SMITHSONIAN INSTITUTION Contributions from the United States National Herbarium Volume 42: 1-144 Identifying Harmful Marine Dinoflagellates by Maria A. Faust and Rose A. Gulledge Department of Systematic Biology - Botany, National Museum of Natural History Washington, DC 2002 ABSTRACT Faust, Maria A. and Rose A. GuUedge. Identifying Harmful Marine Dinoflagellates. Smithsonian Contributions from the United States National Herbarium, volume 42: 144 pages (including 48 plates, I figure and 1 table). - A taxonomic identification and reference guide of 48 harmful marine dinoflagcUatc species present in the world's oceans, fhis guidebook illustrates the morphology and ta\onom\ of harmlul marine dinoOagellates of the following genera: Alexandrium, Dinophysis, Gymnodiniiim. Ostreopsis, Prorocentrum, Coolia, Cochlodinium, Gambierdiscus, Gonyaulax. Gyrodinium, Lingulodinium, and Pfiesteria. These organisms have been implicated in marine life mortality events and/or seafood-borne human diseases. Some species cause problems due to red tide conditions, others produce toxins; e.g. brevetoxins. ciguatoxins, dinoph\sisloxins. and ichlh>otoxins. Detailed taxonomic descriptions of plate and thecal morpholog\, and cellular structure are presented. Taxonomic treatment of species includes nomenclatural types, type locality, synonyms, and etymology. Information is also available on species reproduction, ecology, biogeograph\, distribution, and habitat and locality. Species illustrations presented as scanning electron micrographs, ditTerential interference contrast and epilluorescence light micrographs, as well as line drawings. A comprehensive glossary list and literature reference section is included. Kofoidian plate tabulation followed for armored species plate designation. The International Code of Botanical Nomenclature (ICBN) was followed for the taxonomical treatment of species. This fully illustrated laboratory guide is intended for the researcher, instructor, and the student; it is the most comprehensive reference manual for identifying harmful dinotlageliate taxa. It can also serve as a field guide for marine biologists and environmental researchers. DATE OF PUBLICATION: April 2002 Cover Design: Illustrations by Alice Tangerini; front Prorocentnim hojjmaiuuaimm 1 aust; back Prorocentrum ruelzlerianum Faust. Contributions of the United States National Herbarium (IS.SN 0097-1618) Department of Systematic Biology - Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0166, USA. POSTMASTER; Send address changes to Contributions US National Herbarium, Department of Systematic Biology - Botany, National Museum of Natural Historv. Smithsonian Institution. Washington, DC, 20560-0166, USA. The paper used in this publication meets the minimum requirements of the .American National Standard for I'ermaneiicc of i'apcr for Printed Librar\ Materials /.3M 4.S-19S4 Contributions from the U.S. National Herbarium was first published in 1800 by The United St;ites Deparlmcnt of Agriculture. From July I, 1902 forward it was published as a Bulletin of the United Slates National Museum. The series was discontinued after volume 38, 1974, and has been revived with volume 39, as a venue for publishing longer taxonomic papers, checklists, floras, and monographs, produced by staff and associates at the U.S. National Herbarium. It is externally peer reviewed, and published at irregular intervals. Subscription and other correspondence should be addressed to CUSNH, Department of Systematic Biology - Botany. National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0166, USA e-mail CUSNH@nmnh.si.edu. The present issue is available for free while supplies last. CONTENTS INTRODUCTION DEDICATION IDENTIFYING HARMFUL MARINE DINOFLAGELLATES ACKNOWLEDGEMENTS IDENTIFYING HARMFUL MARINE DINOFLAGELLATE SPECIES Alexandrium acatenella A. catenella A. mlmttum A. monilatum A. ostenfeldii A. pseudogonyaulax A. tamarense A. tamiyavanichi Cochlodinium polykrikoides Coolia monotis Dinophysis acuminata D. acuta D. caudata D. forta D. miIra D. norvegica D. rotimdata D. sacculus D. tripos Gamhierdiscus toxicus Gonyaulax polygramma Gymnodmium breve G. catenatum G. mikimotoi G. pulchellum G. sangiiineum G. veneftcum Gyrodinium galatheanum Lingulodinium polyedrum Noctiluca scintillans Ostreopsis heptagona O. lenticularis O. mascarenensis O. ovata O. siamemis Pfiesleria piscicida Prorocentrum arenarium P. balticum P. belizeanum P. coticavum P. faustiae 5 5 5 9 10 10 97 11 98 13 99 14 100 15 101 17 102 18 103 20 104 21 105 22 106 23 107 25 108 26 109 28 110 29 111 30 112 32 113 33 114 34 115 35 116 37 117 37 118 39 119 40 120 41 121 42 122 43 123 44 124 45 125 46 126 47 127 49 128 50 129 51 130 52 131 53 132 54 133 55 134 56 135 57 , 136 58 137 p. hoffmannianum 59, 138 P. lima 60, 139 P. maculosum 61, 140 P. mexicanum 62, 141 P. micans 63, 142 P. minimum 65, 143 P. ruetzleriamim 66, 144 GLOSSARY 67 BIBLIOGRAPHY 73 Harmful Marine Dinoflagellates Identifying Harmful Marine Dinoflagellates Maria A. Faust and Rose A. Gulledge INTRODUCTION Interest in identifying harmful dinoflageilate species has received worldwide recognition in recent years due to the increase in red tides, fish kills, and shellfish poisoning events repoited from coastal marine ecosystems (Hallegraeff 1991). The publication, Identifying Harmful Marine Dinoflagellates, is an effort by the authors to present a fully illustrated identification guide for harmful dinoflageilate taxa. The user will recognize general information on dinoflageilate morphology and other criteria used in species identification. Each taxon is presented with a species overview, and a taxonomic description of cell and thecal plate morphology, reproduction, life cycle, ecology, toxicity, species comparison, habitat and locality, and etymology. This is supplemented with a number of high-resolution light and scanning electron photomicrographs and line drawings. Taxonomic treatment of harmful dinoflageilate taxa includes nomenclatural types, type locality, and synonyms. The nomenclatural name of a species is taken from the original publication of the taxa, with the exception of those where the type species is not known. Species names used in this publication are valid as of those published by 2000. An extensive glossary of terms and relevant literature citations are also provided. This guide will be useful to teachers, researchers and students, as well as professionals involved in environmental water quality assessment and management, fisheries and aquacuhure, and public health. DEDICATION Dr. Maria Faust would like to dedicate this work to her mentor. Dr. Grethe Hasle, Professor of Marine Botany, University of Oslo, Norway. Dr. Hasle has devoted much of her life to teaching; and sharing her understanding of the patterns and order in the diversity of marine phytoplankton species, their morphological relationships, and their global distribution. Nearly 19 years ago Dr. Faust was introduced to identifying marine plankton in a course taught by Dr. Hasle. To this day she is still fascinated by the beauty and diversity of dinoflageilate structures and morphological patterns which manage to restore one's perspective and faith in nature. IDENTIFYING HARMFUL MARINE DINOFLAGELLATES Dinoflagellates are unicellular eukaryotic microorganisms. They are free swimming protists with a forward spiraling motion propelled by two dimorphic flagella. They possess a large nucleus with condensed chromosomes, chloroplasts, mitochondria and golgi bodies. Biochemically, photosynthetic species have chlorophylls a and c, and light harvesting pigments peridinin, fucoxanthin and xanthophylls. Dinoflagellates mainly reproduce asexually via binary fission, but some species reproduce sexually and form resting cysts. Their nutrition varies from autotrophy (photosynthesis) to heterotrophy (absorption of organic matter) to mixotrophy (autotrophic cells engulf prey organisms). These features are species-specific (Spector 1984). Dinoflageilate species are adapted to a variety of habitats: from pelagic to benthic. from temperate to tropical seas, and from estuaries to freshwater. Many species are cosmopolitan and can survive in variety of habitats: in the plankton, or attached to sediments, sand, corals, or macroalgal surfaces. Some species produce resting cysts that can survive in sediments for an Harmful Marine Dinoflagellates extended period of time, and then germinate to initiate blooms {Spector 1984). Dinotlagellate 'blooms' (cell population explosions) can cause discoloration of the water (known as red tides) which can have harmful effects on the surrounding sea life and their consumers: mass mortalities in llsh, invertebrates, birds, and mammals. When toxic species are in bloom conditions the toxins can be quickly carried up the food chain and indirectly passed onto humans via tlsh and shellfish consumption, sometimes resulting in gastrointestinal disorders, permanent neurological damage, or even death. While harmful dinotlagellate blooms are at times a natural phenomenon and have been recorded throughout history, in the past two decades the public health and economic impacts of such events appear to have increased in frequency, intensity and geographic distribution (Taylor 1987). Toxin production and red tide events of harmful marine dinoflagellates are summarized in Table I. Table 1. Toxin production and red tide events of harmful marine dinoflagellates Species Keel Tide Produced loxin Reference Alexandrium acatenella A. catenella A. minutum A. moniiatum A. ostenfetdii A. pseudogonyaulax A. tamareuse A. lamiyavantchi YES YES YES PSP toxins Prakash & Taylor 1966 NO NO Iclithyotoxins; PSP toxins: CI -4, GTX, Prakash et al. 1971, Fukuyo 1985, Fukuyo SXT et al. 1985, Oeata & Kodama 1986 PSP toxins: GTX 1-4 Oshimaetal. 1989 YES Ichthyotoxins PSP toxins: GTXl, SXT Gates & Wilson 1960, Ray & Aldrich 1967, Schmidt &Loeblich 1979 Mild PSP toxins; Spirilides Goniodomin A Cembella et al. 1987, 1988,2000 Murakami et al. 1988 YES Strong PSP toxins: GTX 1-5, NSXT, SXT Larsen & Moestrup 1989. Shimizu et al. 1975, Oshimaetal. 1977 NO Strong PSP toxins: G IX, SXT Fukuyo et al. 1989, Kodama et al. 1 988 Cochiodinium polykrikoides YES Ichthyotoxins Yuki & Yoshimatsu 1989, K.im 1998, Ho & Zubkofri979 Coolia monotis NO Cooliatoxin Nakajimaetal. 