achromaticum Lebour -f 34,99 Gymnodinium rotundatum Klebs + + + Gymnodinium agile Kofoid et Swezy + 4- 76,149 Gymnodinium rubrocinctum Lebour + 76 .. Gymnodinium agiliforme Schiller 4- + 4- + Gymnodinium scopulosum Kofoid et Swezy + 76 Gymnodinium albulum Lindemann43 + 76 Gymnodinium semidivisum Schiller + + 75,76,77,145 Gymnodinium amphora Kofoid et Swezy 4- 76 Gymnodinium simplex (Lohmann) Kofoid et Swezy43 -t + + + + Gymnodinium arcticum Wulff + + + + Gymnodinium situla Kofoid et Swezy 4- 149 Gymnodinium attenuatum Kofoid et Swezy + + 76,149 Gymnodinium sphaericum Calkins + 149 Gymnodinium auratum Kofoid et Swezy + + 76,149 Gymnodinium sphaeroideum Kofoid + + 75,149 Gymnodinium aureolum (Hulburt) G. Hansen44 + + + + + Gymnodinium sulcatum Kofoid et Swezy + 76 75 Gymnodinium aureum Kofoid et Swezy + H- 101,149 Gymnodinium translucens Kofoid et Swezy + Gymnodinium baccatum Balech + 34 Gymnodinium tridentatum Schiller + 149 Gymnodinium biconicum Schiller + + + + Gymnodinium variabile Herdman + + + 75,76,77,102, 149 Gymnodinium canus Kofoid et Swezy + + 8 6 ,149 Gymnodinium caput Schiller + 4- + + 76,145,149 Gymnodinium vestificii Schütt52 + + 76,147 Gymnodinium carinatum Schilling + 127 Gymnodinium voukii Schiller + + Gymnodinium catenatum Graham45-46 + + 13,56 Gymnodinium wulffii Schiller + + 77,149 Gymnodinium cinctum Kofoid et Swezy + 76 Gyrodinium Kofoid et Swezy 1921 emend. Gymnodinium conicum Kofoid ei Swezy47 + + 22,149 G. Hansen et Moestrup (=Gymnodinium Gymnodinium corii Schiller 4- + -f + Stein 1878partim.) Gymnodinium costatum Kofoid et Swezy + 99 Gyrodinium acutum (Schütt) Kofoidet Swezy + + + 222 F. Gómez Checklist of Mediterranean free-living dinoflagellates 223

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference n< Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no.

Gyrodinium biconicum Kofoid et Swezy + 149 Greuetodinium cylindricum (Greuet) Loeblich III 67 + 62 Gyrodinium conicum Schiller + 145 Nematodinium Kofoid et Swezy 1921 (= Pouchetia Gyrodinium contortum (Schütt) Kofoid et Swezy53 + + + + + Schütt 1895) Gyrodinium corsicum Paulmier, Beríand, Billard et + + + Nematodinium armatum (Dogiel) Kofoid et Swezy68 + + 38,76,93 Nezan Nematodinium torpedo Kofoid et Swezy + + 81,102 Gyrodinium crassum (Pouchet) Kofoid et Swezy + + + Proterythropsis Kofoid et Swezy in Kofoid Gyrodinium cuneatum Kofoid et Swezy + 75 Proterythropsis crassicaudata Kofoid et Swezy + 101,102 Gyrodinium fissum (Levander) Kofoid et Swezy54 + 127 Warnowia Lindemann 1928 (= Pouchetia Schütt) Gyrodinium fusiforme Kofoid et Swezy + + + + + + + Warnowia compacta (Schütt) Schiller + 147 Gyrodinium glaebum Hulburt + + + Warnowia dohrnii Zimmermann + + 101,177 Gyrodinium herbaceum Kofoid et Swezy. + + + Warnowia fusus (Schütt) Lindemann + + + + + Gyrodinium lachryma (Meunier) Kofoid el Swezy55 + + + + Warnowia juno (Schütt) Schiller + + 101,147 Gyrodinium longum (Lohmann) Kofoid et Swezy + + 75,101 Warnowia maculata (Kofoid et Swezy) Lindemann + 101 Gyrodinium nasutum (Wulff) Schiller + 149 Warnowia polyphemus (Pouchet) Schiller + + 120,149 Gyrodinium ochraceum Kofoid et Swezy + + 77,145 Warnowia pulchra Schiller + + + 61,145 Gyrodinium ovatum (Gourret) Kofoid et Swezy + + 60,81 Warnowia rosea (Pouchet) Schiller + + 49,121 Gyrodinium ovum (Schütt) Kofoidet Swezy + + 144,145,147, Warnowia violescens Kofoid et Swezy + 81 166 NoctilucalesHaeckel '1894 Gyrodinium parvulum (Schütt) Kofoidet Swezy + + + Kofoidiniaceae Taylor 1976 Gyrodinium pellucidum (Wulff) Schiller + + + Cymbodinium Cachón et Cachón 1967 Gyrodinium pepo (Schütt) Kofoid et Swezy56 + + + Cymbodinium elegans Cachón et Cachón + 16 Gyrodinium pingue (Schütt) Kofoidet Swezy57 + + + + Kofoidinium Pavillard 1928 Gyrodinium rubricaudatum Kofoid et Swezy + 76 Kofoidinium pavillardii Cachón et Cachón® + 17 Gyrodinium spirale (Bergh) Kofoid et Swezy + + + + -f- -H Kofoidinium splendens Cachón et Cachón + + 1,17 Gyrodinium varians (Wulff) Schiller + + Kofoidinium velelloides Pavillard + + + + + + + Karenia G. Hansen eí Moestrup 2000 Pomatodinium Cachón et Cachón 1966 Karenia brevis (Davis) G. Hansen et Moestrup58 +• + 59,81 Pomatodinium impatiens Cachón et Cachón + + 15,99 Karlodinium J. Larsen 2000 Spatulodinium Cachón et Cachón 1968 Karlodinium micrum (Leadbeater et Dodge) + 1 Spatulodinium pseudonoctiluca (Pouchet) Cachón et + + 18,173,175 J. Larsen59 Cachón ex Loeblich et Loeblich III Katodinium Fott 1857 (= Massartia Conrad 1926) Leptodiscaceae Kofoid 1916 Katodinium glaucum (Lebour) Loeblich III 60 + + + Cachonodinium Loeblich III 1980 Katodinium tubulatum (Rampi) Sournia61 ■t- 140 Cachonodinium caudatum (Cachón et Cachón) + 18 Plectodinium Biechelerl93462 Loeblich III 70 Plectodinium nucleovolvatum Biecheler + + + + Craspedotella Kofoid 1905 Pseliodinium Sournia 1972 Craspedotella pileolus Kofoid + i7 Pseliodinium vaubanii Sournia + + + + + + Leptodiscus Hertwig 1877 (= Pratjetella Torodinium Kofoid et Swezy 1921 Lohmann 1920) Torodinium robustum Kofoid et Swezy + + -t ■i- + + Leptodiscus medusoides Hertwig71 + + + 18> 70,99,171 Torodinium teredo (Pouchet) Kofoid et Swezy63 + + + + 2,118,145,14i Leptophyllus Cachón et Cachón 1964 (= Abedinium Polykrikaceae Kofoid et Swezy 1921 Loeblich et Loeblich III 1966) Pheopolykrikos Chatton emend. Matsuoka et Leptophyllus dasypus Cachón et Cachón 72 + 14 Fukuyo 1986 Petalodinium Cachón et Cachón 1969 Pheopolykrikos beauchampii Chatton + 24 Petalodinium porcelio Cachón et Cachón + 18 Pheopolykrikos hartmannii (Zimmermann) + 177 Scaphodinium Margalef (= Leptospathium Matsuoka et Fukuyo Cachón et Cachón 1964) Polykrikos Btitschli 1873 Scaphodinium mirabile Margalef73 + + + + Polykrikos kofoidii Chatton + + + + + Noctilucaceae Kent 1881 Polykrikos schwartzii Bütschli + + + + Noctiluca Suriray ex Lamarck 1816 Ptychodiscaceae Lemmermann 1899 Noctiluca scintillans (Macartney) Kofoid + + + + + + + + + Ptychodiscus Stein 1883 Protodiniferaceae Kofoid et Swezy 1921 Ptychodiscus noctiluca Stein64 + + 123 137 Pronoctiluca Fabre-Domergue 1889 Warnowiaceae Lindemann 1928 Pronoctiluca pelagica Fabre-Domergue + + + Erythropsidinium P.C. Silva 1960 (= Erythropsis Pronoctiluca rostrata F.J.R. Taylor74 + Hertwig 1884) Pronoctiluca spinifera (Lohmann) Schiller + + + + Erythropsidinium agile (Hertwig) PC. Silva65 + + 33 147 OxyrrhinalesSournia 1984 Erythropsidinium minor (Kofoid et Swezy) P.C. Sil va + 101 102 Oxyrrhinaceae Sournia 1984 Erythropsidinium pavillardii (Kofoid et Swezy) + + 61 [on Oxyrrhis Dujardin 1841 P.C. Silva66 Oxyrrhis marina Dujardin + + + + + Greuetodinium Loeblich III 1980 (= Leucopsis PeridinialesHaeckel 1894 Greuet 1968) Amphidomataceae Sournia 1984 224 F. Gómez Checklist of Mediterranean free-living dinoflagellates 225

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference] Dinophyceae West ei Fritsch 1927 Alb Alg Bal Tyr Lig lon Adr Aeg Lev Reference no.

Amphidoma Stein 1883 (=Pavillardinium Ceratium teres Kofoid 4- 4- 4- 4- 4- 4- 4- -f + De-Toni 1936 partim., Murrayella Kofoid 1907) Ceratium trichoceros (Ehrenberg) Kofoid 4- 4- 4- 4- 4- 4- 4- + + Amphidoma caudata Halldal75 Ceratium tripos (Müller) Schiller 4- 4" 4- 4- 4" 4- 4- + + Amphidoma elongata Kofoid et Swezy 149 Ceratium volans Cleve87 + 4- 4- 4- 4- 4- 4- Amphidoma nucula Stein76 33,8 Ceratium vultur Cleve 4- 4- 4- 4- + + 4- Ceratiaceae Kofoid 1907 Ceratocorythaceae Lindemann 1928 Ceratium Schrank 1793 Ceratocorys Stein 1883 Ceratium arietinum Cleve + + 4- 4- 4- 4- 4- + 4- Ceratocorys armata (Schütt) Kofoid -f 4- 4- 4- 4- 4- + 4- Ceratium azoricum Cleve 4- + + 4- 4- 4- + Ceratocorys gourretii Paulsen 4- 4- 4- 4- + + 4- Ceratium belone Cleve 4- + + 4- 4- 4- + 4- Ceratocorys horrida Stein 4- 4- 4- 4- + + + 4- Ceratium breve (Ostenfeld et Schmidt) Schröder 4- 4- + 4- Cladopyxidaceae Poche 1913 Ceratium brunellii Rampi77 4 - 134 Cladopyxis Stein 1883 (= Micracanthodinium Ceratium buceros (Zacharias) Schiller 4- + 4- 4- 4 - 4- 4- 4- 4- Deflandre 1937 partim.) Ceratium candelabrum (Ehrenberg) Stein + + 4- 4- 4- 4- 4- 4" 4- Cladopyxis brachiolata Stein Ceratium carriense Gourret + + 4- 4- 4 - 4- 4- 4- 4- Cladopyxis caryophyllum (Kofoid) Pavillard88 Ceratium claviger Kofoid78 4- + 4- Cladopyxis quadrispina Pavillard89 Ceratium coarctatum Pavillard 4- 4- 4- 4- 4- 4- 4- Cladopyxis spinosa (Kofoid) Schiller90 64,168 Ceratium concilians Jörgensen 4- 4- 4- 4- 4- 4- 4- 4- Palaeophalacroma Schiller 1928 Ceratium contortum (Gourret) Cleve 4~ 4- 4- + 4 - 4* 4- 4- 4- Palaeophalacroma unicinctum Schiller91 Ceratium contrarium (Gourret) Pavillard79 + 4- 4- + 4 - 4- 4- 4- 4- Palaeophalacroma verrucosum Schiller Ceratium declinatum (Karsten) Jörgensen + 4- 4- + 4- 4- 4- 4- + Crypthecodiniaceae Biecheler ex Chatton 1952 Ceratium deflexum (Kofoid) Jörgensen + 4- 4- 4- Crypthecodinium Biecheler 1938 Ceratium denticulatum (Jörgensen) Paulsen80 + 4- 4- 82,119,168 Crypthecodinium cohnii (Seligo) Chatton92 Ceratium digitatum Schütt 4- + 4- 4- 4- 4- 4- Goniodomataceae Lindemann 1928 Ceratium egyptiacum Halim81 4- 1,65,86 Goniodoma Stein 1883 (= Triadinium Dodge 1981, Ceratium euarcuatum Jörgensen 82 4- + + + 4- 4- 4- 4- 4- = Heteraulacus Diesing 1850 partim.) Ceratium extensum (Gourret) Cleve83 4- 4- 4* + 4- 4- 4- 4- 4- Goniodoma acuminatum (Ehrenberg) Stein 93 Ceratium falcatiforme Jörgensen + 4- 4- 4- 4- Goniodoma sphaericum Murray et Whitting94 Ceratium falcatum (Kofoid) Jörgensen + 4- 4- 4- 4- 4- 4- 4- Pyrodinium Plate 1906 Ceratium furca (Ehrenberg) Claparède ei Lachmann + 4 - 4- 4- 4- 4- 4 - 4- 4- Pyrodinium bahamense Plate95 40 Ceratium fusus (Ehrenberg) Dujardin + 4 - 4- 4- 4- 4- 4- 4- 4- Gonyaulacaceae Lindemann 1928 Ceratium geniculatum (Lemmermann) Cleve + 4- 4- 4- Alexandrium Halim emend. Balech 1989 Ceratium gibberum Gourret + 4- 4- 4- 4- 4- 4- 4- (= Protogonyaulax F.J.R. Taylor 1976) Ceratium gravidum Gourret 4- 4- 4- 4- 4- 4- 4- 4- Alexandrium andersonii Balech 27 Ceratium hexacanthum Gourret 4~ 4- 4- 4- 4- 4- 4- 4- Alexandrium balechii (Steidinger) Balech 110,156 Ceratium horridum (Cleve) Gran84 + 4- 4- 4- 4- 4- 4- 4- Alexandrium catenella (Whedon et Kofoid) Balech Ceratium incisum (Karsten) Jörgensen 4- 4- 4- 4- 4- 4- 4- Alexandrium compressum (Fukuyo, Yoshida et 20 Ceratium inflatum (Kofoid) Jörgensen + 4- 4- 4- 4- + 4- 4- Inoue) Balech 30 Ceratium kofoidii Jörgensen 4- 4- 4- 4- 4- + 4- 4- Alexandrium concavum (Gaarder) Balech 6,30 Ceratium limulus (Gourret ex Pouchet) Gourret 4~ 4- 4- 4- 4- 4- + 4- 4- Alexandrium foedum Balech 30 Ceratium lineatum (Ehrenberg) Cleve 4- 4- 4- Alexandrium insuetum Balech 30 Ceratium longirostrum Gourret + 4- 4- 4- 4- 4- 4* 4- + Alexandrium kutnerae (Balech) Balech 30 Ceratium longissimum (Schröder) Kofoid 4- 4- 4- 4- 4- 4- 4- Alexandrium leei Balech 30 Ceratium lunula (Schimper ex Karsten) Jörgensen + 4- 4- 4- + 4- Alexandrium margalefii Balech Ceratium macroceros (Ehrenberg) Cleve + 4- + 4- 4- 4- 4- 4- 4- Alexandrium minutum Plalim96 87,107 Ceratium massiliense (Gourret) Karsten + 4- + 4- 4- 4- 4- 4- 4- Alexandrium ostenfeldii (Paulsen) Balech et Tangen9 20,30,71 Ceratium minutum Jörgensen + 4- 4- 4- Alexandrium pseudogonyaulax (Biecheler) Ceratium paradoxides Cleve85 + 4- 4- 4- Horiguchi ex Kita et Fukuyo Ceratium pavillardii Jörgensen + 4- + 4- 4- 4- 4- 4- 4- Alexandrium tamarense (Lebour) Balech Ceratium pentagonum Gourret 4- + 4" 4- 4- 4- 4- 4- Alexandrium taylorii Balech Ceratium platycorne Daday 4* + 4- 4- 4- + 4- 4- Amylax M eunier 1910 (= Gonyaulax Diesing Ceratium praeolongum (Lemmermann) + + + 4- 1866 partim. ) 1 Kofoid ex Jörgensen Amylax buxus (Balech) Dodge98 87 Ceratium pulchellum Schröder 4- 4- + + 4- 4- + 4- 4- Amylax triacantha (Jörgensen) Sournia99 Ceratium ranipes Cleve + + + 4- 4- + 4- 4- Gonyaulax Diesing 1866 (= A m ylax Meunier Ceratium reflexum Cleve + 150 1910 partim.) 4- 4- 4- 76,138,145 Ceratium schroeteri Schröder 4- 4- + 4- Gonyaulax africana Schiller Ceratium setaceum Jörgensen + 4- 4- 4- 4- 4" 4- + 4- + + 4- 4- + 4- Gonyaulax birostris Stein 4- 33,76,100,119 Ceratium strictum (Okamura et Nishikawa) Kofoid + 4- 4- + 4- 4- Gonyaulax brevisulcatum Dangeard Ceratium symmetricum Pavillard 4- 4- 4- 4- 4- + 4- 4- 4- 4- 4- + 4- 4- Gonyaulax diegensis Kofoid100 4- 4- 4- 4- + 4- 4- Ceratium tenue (Ostenfeld ei Schmidt) Jörgensen 86 4- 4- 4- 4- 4- Gonyaulax digitale (Pouchet) Kofoid101 226 F. Gómez Checklist of Mediterranean free-living dinoflagellates 227

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference r Dinophyceae West ei Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no.