1981, Holmes et al. 1995 Dmophysis acuminata YES DSP toxins: OA Cembella 1989, Lee et al. 1989, Kat 1985 D. acuta YES DSP toxins: DTX 1,0 A Lee et al. 1989, Yasumoto 1990 D. caudata YES Ichthyotoxins Okaichi 1967 D. fortil NO DSP toxins: DTX 1 -2, OA Lee et al. 1989, Yasumoto 1990 Harmful Marine Dinoflagellates Species Red Tide Produced 1 oxin Reference D. mitra NO DSP toxins: DTX1,0A Lee et al. 1989 D. norvegica D. rotundata D. saccuhts D. tripos YES NO YES NO DSP toxins: DTXl.OA DSP toxins: DTXl DSP toxins: OA DSP toxins: DTXl Cembella 1989, Lee et al 1989. Yasumoto 1990 Leeetal. 1989 Masselin et al, 1992, Giacobbe et al. 1995, Delgadoetal. 1996 Leeetal. 1989 Gambierdiscus toxicus Gonyaulax polygramma Gymnodinium breve G. catenatum NO Ciguatoxin, Gambieric acid, Maitotoxin Murataet al.! 990, Yasumoto et al. 1977, 1987, 1993, Yokoyamaetal. 1988 YES Fish and shellfish kills due to anoxia after Hallegraeff 1991, Koizumi et al. 1996 red tide YES YES NSP toxins: Brevetoxins PSP toxins Baden 1983, Baden et al. 1982. Hughes 1979 Morey-Gaines 1982, Mee et al 1986 G. mikimotoi G pulchellum G. sanguineum G. veneficum YES NSP toxin: Gymnodimine; Ichthyotoxins Hallegraeff 1991. Seki et al. 1996 Lingulodinium polyedra Noctiluca scintiUans Ostreopsis heptagona O. lenticularis YES YES NO Gyroduuum galatheanum YES YES Ichthyotoxins Ichthyotoxins Ichthyotoxins lchth>otoxins PSP toxins: SXT Onoue et al. 1985, Onoue & Nozawa 1989, Steidingeretal. 1998 Cardvvell et al. 1979, Tindal! et al. 1984. Carlson & Tindali 1985 Ballantine 1956, Abbott & Ballantine 1957, Dodge 1982 Braarud 1957, Steemann Nielsen & Aabye Jensen 1957, Pieterse & Van Der Post 1967 Bruno etal. 1990 YES Fish and shellfish kills due to high levels Okaichi & Nishio 1976 of ammonia atler red tide NO NO Unnamed toxin OTX, Unnamed toxin J. Babinchak {according to Norris et al. 1985) Tindali et al. 1990, Ballantine et al. 1988 O. mascarenensis NO Ciguatera toxin? Quod 1994, Morton, S.L. (personal communication) O. ovata NO Unnamed toxin Nakajimaetal. 1981 Harmful Marine Dinoflagellates Species Red Tide Produced Toxin Reference O. siamensis NO Unnamed toxin Nakajimaet al. 1981, Usami et al. 1995 Pfiesteha piscicida NO Ichthyotoxins Burkholder et a!. 1995, Noga et al. 1996, Burkholder& Glasgow 1997 Prorocentrum arenarium NO DSP toxins: OA Ten-Hage et al. 2000 P. balticum YES Unknown toxin Siiva 1956, 1963, Numann 1957 P. belizeaintm NO DSP toxins: DTX1,0A Morton etal. 1998 P concavum NO DSP toxins: OA. FAT, Unnamed toxin Tindall et al. 1984, Tindall et al. 1989, Dickevetal. 1990, Huetal. 1993 P. fausttae NO DSP toxins: DTX1,0A Morton 1998 P hoffmanniamim NO DSP toxins: OA, FAT Aikman et al. 1993 P. lima NO DSP toxins: DTX 1 ,2,4, OA, FAT, prorocentrolide Murakami et al. 1982, Yasumoto et al. 1987, Torigoe et al. 1988, Tindall et al. 1989, Lee etal. 1989, Marr etal. 1992, Hu etal. 1993, 1995 P. maculosiim NO DSP toxins: OA, Prorocentrolide B Dickcv et ai. 1990, Hu et al. 1996 P. mexicanum NO FAT Steidinger 1983, Carlson 1984, Tindall et al. 1984 P. inkam YES Shellfish kills Pinot & Silva 1956, Horstman 1981 P. minimum NO DSP toxin: Venerupin Nakazima 1965, 1968, Smith 1975, Okaichi & Imatomi 1979, Tangen 1983, Shiniizu 1987 P. ruetzlerianum NO Unnamed toxin Quod (personal communication) ABBREVIATIONS: DSP = diarrhetic shellfish poisoning; DTXi, DTX2, DTX3, DTX4 = dinophysistoxins; FAT = fast acting toxin; GTXl, GTX2, GTX3, GTX4, GTX5 = gonyautoxins; NSP = neurotoxic shellfish poisoning; NSXT = neosaxitoxin; OA = okadaic acid; OTX = ostreotoxin; PSP = paralytic shellfish poisoning, SXT = saxitoxin. Dinoflagellates exhibit a wide divergence in morphology and size that are essential features used to identify species, as well as surface ornamentation (pores, areolae, spines, ridges, etc.). Armored or thecate species, those that possess a multi-layered cell wall, can be Harmful Marine Dinoflagellates distinguished from unarrnored or atiiecate species, those that lack a cell wall. Surface morphology of thecate cells, often critical to proper identification, can be discerned after cell fixation. However, identification of athecate species is mainly based on live cells since many morphological features may by destroyed by fixation (Steidinger & Tangen 1996). orientation is also used: the forward end when the cell moves is called the apical pole: the opposite end is the antapical pole. Desmokonts are laterally flattened species with two large lateral plates: right valve and left valve. In lateral view the right valve reveals flagellar placement in the anterior V-shaped depression (Fig. 1 A). Dinokonts are, in general, divided into 2 main sections (epitheca and hypotheca) and divided by a girdle (cingulum) (Fig. 1 B-F). The side the tlagella arise from is the ventral side, the opposite side is the dorsal. Ventral view (Fig. 1 B) reveals the position of the flagella in relation to the cingulum and sulcus (Taylor 1987). Other important features include position of the cingulum and whether it is displaced or not (Fig. I B). If displaced and the left side is more anterior, the displacement is left-handed. If the opposite is true, it is right-handed. The former is much more common. The degree of displacement is given in cingulum widths (Taylor etal. 1995). In thecated species the plate pattern, or tabulation, is crucial (see Balech & Tangen 1985) (Fig. 1 C, E, F). The description of new species or any critical taxonomy requires complete elucidation of the plate pattern, which can be difficult, requiring special techniques (see Steidinger et al. 1996). ACKNOWLEDGEMENTS Fig. 1. Identity'ing dinotlagellates: A. lateral \ic\\ of a desmokont cell type (two dissimilar tlagella apically inserted); B. ventral view of a dinokont cell type (two dissimilar tlagella vctitrally inserted); C. ventral view of a thecate peridinioid cell; D. ventral view of an athecate gymnodinoid cell; E. apical view of cpithocal plates; F. antapical view of hypothecal plalcs. Ch = chloroplasts; N = nucleus; Po = apical pore plate; SL = sulcal list (Figs. A-B redrawn from Steidinger & Tangen 1996; Figs, C-F redrawn from Taylor 1987) Another distinction used in dinoflagellate identification is morphological cell type (Fig. 1 A, B): 1. desmokont type where two dissimilar flagella are inserted apically (e.g. Prorocentrum)\ and 2. dinokont type where two dissimilar flagella are inserted ventrally (e.g. Alexandrium). Terminology to describe Dr. Maria Faust thanks Dr. Klaus Ruetzler, Curator of Sponges, National Museum of Natural History, Smithsonian Institution, for introducing her to the magnificent world of coral reef-mangrove ecosystems at Belize and encouraging her studies. We are greatly indebted to Drs. Patricia A. Tester (National Ocean Service, NOAA) and Steve L. Morton (Marine Biotoxin Program, NOAA) for contributing photomicrographs and critically reviewing the manuscript. We thank S.H. Brawiey, editor of Journal of Phycology, for permission to use published pictures (University of Maine), and D.G, Mann, editor of Phycologia, for permission to use published pictures (Royal Botanic Garden Edinburgh). We also thank the following scientists and colleagues 10 Harmful Marine Dinoflagellates for providing photomicrographs of harmful dinoHageliate species: Drs. C. Andreis (University of Milan), G.T. Boalch (The Laboratory-Citadel Hill), S. Blackburn (CSIRO Marine Research), J.M. Burckholder (North Carolina State University), B. Dale (Universiry of Oslo), J.D. Dodge (Royal Holloway College), Y. Fukuyo (University Tokyo), D. Grzebyk (CREMA-L Houmeau, CNRS-IFREMER). G. Hallegraeff (University of Tasmania), G. Honsell (University of Udine), T. Horiguchi (Hokkaido University), J. Larsen (University of Copenhagen), J. Lewis (University of Westminster), A.J. Lewitus (University of South Carolina), L. Mackenzie (Cawthron Institute), K. Matsuoka (Nagasaki University), M. Montresor ('A. Dohrn' Zoological Station), T. Nishijima (Kochi University), D.R. Norris (Florida Institute of Technology), A. Prakash (Bedford Institute of Oceanography), K.A. Steidinger (Florida Marine Research Institute), H. Takayama (Hiroshima Fisheries Experiment Station), F.J.R. Taylor (University of British Columbia), S. Toriumi (Higashi Senior High School), K. Yuki (Matoya Oyster Research Laboratory) and A. Zingone ('A. Dohrn' Zoological Station). We also thank Don Hulbert for technical help (Smithsonian Office of Imaging, Printing & Photographic Services). Finally, we wish to express our appreciation to Dr. P.M. Peterson, editor, for his useful suggestions to improve the clarity of presentation of this work. Identifying Harmful Marine Dinoflagellate Species Alexandrium acatenella (Whedon el Kofoid) Balech, 1985 Plate l,Figs. 1-4 Species Overview: Alexandrium acatenella is an armoured, marine, pianktonic dinoflagellate. It is associated with toxic PSP blooms in Pacific coastal regions. Taxonoinical Description: A non-chain forming species, cells of A. acatenella are small to medium sized, longer than wide, and angular to round in ventral outline (Figs. 1,2). A characteristic ventral pore is present (Fig. 3). Two short antapical spines are present; no apical horn (Fig. 3). The thecal surface is sculptured with large and small pores. Cells range in size between 35-51 (im in length and 26-35 ^m in transdiameter width (Whedon & Kofoid 1936; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). INoinciiclatura! Types: HolotNpe: Gonyuulux acatenella Whedon and Kofoid, 1936; 31, 33-34, figs. 8-13 Type Locality; NW Pacific Ocean; San Diego, California, USA Synonyms; Gonyaulax acatenella Whedon and Kofoid, 1936 Protogonyaulax acatenella (Whedon and Kofoid) Taylor, 1979 Thecal Plate Description: The plate formula for A. acatenella is; Po, 4', 6", 6c, 9s, 5'", T". The epitheca in this species is longer than the hypotheca; often it is equal to the length of the hypotheca plus the cingulum. The cone-shaped epitheca is low with convex sides (Figs. 1-3). The apical pore complex (APC) is roughly rectangular. The apical pore plate (Po) is broadiy oval and narrows ventrally; it bears a relatively large and comma-shaped foramen (Fig. 4). The first apical plate (1') comes in direct contact with the Po, and also bears the characteristic ventral pore (vp) (Fig. 4)(Whedon & Kofoid 1936; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). The