Gonyaulax elegans Rampi 138 Heterodinium scrippsi Kofoid114 + + + 64,89,90,91, Gonyaulax fragilis (Schütt) Kofoid 124,175 Gonyaulax glyptorhynchus Murray et Whitting102 95 Heterodinium sinistrum Kofoid et Adamson115 + l Gonyaulax gracilis Schiller 52,167 Heterodinium whittingae Kofoid + 91,92,124, Gonyaulax highleyii Murray et Whitting103 114 125,139 Gonyaulax hyalina Ostenfeld et Schmidt + + '138,173, Ostreopsidaceae Lindemann 1928 175 Coolia M eunier 1919 (=Ostreopsis J. Schmidt Gonyaulax ligustica Rampi + + 118,138 1901 partim .) Gonyaulax kofoidii Pavillard104 + + + + 4- + + Coolia m onotis Meunier116 + + + + Gonyaulax minuta Kofoid et Michener105 -t + + + + Ostreopsis I Schmidt 1901 Gonyaulax monacantha Pavillard + + + + + + 4- 4- + Ostreopsis ovata Fukuyo + 162 Gonyaulax monospina Rampi 4- + 20,138,1 Ostreopsis siamensis J. Schmidt + + 151,161 Gonyaulax orientalis Lindemann + 76 Oxytoxaceae Lindemann 1928 Gonyaulax ovalis Schiller106 + + + + Centrodinium Kofoid 1907 (= Pavillardinium De-Tonii Gonyaulax pacifica K o fo id 104-107 + + + + + 4- + + 1936 partim . , Murrayella Kofoid 1907partim.) Gonyaulax polygramma Stein + + + + 4* 4- + + + Centrodinium biconicum (Murray et Whitting) + + 81,138 Gonyaulax rostrata Dangeard + 168 F.J.R. Taylor 117 Gonyaulax rotundata Rampi + 4- 22,138 Centrodinium complanatum (Cleve) Kofoid + + + Gonyaulax rugosum Wailes103 4- 114,149 Centrodinium elongatum Kofoid + 33 Gonyaulax scrippsae Kofoid + 4- + + Centrodinium eminens Böhm + + 91,138,175 Gonyaulax sphaeroidea Kofoid + '135 Centrodinium intermedium Pavillard + + 4- Gonyaulax spinifera (Claparède et Lachmann) + + + + + 4- 4- 4- + Centrodinium m axim um Pavillard + + + + Diesing108 Centrodinium pavillardii F.J.R. Taylor118 + + + + + + Gonyaulax trottii Rampi + 138 Centrodinium splendidum (Rampi) F.J.R. Taylor119 + 131,138 Gonyaulax turbynei Murray et Whitting + + + + + Corythodinium Loeblich et Loeblich III 1966 Gonyaulax unicornis Lebour + 64,76 (= Oxytoxum Stein 1883partim.) Gonyaulax verior Sournia109 + + + + + + + Corythodinium belgicae (Meunier) F.J.R. Taylor103 + + 76,81,114 Lingulodinium Wall emend. Dodge 1989 Corythodinium compressum (Kofoid) F.J.R. Taylor + + + Lingulodinium milneri (Murray et Whitting) Dodge1 86 Corythodinium constrictum (Stein) F.J.R. Taylor + + + + + + + + + Lingulodinium polyedra (Stein) Dodge111 Corythodinium cristatum (Kofoid) F.J.R. Taylor + + + Protoceratium Bergh 1881 (= Gonyaulax Corythodinium curvicaudatum (Kofoid) F.J.R. Taylor + 168 Diesing 1866 partim.) Corythodinium diploconus (Stein) F.J.R. Taylor + + + 64,88,175 Protoceratium areolatum Kofoid Corythodinium elegans (Pavillard) F.J.R. Taylor + + + + + Protoceratium pepo Kofoid ei Michener 138 Corythodinium frenguellii (Rampi) F.J.R. Taylor + + + 138,168,175 Protoceratium reticulatum (Claparède et Corythodinium reticulatum (Stein) Loeblich et + + + + + + + Lachmann) Bütschli112 Loeblich III Protoceratium spinulosum (Murray et Corythodinium tessellatum (Stein) Loeblich et + + + + + + + + + Whitting) Schiller Loeblich III Pleterodiniaceae Lindemann 1928 Oxytoxum Stein 1883 Heterodinium Kofoid 1906 Oxytoxum aceratum Rampi + 138 Heterodinium agassizii Kofoid 125 Oxytoxum adriaticum Schiller + + + + Heterodinium balechii Rampi 138,168 Oxytoxum areolatum Rampi + + + 22,68,131 Heterodinium crassipes Schiller 145 Oxytoxum brunellii Rampi120 + + + 64,74,118,131 Heterodinium debeauxii Rampi 131 Oxytoxum caudatum Schiller + + + + Heterodinium dispar Kofoid ei Adamson 34.99.138 Oxytoxum coronatum Schiller + + + 140,145,175 Heterodinium doma (Murray et Whitting) Kofoid 131 Oxytoxum crassum Schiller + + + + Heterodinium dubium Rampi 131 Oxytoxum cribosum Stein + 140 Heterodinium fides Kofoid 1 Oxytoxum curvatum (Kofoid) Kofoid121 + + H- + Heterodinium globosum Kofoid 124,125,131 Oxytoxum depressum Schiller + + + Heterodinium grahamii Rampi 64.138 Oxytoxum elongatum Wood + 81 Heterodinium inaequale Kofoid113 64,121 Oxytoxum gladiolus Stein + + + + + + Heterodinium kofoidii Pavillard 90,122,145 Oxytoxum globosum Schiller122 + + + Heterodinium laticeps Léger 90 Oxytoxum laticeps Schiller + + + + + + + + Heterodinium leiorhynchum (Murray et Whitting) Oxytoxum longiceps Schiller123 + + + -t + Kofoid Oxytoxum longum Schiller + + + + + Heterodinium mediterraneum Pavillard Oxytoxum milneri Murray et Whitting124 + + + + + + Heterodinium milneri (Murray et Whitting) Kofoid 64,124,129, Oxytoxum minutum Rampi + + + + -T 140 175 Oxytoxum obesum Rampi + Heterodinium minutum Kofoid et Michener 34,81 Oxytoxum obliquum Schiller + + + + + + + + Heterodinium murrayi Kofoid 49,64,131,139 Oxytoxum ovale Schiller125 + + Heterodinium rigdenae Kofoid 1 Oxytoxum pachyderme Schiller ex F.J.R. Taylor 4- 145 228 F. Gómez Checklist of Mediterranean free-living dinoflagellates 229

Table II. (continued) Table II. (continued)

Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Dinophyceae West et Fritsch 1927 Dinophyceae West et Fritsch 1927 Reference no. Alb Alg B a i Tyr Lig Ion Adr Aeg Lev Reference :

Oxytoxum parvum Schiller126 + + +' + + Protoperidinium conicoides (Paulsen) Balech + 40 Oxytoxum punctulatum Rampi127 + + 55,138 Protoperidinium conicum (Gran) Balech + + + + + + + + + Oxytoxum radiosum Rampi + 131,138 Protoperidinium crassipes (Kofoid) Balech147 + + + + + + + + Oxytoxum rampii Sournia128 + + 29,140 Protoperidinium curvipes (Ostenfeld) Balech148 + + + + + Oxytoxum scolopax Stein + + + + + + + + + Protoperidinium deficiens (Meunier) Balech + 86 Oxytoxum sphaeroideum Stein + + + + + + Protoperidinium depressum (Bailey) Balech + + + + + + + + + Oxytoxum spinosum Rampi + 64,131 Protoperidinium diabolus (Cleve) Balech149 + + + + + + + + + Oxytoxum subulatum Kofoid + + 29,91,105 Protoperidinium divergens (Ehrenberg) Balech + + + + + + + + Oxytoxum turbo Kofoid + + + Protoperidinium elegans (Cleve) Balech + + + Oxytoxum variabile Schiller129 + 4* + + + + + Protoperidinium excentricum (Paulsen) Balech + + + Oxytoxum viride Schiller + + + + + Protoperidinium exiguipes (Mangin ex Halim) Dodge + 40 Pavillardinium De-Toni 1936 (= Amphidoma Protoperidinium fimbriatum (Meunier) Balech + 76 Stein 1883 partim., Murrayella Kofoid 1907) Protoperidinium finitimum Balech150 + + + + + Pavillardinium ovale (Pavillard) De-Toni130 29,123 Protoperidinium globulus (Stein) Balech131 + + + + + + + + Schuettiella Balech 1988 (= Gonyaulax Diesing 1866 Protoperidinium grande (Kofoid) Balech + + + + t- + + partim., Oxytoxum Stein 1883 partim.) Protoperidinium granii (Ostenfeld) Balech + + + + + + + + H- Schuettiella mitra (Schütt) Balech131 Protoperidinium heteracanthum (Dangeard) Balech + + 77,99 Peridiniaceae Ehrenberg 1828 Protoperidinium hirobis (Abé) Balech + + 1,144 Calcigonellum Deflandre 1948 Protoperidinium inclinatum (Balech) Balech + 99 Calcigonellum infida Deflandre emend. Montresor132 39 Protoperidinium inflatum (Okamura) Balech + + + + + + Calciodinellum Deflandre 1947 Protoperidinium latispinum (Mangin) Balech H- + + + Calciodinellum levantinum Meier, Janofske et 106 Protoperidinium leonis (Pavillard) Balech + + + + + + + Willems133 Protoperidinium ligusticum (Rampi) Balech + 138 . Calciodinellum operosum Deflandre132 39 Protoperidinium maranense Tolomio + 163,165 Diplopelta Stein ex Jörgensen 1912 (= Dissodium Protoperidinium marielebourae (Paulsen) Balech + + + + + Abé 1941 partim.) Protoperidinium mediterraneum (Kofoid) Balech + + + + + + + + + Diplopelta bomba Stein ex Jörgensen 134 Protoperidinium minutum. (Kofoid) Loeblich III + + + + + Diplopelta symmetrica Pavillard135 Protoperidinium mite (Pavillard) Balech + + + + + Diplopsalis Bergh 1881 (= Dissodium Protoperidinium nipponicum (Abé) Balech 152 + + 40,150 Abé 1941 partim.) Protoperidinium nudum (Meunier) Balech153 + 98,99 Diplopsalis lenticula Bergh136 Protoperidinium oblongum (Aurivillius) + + + + + + Diplopsalopsis Meunier emend. Balech 1988 Parke et Dodge Diplopsalopsis orbicularis (Paulsen) Meunier137 150 Protoperidinium obtusum (Karsten) Parke et Dodge + 150 Diplopsalopsis latipeltata Balech et Borgese 28,144 Protoperidinium oceanicum (Vanhoffen) Balech + + + + + + •r + + Kryptoperidinium Lindemann 1924 Protoperidinium oviforme (Dangeard ) Balech + + + + (= Glenodinium Ehrenberg 1837 partim.) Protoperidinium ovum (Schiller) Balech + + + -t + + + Kryptoperidinium foliaceum (Stein) Lindemann138 Protoperidinium pallidum (Ostenfeld) Balech + + + + + + + Oblea Balech ex Loeblich et Loeblich III 1966 Protoperidinium parthenopes Zingone et Montresor + 178 Oblea rotunda (Balech) Balech ex Sournia139 32,149 Protoperidinium pedunculatum (Schlitt) Balech + + + + + + Pentapharsodinium Indelicate et Loeblich III 1986 Protoperidinium pellucidum (Schütt) Balech + + + + + + + + (= Peridinium Ehrenberg 1831 partim.) Protoperidinium pentagonum (Gran) Balech + + + + + + + + Pentapharsodinium tyrrhenicum (Balech) 6,111 Protoperidinium punctulatum (Paulsen) Balech + + + + + + + Montresor, Zingone et Marino140 Protoperidinium pyriforme (Paulsen) Balech + + + + + + + + + Peridinium Ehrenberg 1831141 Protoperidinium quarnerense (Schröder) Balech + + + + + + + Peridinium quinquecorne Abé 142 Protoperidinium schilleri (Paulsen) Balech + + + Preperidinium Mangin 1913 (= Diplopeltopsis Protoperidinium simulum (Paulsen) Balech + + + + Pavillard 1913, Zygabikodinium Loeblich et Protoperidinium sinaicum (Matzenauer) Balech + + 76,77,149 Loeblich III 1970) Protoperidinium solidicorne (Mangin) Balech154 + + + r* + Preperidinium meunieri (Pavillard) Elbrächter 143 Protoperidinium sphaericum (Murray et Whitting) + + + + + Protoperidinium Bergh emend. Balech 1974141 Balech (= Peridinium Ehrenberg 1831 partim., Protoperidinium sphaeroides (Dangeard) Balech155 + + + Minuscula Lebour 1925) Protoperidinium sphaeroideum (Mangin) Balech155 + 149 Protoperidinium abei (Paulsen) Balech144 + + + + + + Protoperidinium steinii (Jörgensen) Balech + + + + + + + + -f- Protoperidinium anthonyi (Fauré-Fremiet) Balech + 76 Protoperidinium subinerme (Paulsen) Loeblich III + + + + + + + Protoperidinium bipes (Paulsen) Balech145 + + + + + + + Protoperidinium thorianum (Paulsen) Balech + + + + 146 63,64 Protoperidinium bispinum (Schiller) Balech + + + + + Protoperidinium tregouboffii (Halim) Balech136 + Protoperidinium brevipes (Paulsen) Balech + + + Protoperidinium tristylum (Stein) Balech + + + 76,168,17! Protoperidinium brochii (Kofoid et Swezy) Balech + + + + + + -f- + + Protoperidinium tubum (Schiller) Balech + + + + Protoperidinium bulla (Meunier) Balech + 76 Protoperidinium tumidum (Okamura) Balech + + + Protoperidinium cerasus (Paulsen) Balech + + + + + + + Protoperidinium variegatum (Peters) Balech + 127 Protoperidinium claudicans (Paulsen) Balech + + + + + + + + + Protoperidinium wiesneri (Schiller) Balech157 -r 230 F. Gómez Checklist of Mediterranean free-living dinoflagellates 231