post-median cingulum is deeply excavated, and displaced in a descending fashion about 1 time its width without overhanging. Narrow lists are present on the cingulum (Figs. 1- 3). The deeply excavated sulcus widens Harmful Marine Dinoflagellates 11 posteriorly flaring to the right, slightly invading the hypotheca. The short hypotheca is broadly rounded with two posterior antapical spines (Figs. 1-3). The antapex region between the spines is slightly concave (Whedon & Kofoid 1936; Balech 1995; Taylor et a!. 1995; Steidinger & Tangen 1996). [Morphology and Structure: A. acatenella is a photosynthelic species with elongated chloroplasts. Cells can be highly pigmented and reddish-brown in color. The elliptical nucleus is C-shaped and equatorial (Whedon & Kofoid 1936; Prakash & Taylor 1966; Balech 1995). Reproduction: A. acatenella reproduces asexually by binary tlssion (Whedon & Kofoid 1936). Ecology: A. acatenella is a plankton ic species associated with paralytic shelltTsh poisoning (PSP) events and red tides. Populations are most abundant in neritic waters at 15T. A bloom event in British Columbia caused four human illnesses and one death in 1965, the first reported PSP outbreak associated with A. acatenella. Cell densities during this red tide were as high as 13.5 X 10^ cells/L (Whedon & Kofoid 1936; Prakash & Taylor 1966). Toxicity: Alexandrium acatenella is a known PSP toxin-producing dinoflagellate species responsible for several illnesses and one death in British Columbia (Prakash & Taylor 1966). Species Comparison: A. acatenella is very similar morphologically (size, shape and thecal plate formula) to the toxic Atlantic species, A. tamarense. Differences lie in the general shape of the cell, thecal sculpture, length of epitheca in relation to the hypotheca, and size and shape of the apical plates. The former species is roundish, while the latter is wider (shoulders) and roughly pentagonal. Thecal plates in A. acatenella are clearly porolated, while in A. tamarense they are relatively smooth. The epitheca in A. acatenella is distinctly longer than the hypotheca; they are nearly equal in A. tamarense. The size and shape of the apical plates differ in these two species (Balech 1995). A. acatenella also shares some common characteristics of A. catenella. However, the former species is a non-chain former without a posterior attachment pore, bears a ventral pore on r, and is usually found in warmer waters (Prakash & Taylor 1966; Balech 1995). Habitat and Locality: Alexandrium acatenella is widely distributed in Pacific coastal waters. Populations have been recorded from the north Pacific coast of the United States and Canada, Japan, Argentina and northern Chile (Whedon & Kofoid 1936; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). Alexandriiim caten ella (Whedon et Kofoid) Balech, 1985 Plate 2, Figs. 1-6 Species Overview: Alexandrium catenella is an armoured, marine, planktonic dinoflagellate. It is associated with toxic PSP blooms in cold water coastal regions. Taxonomical Description: A chain-forming species, A. catenella typically occurs in characteristic short chains of 2, 4 or 8 cells (Figs. 1,2). Single cells are round, slightly wider than long, and are anterio-posteriorly compressed. A small to medium sized species, it has a rounded apex and a slightly concave antapex (Fig. 1). The thecal plates are thin (Fig. 3) and sparsely porulated. Cells range in size between 20-48 |im in length and 18-32 ^m in width (Fukuyo 1985; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gonyaulax catenella Whedon and Kofoid, 1936: 25-31, figs. 1-7, 14.15 Type Locality: NW Pacific Ocean: San Diego, California, USA Synonyms: Gonyaulax catenella Whedon and Kofoid, 1936 Protogonyaulax catenella (Whedon and Kofoid) Taylor, 1979 Thecal Plate Description: The plate formula for A. catenella is: Po, 4', 6", 6c, 8s, 5"', 2"". The epitheca and hypotheca are nearly equal in height. The hypotheca bears prominent sulcal 12 Harmful Marine Dinoflagellates lists that resemble spines (Fig. 1). In chain forms, anterior attachment pores (aap) and posterior attachment pores (pap) are present (Fig. 4)(Fukuyo 1985; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). The apical pore complex (APC) is broad, triangular and widens dorsally (Figs. 3,4). The apical pore plate (Po) houses the characteristic fishhook shaped foramen, and, if catenate, an ellipsoidal aap (Fig. 4). There are two diagnostic features of this species: a.) the First apical plate, r, comes in contact with the Po (Fig. 3); and b.) a ventral pore (vp) is absent. The median cingulum is lipped, deeply concave, and is displaced in a descending fashion one time its width (Figs. 1,5). The sulcus, with prominent lists, is deeply impressed and widens posteriorly (Figs. 1,5). The wide posterior sulcal plate houses the pap near the right margin (Fukuyo 1985; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Morphology and Structure: A. catenella is a photosynthetic species with numerous yeilow- green to orange-brown chloroplasts. The nucleus is large and U-shaped (Whedon & Kofoid 1936). Reproduction: A. catenella reproduces asexually by binary fission. This species also has a sexual cycle with opposite mating types (heterothallism). After gamete fusion, a planozygote forms which then encysts into a characteristic resting cyst (Fig. 6)(Yoshimatsu 1981). Ecology: A. catenella is a planktonic dinotlagellate species associated with deadly paralytic shellfish poisoning (PSP) events mostly in the Pacific Ocean. Red tides of this species have also been observed (Fukuyo 1985). This species produces a colorless resting cyst as part of its life cycle which cannot be distinguished from the cyst produced by A. tamarense (Fig. 6). The cyst is roughly ellipsoidal with rounded ends; it is covered by a smooth wall and a mucilaginous substance. Cysts have a wide size range: 38-56 jim in length to 23-32 fim in width (Fukuyo 1985; Hallegraeff 1991; Meksumpunetal. 1994). Toxicity: Alexandrium catenella is a known toxin-producing dinoflagellate species; it is the first species ever linked to PSP (Fukuyo 1985; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995). A. catenella produces strong PSP toxins which are transmitted via tainted shellfish. These toxins can affect humans, other mammals, fish and birds: c]-c4 toxins, saxitoxins (SXT) and gonyautoxins (GTX)(Schantz et al. 1966; Prakash et al. 1971). Moreover, Ogata and Kodama (1986) report production of ichthyotoxins in cultured media of .4. catenella. This species is responsible for numerous human illnesses and several deaths after consumption of infected shellfish. Toxic blooms and PSP in shellfish have been reported in Chile (Avaria 1979), Japan (Onoue et al. 1980; 1981a; 1 98 1 b), California (Sharpe 1981) and most of the Pacific coast of the U.S.A. (Nishitani & Chew 1988). Species Comparison: A. catenella is very similar morphologically (size, shape and thecal plate formula) to A. tamarense. Differences lie in the shape of the Po, and presence or absence of a vp. The Po in the former species is slightly smaller, and the vp is absent (Fukuyo 1985). Molecular testing conducted on A. catenella from Japan and A. tamarense from Japan and the U.S.A. revealed a close genetic relationship between the two species, however they remain distinct (Adachietal. 1995). Chains of this species are quite distinctive, but can resemble A. tamiyavanichi; however, A. tamiyavanichi is a warm water species and can be distinguished from A. catenella by its conical shape (Taylor et al. 1995). Habitat and Locality: Alexandrium catenella is widely distributed in cold temperate coastal waters. Populations have been recorded from the west coast of North America (from California to Alaska), Chile, Argentina, western South Africa, Japan, Australia and Tasmania (Fukuyo 1985 Fukuyo et al. 1990; Hallegraeff 1991 Hallegraeff et al. 1991; Taylor et al. 1995 Steidinger & Tangen 1996), Harmful Marine Dinotlagellates 13 Alexandriiim minuturn Halim, 1960 Plate 3, Figs. 1-6 Species Overview: Alexandrium minutum is an armoured, marine, pianktonic dinoflagellate. It is a widely distributed species associated with toxic PSP blooms in coastal regions. Taxononiical Description: Cells of A. minutum are small, nearly spherical to ellipsoidal, somewhat dorsoventrally flattened and occassionally longer than wide (Figs. 1,2). Cells are single with a characteristic ventral pore on the first apical plate, 1' (Figs. 1-4). Thecal plates thin. Thecal surface ornamenation can vary from light to heavy reticulation (mostly confined to the hypotheca) with small scattered pores. Intercalary bands are present (Figs. 1-3). Large range in size in this species: between 15-30 \iu\ in length and 13-24 fim in transdiameter width (Balech 1989; 1995; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996; Hwang et al. 1999). Nomenclatural Types: Holotype: Alexandrium minutum Halim, I960: 101, figs, la-g Type Locality: Mediterranean Sea: Alexandria Harbor, Egypt Synonyms: Alexandrium ibericum Balech, 1985b Alexandrium lusitanicum Balech, 1985b Thecal Plate Description: The plate formula for A. minutum is: Po, 4', 6", 6c, 10s, 5'", 2"". The epitheca is larger than the hypotheca (Figs. 1,2). The apical pore complex (APC) is oval to broadly triangular and pointed posteriorly (Fig. 3). The apical pore plate (Po) is large, narrow and oval with a wide comma-shaped foramen (Figs. 3,5). The Po can be either in direct contact with the first apical plate (!') (Figs. 3,5a) or indirectly connected via a thin suture (thread- like process)(Fig. 5b). A characteristic ventral pore is located on the slender and rhomboidal l' plate (Figs. 2-4). The distinctive sixth precingular plate (6") is long and narrow (Fig. l)(Baiech 1989; 1995; Hallegraeff 1991; Taylor et at. 1995; Steidinger & Tangen 1996; Hwang et a!. 1999). The epitheca is hemielliptical to conical with convex sides (Figs. 1,2). The apex is broadly rounded. The short hypotheca is hemielliptical with a convex to fiat antapex (Figs. 1.2). The deeply excavated cingulum is displaced in a descending fashion one time its width with thickened margins (Figs. 1,2). The sulcus is shallow with narrow lists (Figs. L2)(Balech 1989; 1995; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996; Hwang et al. 1999). Morphology and Structure: A. minutum is a photosynthetic species with an elliptical nucleus (Balech 1989; 1995). Reproduction: A. minutum reproduces asexually by binar> fission. This species also has a sexual cycle that produces a characteristic resting cyst (Fig. 6)(Bolchetal. 