Table II. (continued) Table II. (continued)

Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no. Dinophyceae West et Fritsch 1927 Alb Alg Bai Tyr Lig Ion Adr Aeg Lev Reference no.

Scrippsiella Balech ex Loeblich III 1965158 Prorocentrum balticum (Lohmann) Loeblich III + + + + + + + Scrippsiella lachrymosa Lewis 39,112 Prorocentrum belizeanum Faust + + 151 Scrippsiella precaria Montresor et Zingone 109 Prorocentrum cassubicum (Woloszynska) Dodge + + 75,76,83 Scrippsiella ramonii Montresor 39,108 Prorocentrum compressum (Bailey) Abé ex Dodge173 + + + + + + + -t 4- Scrippsiella rotunda Lewis 39,117 Prorocentrum concavum Fukuyo + + 151 Scrippsiella spinifera Honsell et Cabrini 72 Prorocentrum cordatum (Ostenfeld) Dodge174-175 + + + + + + + Scrippsiella trochoidea (Stein) Balech ex Prorocentrum dactylus (Stein) Dodge + ■+ + + + Loeblich III 159 Prorocentrum dentatum Stein176 + + + + + + + + Podolampadaceae Lindemann 1928 Prorocentrum emarginatum Fukuyo + 172 Blepharocysta Ehrenberg 1873 Prorocentrum gracile Schritt177 + + + + + + + Blepharocysta hermosillai Carbonell-Moore 19 Prorocentrum lima (Ehrenberg) Dodge + + + + + + Blepharocysta paulsenii Schiller + + + + Prorocentrum maximum (Gourret) Schiller178 + + + + + Blepharocysta splendor-maris (Ehrenberg) Stein + + + + 4- Prorocentrum micans Ehrenberg 179 + + + + + + + + + Mysticella Carbonell-Moore 1994 Prorocentrum minimum (Pavillard) Schiller175 + + + + + + Mysticella striata (Schütt) Carbonell-Moore160 ■t 132 Prorocentrum nanum Schiller180 + + + Podolampas Stein 1883 Prorocentrum nux Puigserveret Zingone + 128 Podolampas bipes Stein + + + + ■t" + + 4- 4- Prorocentrum ovum (Schiller) Dogde + + + + + Podolampas curvatus Schiller + + 76,145 Prorocentrum rostratum Stein + + + + + Podolampas elegans Schütt + + + + + + + + 4- Prorocentrum rotundatum Schiller181 + + + + + + + Podolampas palmipes Stein + + + + + + 4- + 4- Prorocentrum scutellum Schröder 182 + + + + + + + + + Podolampas spinifera Okamura161 + + + + + 4“ 4- 4- 4- Prorocentrum triestinum Schiller + + + + + + + Pyrophacaceae Lindemann 1928 Prorocentrum vaginulum (Stein) Dodge183 + + + + + + Pyrophacus Stein 1883 Prorocentrum venetum Tolomio et Cavolo184 + 164 Pyrophacus horologium Stein emend. Wall et Dale + + + + + + 4- + 4- ProtaspidalesLoeblich III 1970 Pyrophacus steinii (Schiller) Wall et Dale + + + + 4- 4- Entomosigmataceae Chatton 1952 Pyrophacus vancampoae (Rossignol) Wall et Dale162 96,160 Entomosigma Schiller 1925 Peridiniales incertae sedis Entomosigma peridinioides Schiller185 + + + 75,76,145 Ceratoperidinium Margalef ex Loeblich III 1980 Pyrocystales Apstein 1909 Ceratoperidinium mediterraneum Abboud-Abi Saab 163 2 Pyrocystaceae (Schütt) Lemmermann 1899 Ceratoperidinium yeye Margalef ex Loeblich III 163 1,99,169 Dissodinium Klebs in Pascher emend. Fragilidium Balech ex Loeblich III 1965 Elbrächter et Drebes 1978186 (=Helgolandicum von Stosch 1869, Dissodinium pseudolunula Swift ex Elbrächter et Goniodoma Stein 1883partim.) Drebes 187 Fragilidium fissile Balech Pyrocystis Murray ex Flaeckel 1890186 Heterocapsa Stein 1883 (= Cachonina (= Gymnodinium Stein 1883 partim., Loeblich III 1968) Dissodinium Klebs in Pascher emend. Heterocapsa lanceolata Iwataki et Fukuyo 164 126 Elbrächter ei Drebes 1978 partim.) H- Heterocapsa niei (Loeblich III) Morrill et 4- + + Pyrocystis acuta Kofoid + 76,125 Loeblich III 165 Pyrocystis elegans Pavillard + H- + + + + + + + + + + + Heterocapsa rotundata (Lohmann) G. Hansen166 4- + + + Pyrocystis fusiformis (Wy ville-Thomson ex Haeckel) + + + Heterocapsa triquetra (Ehrenberg) Stein + 4- + + + Blackman 188 Micracanthodinium Deflandre 1937 Pyrocystis gerbaultii Pavillard189 + + + (= Cladopyxis Stein 1883partim.) Pyrocystis hamulus Cleve + + + + + Micracanthodinium bacilliferum (Schiller) + + 20,140 Pyrocystis margalefii Léger190 + + 91,104 Deflandre167 Pyrocystis minima (Matzenauer) Schiller191 + + + + + H- + + + Micracanthodinium claytonii (Holmes) Dodge168 + H- 22,140 Pyrocystis noctiluca Murray ex Schütt192 ) + + ■f + + + + Micracanthodinium setiferum (Lohmann) Deflandre169 4- + + + + Pyrocystis obtusa Pavillard Spiraulax Kofoid 1911 (= Gonyaulax Diesing Pyrocystis robusta Kofoid + + + + + + 1866 partim.) Dinoflagellates of uncertain identification Spiraulax jolliffei (Murray et Whitting) Kofoid 4- 4- + + + + + Adinimonas Schiller 1928 Prorocentrales Lemmermann 1910 Adinimonas oviforme Schiller193 + + + + Prorocentraceae Stein 1883 Archaeosphaerodiniopsis Rampi 1943 Exuviella Cienkowski 1881170 Archaeosphaerodiniopsis verrucosa Rampi194 + ^ 5 Exuviella aperta Schiller171 4- + + 77,145,168 Pachydinium Pavillard 1915 Mesoporos Lillick 1937 (= Porella Schiller 1928) Pachydinium mediterraneum Pavillard195 + + + Mesoporos globulus (Schiller) Lillick 4- 4- + + + Mesoporos perforatus (Gran) Lillick + 4- 4- + + + + + + Prorocentrum Ehrenberg 1834170 (= Exuviella Cienkowski 1881) Prorocentrum aporum (Schiller) Dodge 4- 4 - + + + + + Prorocentrum arcuatum Issel172 + + + + + 232 F. Gómez Checklist of Mediterranean free-living dinoflagellates 233

Notes mikimotoi (Miyake et Kominami ex Oda) G. Hansen et sembles C. tripos var. pulchellum (Schröder) López, see Moestrup. Gymnodinium pulchellum is distinguished Sournia (1967). 1 Reported in the Western Mediterranean Sea by Gómez 28 Histioneis splendida Murray et Whitting. from K. mikimotoi by the sigmoid apical groove. See 82 Ceratium arcuatum (Gourret) Pavillard, C. tripos var. and Claustre (2003). These records assigned to Astero­ 29 Ornithocercus serratus Kofoid, O. orbiculatus Kofoid et also Note 44. arcuatum Gourret, non C. arcuatum Cleve. dinium gracile Sournia presented morphological dif­ Michener. 52 This taxon resembles Katodinium glaucum (Lebour) 83 Ceratium fusus var. extensum Gourret. ferences with respect the type species.Asterodinium 30 Histioneis karstenii Kofoid et Michener. Loeblich III. 84 C. tripos var. horridum Cleve, but C. tenue (Ostenfeld et libanum Abboud-Abi Saab requires a more detailed 31 Dinophysis acutoides Balech, Phalacroma acutum 53 Gyrodinium opinum (Schütt) Lebour. Schmidt) Jörgensen, C. intermedium (Jörgensen) Jör­ description. Pavillard. 54 Gyrodinium dominans Hulburt. gensen and C. buceros (Zacharias) Schiller have not 2 The type speciesBrachydinium capitatum F.J.R. Taylor 32 Phalacroma stenopterygium Jörgensen. 53 “lacryma ” (= tear-drop) should be the correct epithet of been considered as synonyms. (Taylor 1963) was replaced by Brachidinium capitatum 33 Pseudophalacroma nasutum (Stein) Jörgensen, Dino­ this taxon. 85 This taxon resembles Ceratium limulus (Gourret ex F.J.R. Taylor due to an etymological error (Taylor 1967). physis nasuta (Stein) Parke et Dixon. 56 Gymnodinium spirale var.pepo Schütt. Pouchet) Gourret. Sournia (1973 p.5) reported that the correction is invalid. 34 Gymnodinium sanguineum Flirasaka, G. splendens 37 Gymnodinium spirale var. pinguis Schütt. 86 C. tenuissimum Kofoid. 3 Rare dinoflageilate epiphytic on Rhodophyceae (see Lebour. 58 Gymnodinium breve Davis, Ptychodiscus brevis (Davis) 87 Ceratium carriense var. volans (Cleve) Sournia. Sournia 1986, p. 36). 35 Amphidinium phaeocysticola Lebour has been consid­ Steidinger. 88 Acanthodinium caryophyllum Kofoid. 4 Rare and insufficiently described taxon (Sournia 1986, ered as a synonym of A. crassum Lohmann. However 59 Reported as Gymnodinium galatheanum Braarud 89 Micracanthodinium quadrispinum (Pavillard) Margalef. p. 36). this synonymy is debatable (Elbrächter 1979). (= Gyrodinium galatheanum (Braarud) Taylor sensu 90 Confusion possible with Cladopyxis brachiolata Stein. 5 According to Taylor (1976 p. 190), the cysts were report­ 36 Amphidinium lacustre Stein, A. schroederi Schiller and Taylor). More recently this taxon,unless Gymnodinium 91 Heterodinium detonii Rampi. ed by Margalef et al. (1954). Gloeodinium Klebs and A. lacustriformis Schiller are often considered as syn­ galatheanum Braarud sensu Kite et Dodge, is consid­ 92 Crypthecodinium setense Biecheler. Hemidinium Stein have been considered as the immo­ onyms. Typically fresh and brackish water species. ered as a synonym of Karlodinium micrum (Leadbeater 93 Goniodoma polyedricum (Pouchet) Jörgensen, Heter­ bile and mobile stage respectively of the life cycle of the 31 Amphidinium klebsii Kofoid et Swezy. et Dodge) J. Larsen (Daugbjerg etal. 2000). aulacus polyedricum (Pouchet) Drugget Loeblich, Tria­ same taxa (see Sournia 1986 p. 67). The continental 38 Considered as a synonym of Amphidinium lacustri­ 60 Massartia glauca (Lebour) Schiller, Gyrodinium glau­ dinium polyedricum (Pouchet) Dodge, Goniodoma species, Hemidinium nasutum Stein and others, are re­ forme Schiller by Dodge (1982 p. 72). cum (Lebour) Kofoid et Swezy, Gymnodinium minutum polyedra Rampi. ported in the Mediterranean waters (e.g., Schiller 39 Gymnodinium filum Lebour. Lebour, Massartia minuta (Lebour) Conrad et Kuf­ 94 Heteraulacus sphaericum (Murray et Whitting) Loe­ 1935-1937,p. 89-92, Vilicicetal. 2002). 40 Taylor (1976 p. 114) reported this taxon from the Liguri­ ferath, Massartia tubulata Rampi. blich III, Triadinium sphaericum (Murray et Whittman) 6 Syracosphaera heimii Lohmann. This taxon was previ­ an Sea. 61 Massartia tubulata Rampi. Dodge. ously considered to be a coccolithophorid and has been 41 Non Cochlodinium helix Kofoid et Swezy (= Cochlo­ 62 Related to the genus Gyrodinium Kofoid et Swezy ac­ 95 Reported as Pyrodinium schilleri (Matzenauer) Schiller scarcely reported in dinoflageilate checklists (see Tan­ dinium helicoides Lebour). cording to Sournia (1986, p. 57). [= Pyrodinium bahamense Plate var. compressum gen et al. 1982). 42 Confusion possible between Cochlodinium poly­ 63 Gymnodinium teredo Pouchet. (Böhm) Steidinger, Tester et Taylor], 7 This taxon resembles Amphisolenia spinulosa Kofoid krikoides Margalef (= C. heterolobatum Silva) and 64 Ptychodiscus inflatus Pavillard, P. carinatus Kofoid. 96 Alexandrium lusitanicum Balech. and Amphisolenia mozambica Sournia. Gymnodinium impudicum (Fraga et Bravo) G. Hansen 65 Erythropsis agilis Hertwig. Probably several species are 97 Reported as Goniodoma ostenfeldii Paulsen by Lecal 8 This taxon presents synonyms as Dinophysis borealis et Moestrup (see Choet al. 2001). reported as E. agile (Hertwig) P.C. Silva according to (1954). Paulsen, D. lachmanni Paulsen, D. boehmii Paulsen or 43 Gymnodinium albulum Lindemann and G. simplex Elbrächter (1979). 98 Reported as Gonyaulax subulata Kofoid et Michener. D. skagii Paulsen. (Lohmann) Kofoid et Swezy may be synonyms. 66 Elbrächter (1979) considered this taxon as a synonym This taxon resembles Amylax triacantha (Jörgensen) 9 Dinophysis dens Pavillard. 44 Tlie North European taxon, Gyrodinium aureolum of E. agile (Hertwig) P.C. Silva. Sournia (Dodge 1982, p. 217). 10 The orthographical similarity of Dinophysis alata Jör­ Hulburt sensu Braarud el Heimdal, is a synonym of 67 To the best of my knowledge, never reported after the 99 Reported as Gonyaulax? triacantha Jörgensen by Lecal gensen, Dinophysis alata Böhm and Dinophysis alata Karenia mikimotoi (Miyake et Kommami ex Oda) G. initial description by Greuet (1968b). (1954). (Wood) Balech is confusing. Vilicic et al (2002) report­ Hansen et Moestrup (= Gymnodinium nagasakiense 68 Pouchetia armata Dogiel, Pouchetia maculata Kofoid et 100 Gonyaulax spinifera sensu Schütt. ed Dinophysis alata (Wood) Balech. Takayama et Adachi) (Hansen et al. 2000). See also Swezy. 101 Gonyaulax digitale Kofoid, Protoperidinium digitale 11 Dinophysis amygdala Balech, Phalacroma ovum Note 51. 69 Kofoidinium lebourae (Pavillard) Taylor (= Gymno­ Pouchet. Schütt, nonDinophysis ovum Schütt. 45 According to Bolch and Reynolds (2002) other taxa dinium lebourae Pavillard). 102 Resembles Gonyaulax birostris Stein. 12 This taxon resembles Phalacroma ovatum (Claparède that also produce microreticulate cysts such asGymno­ 70 Originally described from the Ligurian Sea as Lepto­ 103 Reported by Narusevich and Tokarev (1989) in an un­ et Lachmann) Jörgensen. dinium nolleri EUegaard et Moestrup andG. microretic­ dinium caudatum Cachón et Cachón. determinated location of the Mediterranean Sea. 13 Dinophysis caudata var. diegensis Kofoid. ulatum Bolch et Hallegraeff are present in the Tyrrhen­ 71 Pratjetella medusoides (Hertwig) Loeblich et Loeblich 104 The comments by Schiller (1935-1937, p. 290) on the 14 Dinophysis intermedia Pavillard, Dinophysis laevis ian and Adriatic Seas based on the cysts reported by similarity between Gonyaulax kofoidii and G. pacifica Pouchet. III. Doubtful taxon (Sournia 1986, p. 53). Montresor et al. (1998) and Rubino et al. (2000). 72 Abedinium dasypus (Cachón et Cachón) Loeblich et Kofoid could induce confusion between both taxa 15 Phalacroma odiosum Pavillard. 46 The records of Gymnodinium catenatum Graham by Loeblich III. (Pavillard 1937, p. 16; Taylor 1976, p. 104). 16 Phalacroma mitra Schütt, Phalacroma rapa Stein, Pha­ Carrada et al. (1991), Giacobbe et al. (1995) and Labib 73 Reported from the Ligurian Sea asLeptospathium nav­ 105 Gonyaulax minima Matzenauer. lacroma dolichopterygium Murray et Whitting. (1997) are considered as G. impudicum (Fraga et Bra­ icula Cachón et Cachon-Enjumet (1964) after the de­ 106 Resembles Gonyaulax ovata Matzenauer (Schiller 17 Non Phalacroma ovum Schütt. vo) G. Hansen et Moestrup. scription by Margalef (1963). Balkis (2000) reported 1935-1937, p. 289; Taylor 1976, p. 105). 18 Dinophysis infundibula Schiller. 47 Gymnodinium conicum Kofoid et Swezy (= G. viridis this taxon from the Marmara Sea. 107 Pavillardinium brianii (Rampi) Sournia (=Murrayella 19 Dinophysis lenticula Pavillard. Lebour) is considered as a synonym of Gyrodinium 74 Resembles Pronoctiluca acuta (Lohmann) Schiller. brianii Rampi). 20 Dinophysis reticulata (Kofoid) Balech. 108 viridescens Kofoid et Swezy. Non Gyrodinium conicum 75 Oxytoxum margalefii Rampi, Oxytoxum tonollii Rampi. Gonyaulax levanderi (Lemmermann) Paulsen, Cerato­ 21 Dinophysis acuminata f. reniformis Pavillard, D. pavil­ Schiller. 76 Murrayella spinosa Kofoid, Pavillardinium spinosum corys spinifera Schröder. lardii Schröder, D. reniformis (Pavillard) Kofoid et 48 Gymnodinium punctatum var. grammaticum Pouchet. (Kofoid) Taylor ex Sournia, Amphidoma spinosa (Ko­ 109 Gonyaulax diacantha (Meunier) Schiller, Gonyaulax Skogsberg, D. ventrecta Schiller. 49 Gymnodinium rhomboides Schütt,G. hyalinum Lebour foid) Kofoid et Michener, Gonyaulax rouchii Rampi. longispina Lebour, Amylax diacantha Meunier. 22 Dinophysis sphaeroidea (Schiller) Balech. (= G. lucidum Ballantine in Parke el Dixon). Gyrodini­ 77 This taxon resembles Ceratium incisum (Karsten) Jör­ 110 Gonyaulax milneri (Murray et Whitting) Kofoid, Go­ 23 Dinophysis uracantha Schütt, non Dinophysis uracan­ um striatissimum (Hulburt) G. Hansen et Moestrup has niodoma milneri Murray et Whitting. tha Stein. gensen. been considered as a synonym until the redescription of 78 Ceratium buceros f. claviger (Kofoid) Schiller, Ceratium 111 Gonyaulax polyedra Stein. 24 Dinophysis sphaerica Pavillard G. heterostriatum Kofoid et Swezy by Elbrächter (1994). horridum f. claviger (Kofoid) Sournia. 112 Gonyaulax grindleyi Reinecke, non G. reticulatum Ko­ 25 A possible variety of Histioneis depressa Schiller (Tay­ 50 This brackish waters taxon appears associated with 79 Also reported as Ceratium trichoceros var. contrarium foid et Michener. lor 1976, p. 44). Karenia mikimotoi (Miyake et Kominami ex Oda) G. (Gourret) Schiller. 113 Heterodinium laticinctum Kofoid. 26 Ornithocercus carolinae Kofoid, Histiones francescae 114 Hansen et Moestrup (see also Notes 44 and 51). 80 Ceratium horridum var. lenticulatum Jörgensen, C. Heterodinium pulchrum Böhm, Heterodinium richardii Murray et Whitting. 51 According to Faust and Gulledge (2002) this taxon was buceros f. denticulatum (Jörgensen) Schiller. Pavillard. 27 Ornithocercus assimilis Jörgensen, O. galea (Pouchet) recorded in the Tyrrhenian Sea by Carrada et al. (1991). 115 Heterodinium mediocre f. sinistrum (Kofoid) Kofoid et Abé. 81 Reported as Ceratium pulchellum f. eupulchellum by Confusion possible with species of the complexKarenia Ghazzawi (1939) in the Canal of Suez. This taxon re­ Adamson. 234 F. Gómez Checklist of Mediterranean free-living dinoflagellates 235