1991). Ecology: A. minutum is a pianktonic dinoflagellate species associated with toxic paralytic shellfish poisoning (PSP) events in coastal regions around the world. This species also produces dense (reddish-brown) red tides (Hallegraeff 1991). A red tide of this species reported from Taiwan had cell densities as high as 2.5 X lO' cells/L (Hwang et al. 1999). Another red tide of A. minutum reported from South Australia revealed cell levels of 4.8 X 10' cells/L (Cannon 1990). This species produces a clear resting cyst as part of its life cycle. Cysts vary from hemispherical to circular in shape: cyst circular in apical view (24-29 fim in diameter) (Fig. 6); kidney-shaped in lateral view (15-19 |am long). The cyst wall is covered with mucilage (Bolch et at. 1991). Toxicity: Alexandrium minutum is a strong producer of PSP gonyautoxins (GTX): GTXl, GTX2, GTX3 and GTX4 (Oshima et al. 1989). These toxins can affect humans, other mammals, birds and possibly fish (Hallegraeff et al. 1988; Hallegraeff 1991). This species is responsible for PSP events in Taiwan (Hwang et al. 1999), South Australia (Hallegraeff et al. 1988, Cannon 1990), France (Nezan et al. 1989) and New Zealand (Chang et at. 1995). 14 Harmful Marine Dinoflagellates Habitat and Locality: Alexandrium minutum is widely distributed species found in many coastal areas of the world. Populations have been recorded from Alexandria Harbor, Egypt (Halim I960), Italy (Montresor et al. 1990), northern Adriatic waters (Mediterranean Sea)(Honsell 1993), Turkey (Koray & Buyukisik 1988), Spain and Portugal (as A. /^fr/'a' in the life history of the species; e.g. the development of the megacytic zone. This area can increase before cell division or following gamete fusion (Giacobbe & Gangemi 1997). Biological factors (i.e. life history and nutrition) can explain the presence of different morphotypes in the same locality (Zingone et al. 1998). Reproduction: D. saccidm reproduces asexually by binary fission (Taylor et al., 1995). Giacobbe and Gangemi (1997) reported sexual reproduction in this species. Ecology: Dinophysis sacculus is a planktonic species (Taylor et al. 1995). Blooms have been reported from Portugal, North Atlantic Ocean (Alvito et al. 1990; Sampayo et al, 1990), and Italy, Mediterranean Sea (Zingone et al. 1998). Toxicity: D. sacculus has been found to produce okadaic acid (OA) (Masselin et ai. 1992; Giacobbe et al. 1995; Delgado et al. 1996). It has been linked to diarrhetic shellfish poisoning (DSP) occurrences along the Mediterranean and Atlantic European coasts (Alvito et al. 1990; Sampayo et al. 1990; Lassus & Marcaillou-Le Baut 1991; Belin 1993; Boni et al. 1993; Marasovic et al. 1998). Species Comparison: D. sacculus is most often misidentilled as D. acuminata. The major difference between these two species is the shape of the large hypothecal plates: in D. sacculus they are long and sack-like, whereas in D. acuminata they are shorter, more convex dorsal ly and often more slender posteriorly. D. acuminata also exhibits more pronounced thecal areolation and sulcal list ornamentation, but these are variable characteristics. Moreover, since D. sacculus and D. acuminata rarely occur in the same area with the same importance, the possibility of misidentification is reduced (Zingone et al. 1998). Surface thecal ornamentation in this species is similar to a number of other Dinophysis species: D. acuta, D. cauclata, D. norvegica and D. fortii (HallegraetTiS: Lucas 1988). Etymology: "Sacculus ' (Latin) refers to the sack- like shape of the hypotheca. Habitat and Locality: D. sacculus is distributed widely in cold and temperate waters (Taylor et al. 1995), most often observed in semi-enclosed basins, estuaries and lagoons (Zingone et al. 1998). Populations have mostly been reported from the Mediterranean Sea (Zingone et al. 1998), with a few reports from the Atlantic Ocean (Murray & Whitting 1900; Cleve 1901; 1902). F'ieiiiarks: D. sacculus has a history wrought with identitlcation problems mainly attributable to the morphological variability of this species. This problem is enhanced by the many synonyms and questionable identifications that have accumulated in the literature over the years (see Zingone etal. 1998). Many authors consider Phalacroma to be synonymous with Dinophysis (Steidinger & Tangen 1996). Dinophysis tripos Gourret, 1883 Plate 19, Figs. 1-4 Species Overview: Dinophysis tripos is an armoured, marine, planktonic dinoflagellate species. It is a toxic species common in warm temperate to tropical waters. Taxonomic Description: Species in this genus are laterally compressed with a small, cap-like epitheca and a much larger hypotheca (dorso- ventral depth of epitheca is 1/3 to 1/2 hypotheca). The shape of the cell in lateral view is the most important criterion used for identification (Taylor et al. 1995). D. tripos is a very distinctive species. Cells are large, anterio-posteriorly elongated and asymmetrical with two posterior hypothecal projections; a longer ventral process and a shorter dorsal one (Figs. 1-4). The V-shaped processes are often toothed on their posterior ends (small knob-like spines) (Fig. 1). The well developed left sucal list (LSL) widens posteriorly and is often reticulated (Figs. ]-3)(Larsen & Moestrup 1992; Taylor et al. 1995; Steidinger & Tangen 1996). Harmful Marine Dinoflagellates 35 The thick thecal plates are heavily areolated (Fig. 1). Cell size ranges: 90-125 pm in length and 50-60 ^m In dorso-ventral width (Larsen & Moestrup 1992; Taylor et al. 1995). Nomenclatural Types: Holotype: Dinophysis tripos Gourret, 1883: 114, plate 3, fig. 53 Type Locality: Mediterranean Sea: Gulf of Marseille, France Synonyms: Dinophysis caudata var. tripos (Gourret) Gail, 1950 Thecal Plate Description: The small epitheca is made up of four plates. The cingulum is narrow with two well developed lists, anterior cingular list (ACL) and posterior cingular list (PCL), oriented anteriorly (Figs. 1-4). The ACL is supported by many ribs (Figs. 1,4). The wide ACL forms a narrow, funnel-like structure obscuring the epitheca on the bottom. The sulcus is comprised of several irregularly shaped plates. The flagellar pore is housed in the sulcal area. The prominent wide LSL has a straight margin and is supported by three ribs (Figs. l-4)(Larsen & Moestrup 1992; Taylor et al. 1995; Steidinger &Tangen 1996). The hypotheca, with four large plates, comprises the majority of the cell. It is long, narrowing into two tapered or pointed posterior projections: one short and dorsal, and one longer and ventral (Figs. 1-3). The dorsal projection is sometimes seen with a narrow list connecting two daughter cells during cell division (Fig. 3). The ventral margin of the hypotheca is straight or slightly undulate. The dorsal margin is concave below the cingulum and then convex continuing down to the dorsal projection (Figs. l,2)(Larsen & Moestrup 1992; Taylor et al. 1995; Steidinger &Tangen 1996). Morphology and Structure: D. tripos is a photosynthetic species with chloroplasts (Fig. 2). D. diegensis, a smaller form very similar in morphology to D. tripos with a reduced hypothecal process, is suspected to be a gamete of the latter species (Moita & Sampayo 1993). Reproduction: D. tripos reproduces asexually by binary Fission. Moita and Sampayo (1993) speculate that sexual reproduction, with sexual dimorphism, is part of the life cycle for this species. Ecology: Dinophysis tripos is a planktonic species commonly found in neritic, estuarine and oceanic waters (Steidinger & Tangen 1996). No blooms for this species have been reported (Larsen & Moestrup 1992). Toxicity: D. tripos is associated with diarrhetic shellfish poisoning (DSP) events; it produces Dinophysistoxin-1 (DTXl)(Lee et al. 1989). Species Comparison: Dinophysis tripos can be confused with D. caudata; some cells of D. caudata, bearing a short hypothecal process, can superficially resemble D. tripos. However, D. tripos can be distinguished by the presence of two posterior projections (Larsen & Moestrup 1992; Steidinger & Tangen 1996). Habitat and Locality: D. tripos is widely distributed in tropical and temperate waters, and occasionally is found in colder regions (Larsen & Moestrup 1992; Taylor et al. 1995; Steidinger & Tangen 1996). Remarks: Many authors consider Phalacroma to be synonymous with Dinophysis (Steidinger &. Tangen 1996). Giunhierdiscus toxiciis Adachi and Fukuyo. 1979 Plate 20, Figs. 1-6 Species 0\erview: Gambierdiscus toxicus is an armoured, marine, benthic dinoflagellate species. It is a toxic species that was discovered attached to the surface of brown macroalgae in the Gambler Islands. French Polynesia. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The epitheca and hypotheca are not noticeably different in size. A distinguishing feature is the shape and size of the apical pore complex (APC)(Fig. l)(Faust 1992). Cells of Gambierdiscus toxicus are large, round to ellipsoid (Figs. 1,2,4,5), and flattened anterio-posteriorly. The epitheca and hypotheca 36 Harmful Marine Dinoflagellates are nearly equal in height. The cell surface is smooth with numerous deep and dense pores (Figs. 1,3). Thecal plates are very thick. Cells range in size from 24-60 jim in length, 42-140 ^m in transdiameter, and 45-150 jam in dorso- ventral depth (Adachi & Fukuyo 1979). Nomenclaturai Types: Holotype: Gambierdlscus toxicus Adachi and Fukuyo, 1979: figs. 1-7 Type Locality: South Pacific Ocean: Gambler Islands, French Polynesia Synonyms: Diplopsalis sp. Yasumoto et al., 1977 Thecal Plate Description: The plate formula of Gamblerdiscus toxicus is: Po, 3', 7", 6c, 8s, 5"', Ip, 2""(Faust 1995). The apical pore plate (Po) is oval to ellipsoidal with a characteristic fishhook shaped apical pore (Figs. 1,3), the opening of which is always oriented ventrally. Apical plate 2' is subrectangular and is the largest of the three apical plates (Figs. 1,6) (Adachi & Fukuyo 1979). The epitheca is slightly indented ventrally (Figs. 1,4). The hypotheca is deeply excavated ventrally (Figs. 2,5,6) (Adachi & Fukuyo 1979; Fukuyo 1981; Taylor 1979). In the hypotheca the postcingular plate 1'" is triangular; its right corner extrudes, curves inside, and contacts antapical plate 1"" (Figs. 2,6)(Adachi & Fukuyo 1979; Fukuyo 1981). The posterior intercalary plate (Ip) is broad and pentagonal (Figs. 2,6). When the marginal zone widens during thecal growth, the Ip plate changes its shape to rhomboid (Fukuyo 1981). The cingulum is circular, narrow and deeply excavated, and ascends slightly (Adachi & Fukuyo 1979; Bagnis et al. 1979; Taylor 1979). The cingular wall consists of six plates and measures nearly 5 jim in width. It is bordered by a low, thick ridge which is made up of the folding of pre- and postcingular plates (Figs. l,4)(Adachi & Fukuyo 1979)." The sulcus is short, deeply concave and pouch-like, and is oriented to the right (Figs. 2,5)(Adachi & Fukuyo 1979; Bagnis et al. 1979; Taylor 1979), Along the sulcal margin, an overhanging ridge continues along the edge of postcingular plate 5'", and antapical plates I"" and 2""(Fig. 2) (Adachi & Fukuyo 1979). Morphology and Structure: G. toxicus is a photosynthetic species with yellow to golden- brown chloroplasts and a large crescent-shaped nucleus (Fig. 5)(Adachi & Fukuyo 1979). Reproduction: G. toxicus reproduces asexually by binary fission. Ecology: Cells of G. toxicus are frequently found as epiphytes on macroalgae and dead coral. Different strains apparently exhibit a preference for certain algae; e.g. the Hawaiian strain prefers the red alga Spyridia jllamentosa (Shimizu et al. 1982). Cells readily attach to substrates via mucoid strands originating from the sulcal area (Steidinger & Tangen 1996). Toxicity: G. toxicus is known to produce the following toxins: ciguatoxin (Yasumoto et al. 1987; Murata et al. 1990; Yasumoto et al. 1993); gambieric acid (Yasumoto et al. 1993); and maitotoxin (Yasumoto et al. 1977; 1993; Yokoyama et al. 1988). Species Comparison: This species resembles Heteraidacus in tabulation, but differs by its right-handed girdle torsion, large apical closing plate, and a pouch-like sulcal depression (Taylor 1979). Gamhierdiscus toxicus shares a number of characteristics with G. belizeanus. They both have the same plate formula, and have similar apical pore, cingulum, sulcus, general cell shape (lenticulate and antero-posteriorly compressed), and golden brown chloroplasts. However, they differ in a number of distinct features. Architecturally, both species have similar epithecal plates, but differ in thecal surface morphology: G. toxicus has a smooth surface with scattered tine pores, whereas G. belizeanus has a deeply areolated surface. G. toxicus is considerably larger than G. belizeanus. And plate 1 p is broad in G. toxicus, whereas it is long and narrow in G. belizeanus (Faust 1995). Etymology: The genus 'Ganibierdiscus' was named after the Gambier Islands from which it was discovered and also the discoid shape of the cell. The species name 'toxicus ' is derived from the toxin-producing nature of this species. Harmful Marine Dinoflagellates 37 Habitat and Locality: This species was identified from tropical reefs in the Pacific Ocean (Adachi & Fukuyo 1979; Fukuyo 1981), the Indian Ocean (Quod 1994), and the U.S. Virgin Islands (Carlson & Tindall 1985). Populations have been found in tidal pools and lagoons, as well as in colored sand, in the Caribbean (Faust 1995). In the United States, G. toxicus has been collected in waters around Hawaii (Taylor 1979; Shimizu et al. 1982) and the Florida Keys (Berginann & Alam 1981; Besada et al. 1982; Loeblich& Indelicato 1986). Gonyaulax polygramma Stein, 1883 Plate 21, Figs. 1-6 Species Overview: Gonyaulax polygramma is an armoured, marine planktonic dinoflagellate species. It is a red tide bloom species associated with massive fish and shellfish kills. Taxonomic Description: Cells of Gonyaulax polygramma are medium-sized, elongate and pentagonal (Figs. 1-6). The tapered epitheca bears a prominent apical horn, and exceeds the symmetrical hypotheca (Figs. 1-3). Longitudinal ridges ornament the thecal surface; reticulations are present between the ridges (Figs. 1-3). On mature cells, longitudinal ridges may be thick and spinulous. Cells range in size from 29-66 l^m in length and 26-56 jam in dorso-ventral depth (Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gonyaulax polygramma Stein, 1883: pi. 4, figs. 15-16 Type Locality: unknown Thecal Plate Description: The plate formula for G. polygramma is: Po, 3', 2a, 6", 6c, 4-8s, 6'", 1 ""(Dodge 1989). The epitheca is convex to angular, and bears 12 apical plates (Figs. 1-3). The elliptical apical pore plate (Po) does not extend onto the dorsal side of the cell; it is in direct contact with the first apical plate (T). The 1 ' plate with a ventral pore (vp). The left-handed cingulum is post-median and displaced about 1.5 times its width without overhanging (Figs. 1,2,4,6). The slightly excavated sulcus widens posteriorly; it invades the epitheca slightly (Figs. 1,6). The hypotheca is truncate with straight sides and consists of six plates; 1-3 antapical spines present (Figs. l-4,6)(Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Steidinger & Tangen 1996). Morphology and Structure: G. polygramma is a photosynthetic species with chloroplasts. The large oval nucleus is located posteriorly (Dodge 1982). Reproduction: G. polygramma reproduces asexually by binary fission. Ecology: G. polygramma is a planktonic species commonly found in neritic and oceanic waters (Steidinger & Tangen 1996). This cosmopolitan species is a red tide bloom former associated with shellfish and fish kills. Deadly G. polygramma red tides have been reported from Florida (Steidinger 1968), Japan (Nishikawa 1901; Fukuyo et al. 1990; Koizumi et al. 1996), New South Wales (Hallegraeff 1991), South Africa (Grindley & Taylor 1964), and Hong Kong (Lam & Yip 1990). During a bloom in Uwajima Bay, Japan, in 1994, cell levels peaked at 6.8 X 10"* cell/ml and caused mass mortalities of cultured and natural fish and shellfish stocks (Koizumi et al. 1996). Toxicity: G. polygramma is a non-toxin producing species, but as a red tide species, it has been associated with massive fish and invertebrate kills due to anoxia and high sulfide and ammonia levels resulting from celt decomposition (Hallegraeff 1991; Koizumi et al. 1996). Habitat and Locality: G. polygramma is a cosmopolitan species common in cold temperate to tropical waters worldwide (Hallegraeff 1991; Steidinger & Tangen 1996). Gymnodinium breve Davis, 1948 Plate 22, Figs. 1-4 Species Overview: Gymnodinium breve is an unarmoured, marine, planktonic dinofiagellate species. It is a toxin-producing species 38 Harmful Marine Dinofiagellates associated with red tides in the Gulf of Mexico, off the coast of western Florida. Taxonomic Description: Gymnodiniiim breve is an athecate species; i.e. without thecal plates. Cells are small and dorso-ventrally flattened (Figs. 1-3). The cell is ventrally concave and dorsal ly convex. Cells appear almost square in outline, but with a prominent apical process directed ventrally (Figs. 1,3,4). Cells range in size from 20-40 ^im in width to 10-15 jam in depth, and are slightly wider than long (Steidinger et al. 1978; Steidinger 1983; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca is rounded with a distinctive overhanging apical process (Figs. 1-3). The epitheca is smaller than the hypotheca (Figs. 1- 3). The cingulum is displaced in a descending fashion up to 2 times its width. It houses the transverse tlagellum. The sulcus extends into the epitheca up to the antapex adjacent to the apical process (Fig. 4). It houses the longitudinal tlagellum. An apical groove, present near the distal epithecal end of the sulcus, extends across the apical process onto the dorsal side of the cell (Figs. 1,2). It is not an extension of the sulcus. The wide hypotheca is notched by the sulcus and is slightly bilobed posteriorly (Figs. 1-4). Discharged trichocysts have been observed (Davis 1948; Steidinger et al. 1978; Steidinger 1983; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype; Gymnodinium breve Davis, 1948: 358-360, figs. 1,2 Type Locality: Gulf of Mexico: near Naples. Florida, USA Synonyms: Ptychodiscus brevis (Davis) Steidinger, 1979 Morphology and Structure: Gymnodinium breve is a photosynthetic species with numerous peripheral yellowish-green chloroplasts and nuiliistalked pyrenoids (Figs. 2,3). The large round nucleus is 6-9 jam in diameter and located in the left half of the hypotheca (Figs. 3,4). Lipid globules have also been observed (Fig. 3). This species does not have peridinin as a major accessory pigment (Davis 1948; Steidinger et al. 1978; Steidinger 1983; Taylor et al. 1995; Steidinger & Tangen 1996), Reproduction: G. ^reve reproduces asexually by binary Fission; cells divide obliquely during mitosis. This species also has a sexual cycle: isogamous gamete production, fusion and formation of a planozygote. The planozygote is morphologically similar to the vegetative cell, but larger. The gametes are rounder and slightly smaller than the vegetative cells (18-24 )im in diameter). It is speculated that temperature controls the onset of the sexual cycle since sexual stages only occurred in fall and winter in both field populations and cultures (Walker 1982). Ecology: G. breve is a planktonic oceanic species, though populations have been documented in estuarine systems under bloom conditions. This species is a bloom-former associated with red tides in the Gulf of Mexico, in particular the west coast of Florida. During a bloom cell levels can reach as high as I X 10^ to I X 10^ celis/L. Blooms initiate offshore requiring high salinities (> 30 o/oo) and high temperatures (Steidinger 1975; Steidinger et al. 1978; Steidinger & Tangen 1996). G. breve cells are active swimmers resembling 'falling leaves as they swim slowly, turning over and over through the water'. This species forms cysts under adverse conditions. Chain formation reported in very dense concentrations (Steidinger & Joyce 1973). Toxicity: G. breve is a known toxic species that produces a series of brevetoxins (neurotoxins) (Baden 1983). These toxins are responsible for massive till kills along the west coast of Florida in the Gulf of Mexico. Aerosolization of the toxins (noxious air-borne G. breve fragments from sea spray) has been linked to asthma-like symptoms in humans (Baden et al. 1982). Brevetoxins produce neurotoxic shellfish poisoning (NSP) when consumed (Hughes 1979). These toxins are known to cause human illness and distress; however, the poison is not fatal: no human fatalities have been reported from consumption of G. /^/-fvf-infected bivalves (Steidinger & Joyce 1973). So far NSP has been restricted to the western coast of Florida, but more recently it has been documented for New Zealand as well (Steidinger et a!. 1973; Baden et al. 1982; Taylor etal. 1995). Harmful Marine Dinofla.aellates 39 Habitat and Locality: Gymnodinium breve populations are found in warm temperate to tropical waters, most regularly from the Gulf of Mexico, off the west coast of Florida. G. breve and G. hreve-WkQ species have also been reported from the West Atlantic, Spain. Greece, Japan and New Zealand (Fukuyo et al. 1990; Taylor et al. 1995; Steidinger & Tangen 1996). Gymnodhiiuni eatenatiim Graham, 1943 Plate 23, Figs. 1-7 Species Overview: Gymnodinium catenatum is an unarmoured, marine, planktonic dinoflagellate species. It is a chain-forming, toxin-producing, red tide species associated with PSP events throughout the world. Taxonomic Description: Gymnodinium calenalum is an athecate species; i.e. without thecal plates. This species is typically seen in chain formation with up to 64 cells. Cells are small with morphology varying between single cell (Fig. 1) and chain formation (Figs. 2-4). Single cells are generally elongate-ovoid with slight dorso-ventral compression (Figs. 1,5). The apex is truncate or slightly conical while the antapex is rounded and notched (Figs. 1,5). Chain formers, in general, are squarish-ovoid with anterior-posterior compression (Fig. 3). A characteristic horseshoe shaped apical groove encircles the apex (Fig. l)(Graham 1943; Larsen & Moestrup 1989; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Single cells range in size from 27-43 |im in width to 34-65 ^im in length. Chain-forming cells are slightly smaller with sizes ranging from 27-43 |im in width to 23-60 [im in length; terminal cells are slightly larger (Figs. 2,3), similar to single cells (Graham 1943; Blackburn et al. 1989; Larsen & Moestrup 1989; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca is smaller than the hypotheca, rounded or truncate (Figs. 1,2). In chain- formers, the epitheca is conical (Figs. 2,4). The larger hypotheca tapers slightly posteriorly (Figs. 2,3). It is notched by the sulcus at the antapex creating a bilobed posterior (Fig. 5). The premedian cingulum displays left-handed displacement, about 2 times its width (Figs. 1,2). The transverse flagellum is housed in the deep cingulum (Figs. 1-3). The sulcus is deep and extends almost the full length of the cell: from just beneath the apex to the antapex (Figs. 1- 3)(Graham 1943; Larsen & Moestrup 1989; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gymnodinium catenaium Graham, 1943:259-262, tigs. 1,2 Type Locality: NE Pacific Ocean: Gulf of California, Mexico Morphology and Structure: Gymnodinium catenatum is a photosynthetic species with numerous yellow-brown chloroplasts and conspicuous pyrenoids. The large nucleus is centrally located. Lipid globules are also common (Graham 1943; Larsen & Moestrup 1989; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Reproduction: G. catenatum icproduces asexually by binary tlssion. This species also has a sexual cycle with opposite mating types (heterothallism). After gamete fusion, a planozygote forms, and after two weeks, this form encysts into a characteristic resting cyst (Fig. 6). Nutrient detlciency induces the sexual phase (Blackburn et al. 1989). Ecology: G. catenatum is a planktonic red tide species. The first G. catenatum red tide was reported from the Gulf of California with populations close to 1 X 10^ cells/L (Graham 1943). Populations of this species have been recorded from Mexico (Mee et al. 1986), Japan (Ikeda et al. 1989), Australia (Hallegraeff et al. 1988; 1989), Venezuela (La Barbera-Sanchez et al. 1993). the Philippines (Fukuyo et al. 1993) and Europe (Estrada et al. 1984; Franca & Almeida 1989; Giacobbe et al. 1995). G. catenatum produces a characteristic resting cyst (Fig. 6). Cysts are 42-52 ^m in diameter, spherical and brown. They have a very distinct morphology: the surface is covered with microreticulate ornamentations. These cysts can germinate after just two weeks of dormancy and initiate new populations (Blackburn et al. 1989). 40 Harmful Marine Dinotlagellates Cysts are not only a reseeding tool, but also a disbursement agent: G. catenatum was introduced to Australian waters via ships' ballast water (Haltegraeff& Bolch 1991). Toxicity: G. catenatum is a known paralytic shellfish poison (PSP) toxin producer (Morey- Gaines 1982; Mee et al. 1986). This species is the only unarmoured dinotlagellate known to produce PSP toxins (Taylor et al. 1995). First reports of PSP associated with G. catenatum blooms were recorded in Spain (Estrada et al. 1984). Species Comparison: Gymnodinium catenatum can readily be distinguished from other Gymnodinium species by its long chain formations, however, single cells can easily be misidentitled. Chains of G. catenatum can resemble Alexandrium catenella, an anterio- posteriorly compressed species that forms short- chains, however, this species is a cold-water species and is armoured. Chains of G, catenatum can also be confused with Peridiniella catenata, another armoured chain-forming species. The latter species, however, is not toxic, is a cold- water species and has posterior spines (Larsen & Moestrup 1989; Hailegraeff 1991; Taylor et al. 1995). Gyrodinium impudicum, recently described from Spain, can superficially resemble Gymnodinium catenatum with its similar horseshoe shaped apical groove and its tendency toward chain formation. However, Gyrodinium impudicum is smaller in size, differs in shape, forms shorter chains and is not associated with PSP (Fraga et al. 1995). Habitat and Locality: G. catenatum populations are found in warm, temperate coastal waters. Blooms have been reported in Mexico, Argentina, Europe, Australia and Japan (Hailegraeff 1991). Gymnodiniitm niikimotoi Miyake et Kominami ex Oda, 1935 Plate 24, Figs. 1-7 Species Overview: Gymnodinium mikimotoi is an unarmoured, marine, planktonic dinoflagellate species. It is a common red tide former in Japan and Korea associated with massive fish kills. Taxonomic Description: Gymnodinium mikimotoi is an athecate species; i.e. without thecal plates. Cells are small, broadly oval to almost round (Figs. 1,2) and compressed dorso- ventrally (Figs. 3,4). Cells are slightly longer than wide with a characteristic long and straight apical groove to the right of the sulcal axis (Figs. 1-3). The apical groove extends from the ventral side to the dorsal side of the epitheca (Fig. 3) creating a slight indentation at the apex of the cell (Fig. 2). Cells range in size from 18-40 |im in length to 14-35 fim in width (Takayama & Adachi 1984; Fukuyo et al. 1990; Hailegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca is broadly rounded and smaller than the hypotheca (Figs. 1,2). The hypotheca is notched by the widening sulcus at the antapex resulting in a lobed posterior (Figs. 1,2). The wide and deeply excavated cingulum is pre- median, and is displaced in a descending spiral about 2 limes its width (Figs. 1,5). The sulcus slightly invades the epitheca extending from above the cingulum to the antapex (Figs. l,5)(Takayama & Adachi 1984; Fukuyo et al. 1990; Hailegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). INonienclatural Types: Holot)pe: Gymnodinium mikimotoi Oda, 1935; 35-48', tigs. 1-3 Type Locality; NW Pacific Ocean: Gokasho Bay, Japan Synonyms: Gymnodinium nagasakiense Takayama and Adachi, 1984 Gyrodinium aureolum Hulburt, sensu Braarud andHeimdal, 1970 Morphology and Structure: G. mikimotoi is a photosynthetic species with several oval to round yellow-brown chloroplasts, each with a pyrenoid. The large ellipsoidal nucleus is located in the left hypothecal lobe (Fig. 6)(Takayama & Adachi 1984; Fukuyo et al. 1990; Hailegraeff 1991; Taylor etal. 1995; Steidinger & Tangen 1996). Reproduction: G. mikimotoi reproduces asexually by binary fission; cells divide obliquely Harmful Marine Dinoflagellates 41 during mitosis (Fig. 7)(Yamaguchi & Honjo 1990). Ecology: G. mikimotoi is a planktonic species first described from western Japan (Oda 1935). This species is a recurring bloom former in coastal waters of Japan and Korea; red tides commonly occur in wanner months and are associated with massive fish and shellfish kills (Takayama & Adachi 1984). Reported to be eurythermal and euryhaline, populations of G. mikimotoi could presumably over-winter as motile cells, which could then serve as seed populations for a summer red tide (Yamaguchi & Honjo 1989). Moreover, studies conducted in Omura Bay, Japan, revealed that this species can tolerate anoxic or near anoxic conditions utilizing sulfide from the sediment (lizuka 1972). Cells have a distinct swimming pattern: turning over through water, like a falling leaf (Takayama & Adachi 1984). Toxicity: G. mikimotoi is a toxic species associated with massive kills of benthic invertebrates and of both wild and farmed fishes in coastal waters off Japan and Korea; e.g. in 1933 pearl oyster mortalities near Nagasaki, Japan, resulted in an economic loss of $7 million (Oda 1935). For decades red tides of G. mikimotoi have resulted in devastating marine life mortalities, yet the toxin mechanism and principles are poorly understood. Research indicates that this species produces hemolytic and ichthyotoxic substances (Hallegraeff 1991; Taylor et al. 1995). Recently, Seki et al. (1996) extracted a lipid-soluble toxin, gymnodimine, from shelttlsh in Southland, NZ (dubbed 'Southland toxin') after a Gymnodinium cf mikimotoi red tide event. This toxin produced a quick kill in both mice and fish, but was less toxic than brevetoxins. No reported human illnesses have resulted from consumption of tlsh or shellfish from bloom affected areas (Hallegraeff 1991). Species Comparison: G. mikimotoi resembles G. breve: both species are dorso-ventrally flattened and their nucleus is