116 Ostreopsis monotis (Meunier) Lindemann. 148 Peridinium decipiens var. curvipes Ostenfeld, Protoperi­ 175 Prorocentrum minimum (Pavillard) Schiller and Proro­ trast, Pyrocystis is a permanently free-living pho- 117 Ceratium biconicum Murray et Whitting, Murrayella bi­ dinium subcurvipes (Lebour) Balech. centrum cordatum (Ostenfeld) Dodge may be syn­ totrophic dinoflageilate with a predominant coccoid conica (Murray et Whitting) Pavillard and Pavillardini­ 149 Protoperidinium longipes (Karsten) Balech. onyms (Velikova and Larsen 1999). stage and a flagellated dinospore stage (e.g., Elbrächter um biconicum Rampi are considered synonyms. 130 According to Balech (1976) this taxon is related to the 176 Prorocentrum obtusidens Schiller, also P. monacense etal. 1987). 118 Pavillardinium intermedium (Pavillard) de Toni freshwater species Protoperidinium achromaticum Kufferath described in the Ligurian Sea (Kufferath 187 Pyrocystis lunula (Schütt) Schütt,Dissodinum lunula (= Murrayella intermedia Pavillard), non Centrodinium (Levander) Balech. 1957). (Schütt) Pascher. intermedium Pavillard. 131 Protoperidinium ovatum Pouchet [= P. globulus var. 177 P. hentschelii Schiller, P. sigmoides Böhm, P. macrurus 188 Dissodinium fusiformis (Thompson ex Murray) 119 Pavillardinium splendidum (Rampi) Rampi (= Mur­ ovatum (Pouchet) Schiller, Peridinium ovatum Athanassopoulos. Resembles P. micans Ehrenberg. Matzenauer. rayella splendida Rampi). (Pouchet) Schütt] have been considered as synonyms. ns According to Dodge (1975): Prorocentrum mexicanum 189 Dissodinium gerbaultii (Pavillard) F.J.R. Taylor 120 Resembles Corythodinium reticulatum (Stein) Loeblich 132 This taxon can be confused with Protoperidinium ovum Osorio-Tafall, P. obtusum Ostenfeld, P. brochi Schiller, 190 Léger (1973) reported the presence of this taxon in the et Loeblich III. (Schiller) Balech. P. ovale Schiller, P. ovalis Rampi,P. rampii Sournia. Spanish Mediterranean coast by Margalef et al. (1957). 121 Prorocentrum curvatum Kofoid. 153 Also reported from the Tyrrhenian Sea based on cysts 179 prorocentrum schilleri Böhm in Schiller. Drebes (1981) reported that Pyrocystis margalefii 122 Non Corythodinium globosum (Kofoid) Taylor. by Montresor et al. (1998). iso Prorocentrum nanum Schiller and P. pusillum (Schiller) Léger is probably identical with the resting stages of 123 Oxytoxum sceptrum (Stein) Schröder. 154 Protoperidinium spiniferum (Schiller) Balech. Loeblich were considered as synonyms until Puigserver Dissodinium pseudolunula Swift ex Elbrächter et 124 Oxytoxum challengeroides Kofoid. 155 The orthographic similarity between P. sphaeroides and Zingone (2002). Drebes. 123 Oxytoxum mediterraneum Schiller. (Dangeard) Balech and P. sphaeroideum (Mangin) 181 Prorocentrum cornutum Schiller. 191 Dissodinium minimum Matzenauer. 126 Oxytoxum tenuistriatum Rampi. Balech is confusing (Sournia 1978, p. 29). 182 Prorocentrum sphaeroideum Schiller, P. robustum Oso­ 192 Pyrocystis pseudonoctiluca Wyville-Thomson ex Mur­ 127 This taxon resembles Oxytoxum ovale Schiller 156 This taxon resembles Protoperidinium brachypus rio Tafall. ray, Dissodinium pseudolunula Swift ex Elbrächter et 128 Oxytoxum ligusticum Rampi. (Schiller) Balech. 183 Prorocentrum adriaticum Schiller. Drebes. 129 Oxytoxum gracile Schiller. 157 Protoperidinium angustum (Dangeard) Balech. 184 This taxon resembles Prorocentrum mexicanum Oso­ 193 Regarded as a doubful dinoflageilate by Sournia (1986, 130 Murrayella ovalis Pavillard. See also comments on the 158 Most of the recently described species of Scrippsiella rio-Tafall that is here considered a synonym of Proro­ p. 97). Commonly reported asAdinomonas Schiller. genus by Sournia (1986, p. 73). Balech ex Loeblich are reported from the germination centrum maximum (Gourret) Schiller. 194 Regarded as a doubtful dinoflageilate by Sournia (1986, 131 Gonyaulax mitra (Schütt) Kofoid, Oxytoxum gigas Ko­ of cysts (Montresor et al. 1994, D ’Onofrio et. al 1999). iss Regarded as a doubtful dinoflageilate by Sournia (1986, p. 97). foid. 139 Scrippsiella faeroense Dickensheets et Cox, non Scripp­ p. 37). 193 Doubtful taxon (Sournia 1986, p. 98). Also reported 132 Based on the germination of recent cysts (D’Onofrio siella faeroense (Paulsen) Balech et Soares. 186 Dissodinium is a genus of parasitic dinoflagellates with from the Sicilian coasts or North-Italian lakes by An­ etal. 1999). 160 Reported by Rampi (1941) as Blepharocysta striata a complicate life cycle including planktonic life cycle dreis et al. (1982). 133 Meier et al. (2002) reported 14 species of calcareous di­ Schütt (see Carbonell-Moore 1994). stages similar to those of the genus Pyrocystis. In con- noflagellates from recent cysts (4 new species), only this 161 Podolampas spinifer Pavillard. taxon that germinated from one cyst from the Levan­ 162 Tuberculodinium vancampoae (Rossignol) Wall tine B asin is included. (= Pterospermopsis vancampoae Rossignol). Taylor References 134 Diplopsalis asymmetrica (Mangin) Lindeman, Diplop­ (1976 p. 183) reported the presence of this taxon in the salopsis bomba (Stein) Dodge et Toriumi, Dissodium Mediterranean Sea based on Margalef (1948). 1. Abboud-Abi Saab, M. 1985. Étude quantitative et qual­ 13. Bravo, I., B. Reguera, A. Martinez and S. Fraga. 1990. asymmetricum (Mangin) Loeblich III. 163 One specimen that resembles C. yeye Margalef from the itative du phytoplancton des eaux côtières libanaises. First report ofGymnodinium catenatum Graham on the 133 Considered as a synonym of Diplopelta bomba Stein ex Alborán Sea, one specimen of C. yeye and other unde­ Lebanese Sei. Bull, h 197-222. Spanish Mediterranean coast. In: (E. Granéli, B. Sund­ Jörgensen by Dodge (1982 p. 157). terminated species of this genus were observed from 2. Abboud-Abi Saab, M. 1989. Les dinoflagellés des eaux ström, L. Edler and D. M. Anderson, eds) Toxic Marine 136 Dissodium lenticulum (Bergh) Loeblich III, Glenodini­ the Balearic coasts (unpublished obs.). Ceratoperidini­ côtières libanaises- Espèces rares ou nouvelles du phy­ Phytoplankton. Elsevier. Amsterdam, pp.26-30. um lenticula (Bergh) Schiller. um mediterraneum Abboud-Abi Saab requires a more toplancton marin. Lebanese Sei. Bull. J: 5-16. 14. Cachón, J. and M. Cachon-Enjumet. 1964. Leptospathi­ 137 Diplopsalis orbicularis (Paulsen) Steidinger et Williams. detailed description. 3. Andreis, C., M. D. Ciapi and G. Rodondi. 1982. The the­ um navicula nov. gen, nov. sp. etLeptophyllus dasypus 138 Usually a brackish water species. 164 Iwataki et al. (2002) reported this taxon based on the cal surface of some Dinophyceae: A comparative SEM nov. gen. sp., péridiniens Noctilucidae (Hertwig) du 139 Glenodinium rotundum (Lebour) Schiller. material from the Aegean Sea by Pennik and Clarke approach.Bot. Mar. 25:225-236. plancton néritique de Villefranche-sur-Mer.Bull. Inst. 140 Originally described from the Tyrrhenian Sea as Peri­ (1977), Océanogr. Monaco 62 (1292): 1-12. dinium tyrrhenicum Balech (Balech 1990). 4. Balech, E. 1976. Sur quelques Protoperidinium (Di­ 163 Cachonina niei Loeblich III. 15. Cachón, J. and M. Cachon-Enjumet. 1966. Pomatodini­ 141 Nearly all of the marine species of Peridinium Ehren- noflagellata) du Golfe du Lion.Vie Milieu 16:27-46. 166 Katodinium rotundatum (Lohmann) Loeblich III, Mas­ um impatiens nov. gen. nov. sp. péridiniens Noctilucidae berg have been transferred to Protoperidinium Bergh. 5. Balech, E. 1988. Los Dinoflagelados del Atlántico Su­ sartia rotundata (Lohmann) Schiller, Amphidinium ro­ Kent. Protista logica 2: 23-30. 142 Protoperidinium quinquecorne (Abé) Balech. tundatum Lohmann, Katodinium minutum (Lebour) doccidental. Pub. Esp. Inst. Español Oceanogr. Madrid 16. Cachón, J. and M. Cachón. 1967. Cymbodinium elegans 143 This taxon presents synonyms such asDiplopsalis minor Sournia. 1:310 pp. nov. gen. nov. sp. péridinien Noctilucidae Saville-Kent. (Paulsen) Lindemann, Zygabikodinium lenticulatum 167 Cladopyxis bacillifera Schiller. 6 . Balech, E. 1990. Four new dinoflagellates.Helgoländer Protistologica 3: 313-318. (Manguin) Loeblich et Loeblich III, Diplopeltopsis mi­ 168 Cladopyxis claytonii Holmes. Meeresunters. 44:387-396. nor (Paulsen) Pavillard, Diplopsalis lenticula f. minor 169 Cladopyxis setifera Lohmann, Micracanthodinium 7. Balkis, N. 2000. Five dinoflageilate species new to Turk­ 17. Cachón, J. and M. Cachón. 1967. Contribution à l'étude Paulsen (see Dodge and Toriumi 1993, Elbrächter 1993). bacilliferum (Schiller) Deflandre. ish seas. Oebalia 26: 97-108. des Noctilucidae Saville-Kent, I. Les Kofoidininae Ca­ 144 Non Protoperidinium biconicum (Dangeard) Balech. chón J. et M. Évolution, morphologique et systéma­ 170 The genus Exuviella was included in Prorocentrum by 8. Biecheler, B. 1938. Sur un péridinien cuirassé incolore 145 Minuscula bipes (Paulsen) Lebour. tique. Protistologica 3: 427-444. Dodge (1975). McLachlan et al. (1997) proposed the nouveau Crypthecodinium n. g. setense n. sp. et la 146 Protoperidinium bimucronatum (Schiller) Balech. The separation of both genera. famille nouvelle des Crypthecodiniacées. Bull. Soc. 18. Cachón, J. and M. Cachón. 1969. Contribution à l'étude synonymy between Peridinium sourniai F.J.R. Taylor 171 Schiller (1931-1933, p. 26) reported this taxon asExu­ zool.Fr. 63:9- 13. des Noctilucidae Saville-Kent. Évolution, mor­ and Protoperidinium bispinum (Schiller) Balech is de­ viella (?) aperta Schiller (described from the Adriatic phologique, cytologie, systématique. II. Les Leptodisci- 9. Biecheler, B. 1939. Sur deux péridiniens nouveaux des batable. Sea in 1928). Inadequate description according to nae Cachón J. et M. Protistologica 5: 11-33. 147 According to Schiller (1935, p. 223) Peridinium curtipes eaux saumâtres des environs de Sète.Bull. Soc. zool. Fr. Dodge (1975). 19. Carbonell-Moore, M. C. 1994. On the biogeography of Jörgensen is a synonym of Peridinium crassipes Kofoid, 64:12-18. 172 P. micans var. gibbosum Schiller, P. gibbosum (Schiller) the f am ily Podolampadaceae Lindemann (Dino­ consequently a confusion could be expected.Protoperi­ 10. Böhm, A. 1933. Neue Peridineen aus der Adria. Arch. Schiller, P. blatta Athanassopoulos. Resembles Proro­ phyceae) -vertical and latitudinal distribution. Rev. dinium crassipes (Kofoid) Balech and Protoperidinium Prostistenk. 80:351-354. centrum micans Ehrenberg. Palaeobot. Palynol. 84: 23-44 curtipes (Jörgensen) Balech are different species: 1) 173 According to Dodge (1975): Prorocentrum bidens 11. Bolch, C. J. S. and M. J. Reynolds. 2002. Species resolu­ 20. Caroppo, C. 2000. The contribution of picophytoplank- Protoperidinium crassipes (Kofoid) Balech (= Peridini­ Schiller, P. lebourae Schiller, Exuviella oblonga Schiller tion and global distribution of microreticulate dinofla- ton to community structure in a Mediterranean brack­ um crassipes Kofoid), 2) Protoperidinium curtipes (Jör­ [= Prorocentrum oblongum (Schiller) Taylor], E. lentic­ gellate cysts. /. Plankton Res. 24: 565-578. ish environment./. Plankton Res.22: 381-397. gensen) Balech (= Peridinium crassipes Paulsen 1907, ulata Matzenauer, E. elongata Rampi. 12. Bouquaheux, F. 1971.Gloeodinium marinum nov. sp. non Paulsen 1930). See also Balech (1988, p. 110). 174 Prorocentrum pyriformis (Schiller) Hasle. péridinien dinocapsale.A rch. Protistenk. 113: 314-321. 21. Caroppo, C. and N. Cardellicchio. 1995. Preliminary 236 F. Gómez Checklist of Mediterranean free-living dinoflagellates 237