located in the left half of the hypotheca. However, these species differ in several features: G. mikimotoi does not have an apical process; G. breve cells are flafter (dorso-ventra! compression is greater); and the sulcal invasion of the epitheca is deeper in G. breve (Takayama & Adachi 1984). The Pacific Gymnodinium mikimotoi and the European Gyrodinium aureohim are morphologically similar and have been in a state of taxonomic turmoil for over 20 years (Takayama et al. 1998). They are generally regarded as conspecific, although genetic differences between the two populations do exist (Partensky et al. 1988). Controversy, therefore, still remains over the taxonomic status of the Pacific and European populations. Recently, Takayama et al. (1998) conducted an extensive taxonomic study on the morphological differences between the Pacific Gymnodinium mikimotoi and the European Gyrodinium aureolum. There were several morphological differences reported, namely swimming behavior, cell thickness, and shape and position of nucleus: cells of G. aureolum are thicker; the nucleus of G. aureolum is spherical and central, while that of G, mikimotoi is longitudinally elliptical and located in the left lobe of the hypotheca. Habitat and Locality: G. mikimotoi is a cosmopolitan species commonly found in temperate to tropical neritic waters. Blooms have been reported from Australia, Denmark, Ireland, Japan, Korea, Norway and Scotland (Taylor et al. 1995; Steidinger & Tangen 1996). Gymnodinium pulchelliim Larsen, 1994 Plate 25, Figs. 1-6 Species Overview: Gymnodinium pulchellum is an unarmoured, marine, planktonic dinoflagellate species. This species produces red tide blooms and has been associated with fish and invertebrate kills in Japan and Florida. Taxonomic Description: Gymnodinium pulchellum is an athecate species; i.e. without thecal plates. Cells are small and broadly oval with slight dorso-ventral compression (Figs. 1-5). The ventral surface is flattened; the dorsal surface is rounded. A conspicuous and well- defined sigmoid apical groove is present on the epitheca (Figs. 1,2); the groove is a characteristic reversed S-shape (Fig. 2). Cells range in size 42 Harmful Marine Dinoflagellates from 16-25 [im in length to 11-16 jim in width (Fukuyo et al. 1990; Larsen 1994; Taylor et al. 1995; Steidinger & Tangen 1996; Steidinger et al. 1998). The epitheca is slightly smaller than the hypotheca. The wide and deeply excavated cingulum is premedian, and is displaced in a descending fashion 1-1.5 times its width (Figs. 1,3,6). The sulcus slightly invades the epitheca as a finger-like projection (Fig. 2). The sulcus widens and deepens towards the posterior of the cell creating a bilobed hypotheca (Figs. 1,3,4)( Larsen^ 1994; Taylor et al. 1995; Steidinger & Tangen 1996; Steidinger et al. 1998). INomciK-latural Types: Holotype: Gymnodinlum pulchellum Larsen, 1994:32, fig. 58 Type Locality: Tasman Sea: Hobsons Bay, Melbourne, Australia Synonyms: Gymnudinium type '84-K Onoue et al., 1985 IVlorphology and Structure: G. pulchellum is a photosynthetic species with several yellowish- brown chloroplasts. Pyrenoids are also present (Figs. 3,4). The large nucleus is ellipsoidal and located in the left central part of the cell (Figs. 5,6)(Fukuyo et al. 1990; Larsen 1994; Steidinger & Tangen 1996; Steidinger et al. 1998). Reproduction: G. pulchellum reproduces asexualty by binary fission. Ecology: G. pulchellum is a planktonic species first described from southeastern Australia. This species is a bloom-former associated with extensive fish and invertebrate kills in southeast Florida. During one red tide event waters turned an orange-red color with cell levels recorded as high as 19.7 X 10^' cells/L (Steidinger et al. 1998). Toxicity: G. pulchellum is a toxic species associated with tlsh and invertebrate kills from southeast Florida. The presence of this species at two separate fish kills in the Indian River, FL, suggests it is ichthyotoxic (Steidinger et al. 1998). Onoue et al. (1985) demonstrated that G. pulchellum (as Gymnod'mium type '84-K) is ichthyotoxic. Three toxic fractions have been isolated from this species: neurotoxic, hemolytic and hemaglutinative (Onoue & Nozawa 1989). G. pulchellum is most likely responsible for tlsh kills in the Melbourne, Australia, region (Larsen 1994). Species Comparison: Sharing the same habitat and locale, and the same general shape, G. pulchellum can be confused with G. mikimotoi. G. pulchellum, however, is smaller in size and has a distinctive sigmoid apical groove; the apical groove of G. mikimotoi is straight (Larsen 1994). Etymology: The name 'pulchellum" originates from the Latin word pulchellus, 'beautiful little' (Larsen 1994). Habitat and Locality: This species is found in temperate to tropical neritic waters. It has been reported from Hobsons Bay (Melbourne area), Australia, where it is often common during the austral summer and early autumn (Larsen 1994). It has also been recorded from Tasmanian waters (Hallegraeff 1991), Japanese waters (Fukuyo et al. 1990; Onoue et al. 1985; Takayama 1985) and from the Mediterranean (Carrada et al. 1991). More recently it has been identified in the western Atlantic off the east coast of Florida (Steidinger et al. 1998). Due to its minute size, G. pulchellum may have been greatly overlooked in phytoplankton assessments. Gymnodinium saiigiiineitm Hirasaka, 1922 Plate 26, Figs. 1-4 Species Overview: Gymnodinlum sanguineum is an unarmoured, marine, planktonic dinotlagellate species. This cosmopolitan species is a red tide former that has been associated with fish and shellfish mortality events. Taxonomic Description: Gymnodinlum scmguineum is an athecate species; i.e. without thecal plates. This species is highly variable in size and shape. Cells are large, slightly dorso- ventrally flattened and roughly pentagonal (Figs. 1-3). An apical groove is present (Fig. 2). Cells range in size from 40-80 \\.m in length (Hirasaka 1922; Lebour 1925; Dodge 1982; Fukuyo et al. Harmful Marine Dinoflagellates 43 1990; Hallegraeff 1991; Steidinger & Tangen 1996). The epitheca and hypotheca are nearly equal in size. The epitheca is rounded and conical, and the hypotheca is deeply indented by the sulcus creating two posterior lobes (Figs. 1,2). The median cinguium is left-handed and displaced 1- 2 times its width (Figs. 2,4). The sulcus does not invade the epitheca, but expands posteriorly into the hypotheca (Hirasaka 1922; Lebour 1925; Dodge 1982; Pukuyo et al. 1990; Steidinger Tangen 1996). Nomenclatural Types; Holotype: Gymnodinium sanguineum Hirasaka, 1922:161-164, fig. 1 Type Locality: NW Pacific Ocean: Kozusa-ura, Gokasho Bay, Japan Synonyms: Gymnodinium splendens Lebour, 1925 Gymnodinium nelsonii Martin. 1929 Morphology and Structure: G. sanguineum has numerous large, spindle-shaped, reddish-yellow- brown chloroplasts radiating from the center of the cell (Fig. 4). The large nucleus is slightly off-center (Figs. 3,4). Cells can vary from heavily pigmented to pale yellow or nearly colorless (Hirasaka 1922; Lebour 1925; Dodge 1982; Fukuyo et al. 1990; Steidinger & Tangen 1996). Mixotrophy has been observed for this species: in the Chesapeake Bay G. sanguineum preys on ciliate protozooplankton (Bockstahler & Coats 1993). Reproduction: G. sanguineum reproduces asexually by binary fission; cells divide obliquely during mitosis (Dodge 1982). Ecology: G. sanguineum is a planktonic species common in estuarine and coastal waters. This cosmopolitan species is a bloom-former associated with shellfish and fish kills. The first G. sanguineum red tide was reported from Kozusa-ura, Gokasho Bay, Japan (Hirasaka 1922). Red tide events caused by this species have since been recorded from other coastal regions of Japan (Fukuyo et al. 1990). It is a common red tide bloom species in Australian and New Zealand coastal waters as well (Hallegraeff 1991). G. sanguineum is a common red tide species in the Chesapeake Bay where levels as high as 8.8 X 10 cells/L have been reported (Bockstahler & Coats 1993). One bloom in Coyote Bay, Gulf of California, Mexico, cell densities reached 1.0 X 10^ cells/L (Keifer & Lasker 1975). Robinson and Brown (1983) and Voltolina (1993) observed possible sexual stages of G. sanguineum from a recurrent bloom. They speculate that this species may form resting cysts to reseed a region in the next bloom season. Nakamura et al. (1982) reported that cultures of G. sanguineum can tolerate a wide range of temperatures (13-24 °C) and salinities (15-35 o/oo). Toxicity: G. sanguineum is a red tide species associated with fish and invertebrate kills. Cardwell et al. (1979) reported the acute toxicity of this species to larval stages of two species of oysters in Puget Sound, Washington State. And G. sanguineum is believed to be responsible for at least one reported fish mortality event in Peru (Jordan 1979). Tindall et al. (1984) and Carlson and Tindall (1985) demonstrated one isolate of this species to be potentially toxic; however, the toxin principles have yet to be elucidated. Etymology: The name 'sanguineum ' originates from the Latin word for blood describing the resulting color of the water after a red tide event of this species (Hirasaka 1922). Habitat and Locality: G. sanguineum is commonly found in temperate to tropical neritic waters (Steidinger & Tangen 1996). Blooms have been recorded from Japan (Hirasaka 1922; Fukuyo et al. 1990). Australia and New Zealand (Hallegraeff 1991), and from the Atlantic and Pacific American coasts (Keifer & Lasker 1975; Robinson & Brown 1983; Bockstahler & Coals 1993; Voltolina 1993). Gynuwdhiuim veneficum Baiiantine, 1956 Plate 27, Figs. 1-3 Species Overview: Gymnodinium veneficum is an unarmoured, marine, planktonic dinofiagellate species. This small species has been associated with fish and shellfish mortality events. 