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Staz * Corresponding author: [email protected] plankton composition and biomass from May 1988 to Zool. Napoli 40: 133-237. April 1989.Helgoländer Meeresunters. 44:487-501. Tolomio, C. 1978. Catalogo delle diatomee e delle peridinee Ounissi, M. and H. Fréhi. 1999. Variabilité du microphyto- piu'significative segnalate nelle acque salmastre ital- plancton et des Tintinnida (Protozoaires Ciliés) d'un iane. Mem. Biol. Mar. Oceanogr. 8 : 129-150. Four species of the red algal genusD asya (Dasyaceae, Ceramiales),including two new species, are reported from secteur eutrophe du Golfe d'Annaba (Méditerranée Tolomio, C. 1988. Taxonomical and ultrastructural observa­ Malaysia. Dasya iyengarii B prgesen andD. pilosa (Weber-van Bosse) Millar are reported from Malaysia for the sud-occidentale). Cah. Biol. Mar. 40: 141-153. tion on a Prorocentrum (Dinophyceae) collected in first time, and their morphological features are described along with taxonomic comments.Dasya longifila Ma- Pagou.K. and O. Gotsis-Skretas. 1990. A comparative study southern Corsica coastal waters. Bot. Mar. 31:223-229. suda et Uwai, sp. nov. is chiefly distinguished by its long, monosiphonous, pseudolateral filaments (often longer of phytoplankton in Aegean, Levantine and Ionian seas Tolomio, C. and F. Cavolo. 1985. Presenza di Coolia monotis than the axis) in which intercalary cell divisions take place, and a small number of tetrasporangial stichidia and during March-April 1986.Thalassografica 13: 13-18. Meunier (Dinophyceae, Peridiniales) nelle acque della spermatangial branches per fertile pseudolateral from several related species that possess small-statured, Paulmier, G., B. Berland, C. Billard and E. Nezan. 1995. Gy­ Laguna di Venezia. Oebalia 11: 849-852. sparsely corticated axes.Dasya malaccensis M asudaet Uwai, sp. nov. is characterised by its diminutive thallus (up rodinium corsicum nov. sp. (Gymnodiniales, Dinophy- Tolomio, C. and M. Lenzi. 1996. “Colored waters” in the Or- to 1 0 mm high) with thickly corticated axes, relatively short pseudolaterals, and narrow tetrasporangial stichidia cées), organisme responsable d’une “eau verte” dans l’e- betello and Burano Lagoons (northern Tyrrhenian Sea) and spermatangial branches. This species has no close relatives among small-sized species ofDasya. tang marin de Diana (Corse) en Avril 1994. Cryptogam. from 1986 to 1989. Vie Milieu 46:25-37. Algol 16:11 -94. Totti, C., G. Civitarese, F. Acri, D. Barletta, G. Candelari, E, Polat, S., E. Sarihan and T. Koray. 2000. Seasonal changes in Paschini and A. Solazzi. 2000, Seasonal variability of Introduction dried as voucher herbarium specimens or mounted in the phytoplankton of the NE Mediterranean (Bay of phytoplankton populations in the middle Adriatic sub­ 30% Karo® on microscope slides and deposited in Iskenderun). Turkish J. Bot. 24:1-12. basin. 7. Plankton Res. 22: 1735-1756, The red algal genus D asya belonging to the Dasy­ the Herbarium of the Graduate School of Science, Rabatti, S., F. Bianchi, A. Boldrin, L. Da Ros, G. Socal and Travers, A. and M. Travers. 1963. Contribution a l'étude du aceae (Ceramiales) is characterised by: 1) its radially Hokkaido University (SAP), or in the Seaweed C. Totti. 1994. Particulate matter and phytoplankton in microplancton du canal de Corse. Rapp. P. -v. Réun. branched and sympodially developed axes issuing nu­ Herbarium, Institute of Biological Sciences, Univer­ the Ionian Sea. Oceanol. Acta 1 7:297-307. Commn int. Explor. Mer Méditer. 77:487-493. merous, pigmented, monosiphonous pseudolaterals sity of Malaya (KLU). Sections were made by hand Rampi, L. 1939. Péridiniens rares ou intéressants dans la Vilicic, D. 1985. An examination of cell volume in dominant (usually from every segment) that are subdichoto- using a razor blade and stained with 0.5 % (w/v) mer Ligure. Bull. Soc.fr. Microscopie 8:106-112. phytoplankton species of the Central and Southern mously branched several times and usually persistent; cotton blue in a lactic acid/phenol/glycerol/water Rampi, L. 1939. Ricerche sul fitoplancton del mare Ligure Adriatic Sea, Int. revue ges. Hydrobiol, 70:829 -843. 2 ) tetrasporangial stichidia formed from a branch of (1:1:1:1 [v/v]) solution and m ounted in 50% glyc­ 1.1 Ceratium dele acque di Sanremo. Nuovo Giorn. Bot. Vilicic, D. 1989. Phytoplankton population density and vol­ monosiphonous filaments including pseudolaterals erol/seawater or 30 % Karo® on microscope slides. Ital. N. Ser. 46:299-312. ume as indicator of eutrophication in the eastern part of and bearing whorls of up to 7 tetrasporangia and 2-4 Rampi, L. 1943. Ricerche sul fitoplancton del mare Ligure the Adriatic Sea. Hydrobiologia 174: 117-132. postsporangial cover cells; 3) spermatangial branches 7. Le Goniaulacee delle acque di Sanremo.Att. Soc. Ital. Vilicic, D. 1991. A study of phytoplankton in the Adriatic also produced from a branch of monosiphonous fila­ Sc. Nat. Milano 82: 318-327. Sea after the July 1984 bloom. Int. revue ges. Hydrobiol. ments including pseudolaterals; and 4) the absence of Observations and Discussion Rampi, L. 1945. Osservazioni sulla distribuzione qualitativa 76:197-211. pericarp initials at fertilisation (Parsons 1975,Maggs Dasya iyengarii Borgesen1937:345,f. 16,17 del fitoplancton nel mare Mediterráneo. Att. Soc. Ital. Vilicié, D. and N. Fanuko. 1984. A study of the phytoplank­ and Hommersand 1993, Parsons and Womersley Sc. Nat Milano 84:105. (Figs 1 -9 ) ton in offshore waters of southern Adriatic, January 1998). D asya currently includes more than 80 species Rampi, L. 1950. Ricerche sul fitoplancton del mare Ligure 1980. Nova Thalassia 6: 67-82. that are widely distributed in tropical to temperate Distribution: Tropical regions in the Pacific Ocean 9.1. Peridinium delle acque di Sanremo. Atti Acc. Ligure Viliéic, D., Z. Vucak, A. Skrivanic and Z. Grzetic. 1989. Phy­ seas of the world (Parsons and Womersley 1998). (Price and Scott 1992, present paper) and the Indian Sc. Lett. 7:1-10. toplankton blooms in the oligotrophic open south Adri­ However, no species of the genus had been reported Ocean (Silva et al. 1996, present paper). Rampi, L. 1954. Variazioni stagionali del fitoplancton di su­ atic waters. Mar. Cheni. 28:89-107. from Malaysian waters (Phang and Wee 1991), which perficie raccolto nel golfo di Genova a Punta del Mesco Specimens examined:Kedah: Tanjung Datai Besar Zingone, A., M. M ontresor and D. Marino. 1998. Morpho­ are a very im portant region in assessing the relation­ (La Spezia). Atti Acc. Ligure Sc. Lett. Genova 10: 1- (6°25’51”N, 99°40’25”E; 19.xii.1997; tetrasporangial logical variability of the potentially toxic dinoflageilate 12. ship between the marine algal flora of the Pacific and SAP 093314,093315), Pulau Langkawi. Sarawak: Pu- Dinophysis sacculus (Dinophyceae) and its taxonomic that of the Indian Ocean. In the present paper we re­ lau Talang-Talang Besar (1°55’02”N, 109°46’28”E; Revelante, N. and M. Gilmartin. 1977. The effects of North­ relationships with D. pavillardii and D. acuminata. Eta: port four species ofD asya including two new species ern Italian rivers and eastern Mediterranean ingression J. Phycol. 33: 259-273. 26.V.1998; tetrasporangial SAP 093317-093320), Se- on the basis of our recent collections from Malaysia. matan. Sabah: Pantai Layang Layangan (5°20T3”N, 115°H’43”E; l.vi.1998; tetrasporangial SAP 093321), Pulau Labuan; Pulau Gaya (6°01’21”N, 116°03,01”E; Material and Methods 4 vi 1998; tetrasporangial SAP 093322), Kota Specimens examined were collected at various locali­ Kinabalu; Black Rock (6°06’55”N, U8°00’04”E; ties in Malaysia by us. The specimens were fixed in 16.V.1998; sperm atangial SAP 093316), Sandakan. Terengganu: Tanjung Batu Lepir (5°53’08”N, 10% Formalin in seawater, and later some were