44 Harmful Marine Dinoflagellates Taxonomic Description: Gymnodinium veneficum is an athecate species; i.e. without thecal plates. Cells are small and ovoid without dorso-ventral compression (Figs. 1-3). Cells range in size from 9-18 ^m in length to 7-14 ^im in width (Ballantine 1956; Dodge 1982; Taylor etal. 1995). The epitheca and h>potheca are equal in size. The cell's anterior end is slightly pointed; the epitheca is without an apical groove (Fig. 1). The hypotheca is rounded with a slight indentation at its posterior end (Fig. 2). The deep cingulum is displaced in a descending spiral 1-2 times its width (Figs. 1,3). The sigmoid sulcus slightly invades the epitheca (Figs. 1,3) (Ballantine 1956; Dodge 1982; Taylor et al. 1995). Nomenclatural Types: Holotype: Gymnodinium veneficum Ballantine, 1956:468-474, tigs. 6-17 Type Locality: English Channel: off King William Point, Devonport, United Kingdom Synonyms: Gymnodinium vitiligo Ballantine, 1956 Morphology and Structure: G. veneficum is a photosynthetic species and usually has four irregularly shaped, golden-brown chloroplasts with pyrenoids; occasionally two to eight are present. The large round nucleus is centrally located (Figs. 2,3)(Ballantine 1956; Dodge 1982; Taylor etal. 1995). Reproduction: G. veneficum reproduces asexually by binary fission; cells divide obliquely during mitosis (Ballantine 1956). Ecology: G. veneficum is a planktonic species described from the English Channel (Ballantine 1956). Toxicity: G. veneficum is a known toxic species; it produces an exotoxin lethal to a wide variety of invertebrates and fish (Ballantine 1956; Abbott & Ballantine 1957; Dodge 1982). Species Comparison: In general cell shape and size, G. veneficum can easily be mistaken for G. micrum, a non-toxic species. However, the former species usually has four chloroplasts present and is toxic to invertebrates and fish (Taylor etal. 1995). Habitat and Locality: G. veneficum was described from the English Channel. It may be a wide-spread species, but due to its minute size, it most likely has been greatly overlooked in phytoplankton assessments (Ballantine 1956; Dodge 1982). Gyrodinium galathcanurn (Braarud) Taylor, 1992 Plate 28, Figs. 1-4 Species Overview: Gyrodinium galatheanum is an unarmoured, marine, planktonic dinoflagellate species. It is a common red tide former discovered in Walvis Bay, South Africa, associated with fish kills. Taxonomic Description: Gyrodinium galatheanum is an athecate species; i.e. without thecal plates. Cells are small and oval to round in ventral view (Figs. 1-3). A well-defined apical groove is present ventrally on the anterior of the cell (Figs, 1,2,4). The apical groove can produce a slight indentation at the apex (Fig. I). Cells range in size from 9-17 |im in length to 8-14 fim in width (Braarud 1957; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca and hypotheca are both round (Figs. 1-3). The cingulum is displaced in a descending fashion up to 3 times its width (Figs. 1,2,4). The broad cingulum is deeply excavated and houses the transverse tlagellum (Figs. 1-3). The short and narrow sulcus slightly invades the epitheca adjacent to the apical groove (Figs. l,2,4)(Braarud 1957; Taylor et al. 1995; Steidinger & Tangen 1996). INomenclatural Types: Holot}pe: Gymnodinium galatheanum Braarud, 1957; 137-138, fig. la-e Type Locality: South Atlantic Ocean: Walvis Bay, South Africa Synonyms: Gymnodinium micrum (Leadbeater et Dodge) Loeblich, 111 Woloszynskia micro Leadbeater and Dodge, 1966 Harmful Marine Dinoflagellates 45 Basionym: Gymnodinium galatheanum Braarud, 1957 Morphology and Structure: G. galatheanum is a photosynthetic species witii several round chioroplasts. The large nucleus is round and centrally located (Figs. 3,4). This species does not have peridinin as a major accessory pigment, but has a fucoxanthin derivative and chlorophyll c3 (Braarud 1957; Bjornland & Tangen 1979; Johnsen & Sakshaug 1993; Taylor et al. 1995; Steidinger & Tangen 1996). Reproduction: G. galatheanum reproduces asexually by binary fission. Ecology: G. galatheanum is a bloom-forming planktonic species. Blooms of this species were first recorded from Walvis Bay, South Africa (Braarud 1957). Blooms have since been reported from the Oslofjord, Norway (Bjornland & Tangen 1979) and along the southern coast of Norway (Dahl & Yndestad 1985). Li et al. (2000) recently observed mixotrophic behaviour in G. galatheanum from the Chesapeake Bay. This species was observed to feed on cryptophytes under light and/or nutrient stressed conditions suggesting that this primarily photosynthetic species uses phagotrophy during nutrient-replete conditions to furnish major nutrients necessary for photosynthesis. Toxicity: G. galatheanum is a toxic species associated with fish kills in Walvis Bay, South Africa (Braarud 1957; Steemann Nielsen & Aabye Jensen 1957; Pieterse & Van Der Post 1967). Although this species has been linked to marine life mortalities, mussels and juvenile cod (Nielsen & Stromgren 1991; Nielsen 1993), the toxin principles have yet to be determined (Copenhagen 1953; Pieterse & Van Der Post 1967). Species Comparison: In shape and size Gyrodinium galatheanum resembles two small athecate gymnodinoids, Gymnodinium veneficum and G. micrum (Taylor et al. 1995). Physiologically Gyrodinium galatheanum is closely related to the toxic species Gyrodinium aureulum. Both lack peridinin while both have chlorophyll c3, which is characteristic of several bloom-forming prymnesiophytes (Johnsen & Sakshaug 1993). Habitat and Locality: This species has been reported from cold waters in the North and South Atlantic Oceans: North Sea, British Isles (Larsen & Moestrup 1989); Oslofjord, Norway (Bjornland & Tangen 1979); and Walvis Bay, South Africa (Braarud 1957). G. galatheanum may be a wide-spread species but due to its minute size, it most likely has been greatly overlooked in phytoplankton assessments (Taylor etal. 1995). Lingulodmium poly edrum (Stein) Dodge, 1989 Plate 29, Figs. 1-6 Species Overview: Lingulodinium polyedrum is an armoured, marine, bioluminescent dinoflagellate species. This warm-water species is a red tide former that has been associated with fish and shellfish mortality events. Taxonomic Description: Cells oi Lingulodinium polyedrum are angular, roughly pentagonal and polyhedral-shaped (Fig, I). Cells range in size from 40-54 jiiTi in length and 37-53 ^m in transdiameter width. No apical horn or antapical spines present (Fig. 1). Thecal plates are thick, well defined, and coarsely areolate. Distinct ridges are present along the plate sutures (Figs. 1,2). Numerous large trichocyst pores are present within areolae (Fig. 3)(Kofoid 1911; Dodge 1985; 1989 Lewis & Burton 1988; Fukuyo et al. 1990; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gonyaidax polyedra Stein. 1883: p. 13, pi. 4, figs. 7-9 Type Locality: unknown Synonyms: Gonyaulax polyedra Stein, 1 883 Lingulodinium machaerophorum (Deflandre and Cookson) Wall, 1967 (cyst) Hystrichosphaeridium machaerophorum Deflandre and Cookson, 1955 (cyst) Thecal Plate Description: The plate formula for L polyedrum is: Po, 3', 3a, 6", 6c, 7s, 6"', 2"". The epitheca bears shoulders, nearly straight 46 Harmful Marine Dinoflagellates sides, and an off-center apex which is flattened or slightly pointed (Figs. 1,4). The apical pore plate (Po) contains a raised inner elliptical ridge (Fig. 2). The first apical plate (1') is long and narrow, conies in direct contact with the Po, and bears a ventral pore on its right side (Figs. 1,2,4). The deeply excavated cinguluni is nearly equatorial, and displaced one to two times its width. !t is descending with narrow ribbed lists (Figs. 1,2,4). The deep sulcus invades the epitheca slightly and widens posteriorly. The hypotheca has straight sides and a truncated antapex (Figs. [,2,4)(Kofoid 1911; Dodge 1985; Dodge 1989; Lewis & Burton 1988; Fukuyo et al. 1990; Steidinger & Tangen 1996). Morphology and Structure: L polyedrum is a photosynthetic species with dark orange-brown chloroplasts. The unusual carotenoid, peridinin, is present in the chloroplasts. Also present is a pusule, a C-shaped nucleus, and scintillons (light-emitting organelles)(Kofoid 1911; Schmitter 1 97 1 ; Jeffrey et al. 1975). Reproduction: L. polyedrum reproduces asexualiy by binary fission. Sexual reproduction is also part of the life cycle of this species producing spherical spiny cysts. Ecology: L polyedrum is a bioluminescent planktonic species commonly found in neritic waters. It is responsible for magnificent displays of phosphorescence at night in warm coastal waters (Kofoid 191 1). This warm-water species is a red tide former that has been associated with fish and shellfish mortality events. Deadly red tides have been reported from southern California (San Diego region)(Kofoid 1911; Allen 1921), as well as in the Adriatic Sea (Italy and Yugoslavia) where cell levels as high as 2 X 10 cells/L have been reported (Marasovic 1989; Bruno etal. 1990). This species forms colorless spherical spiny cysts (35-50 )im in diameter). The numerous tapering spines can reach up to 17 ^m in length, al! bearing spinules on their distal ends (Figs. 5,6) (Kofoid 1911; Dodge 1985; 1989; Fukuyo et al. 1990). The cyst of this species is able to fossilize (found in fossil deposits all the way back to the late Cretaceous period): the hystrichosphere (fossilized dinotlagellate cyst) Lingulodinium /luichaerophurum (Detlandre and Cookson) Wall, 1967 was discovered to be the resting spore of L polyedrum (Wall 1967; Fensome et al. 1993). Marasovic (1989) reported production of temporary resting cysts in a waning red tide dominated by L. polyedrum in the Adriatic Sea (Yugoslavia). Near the end of a bloom, the population produced temporary cysts and remained in the plankton. Once environmental conditions were favorable again, the cysts were able to re-seed the area, and thus initiate another red tide event. Toxicity: Bruno et al. (1990) reported the presence of a paralytic shellfish poison (PSP) toxin, saxitoxin, in water samples taken during a bloom of L polyedrum. Habitat and Locality: L. polyedrum is a widely distributed species found in warm temperate and subtropical waters of coastal areas (Kofoid 1911; Dodge 1985; 1989; Steidinger & Tangen 1996). Noctihica schitillans (Macartney) Kofoid et Swezy, 1921 Plate 30, Figs. 1-4 Species Overview: Noctihica scintillam is an unarmoured, marine planktonic dinofiagellate species. This large and distinctive bloom forming species has been associated with fish and marine invertebrate mortality events. Taxononiic Description: Noctihica scintillans is a distinctively shaped athecate species in which the cell is not divided into epitheca and hypotheca. Cells are very large, inflated (balloon-like) and subspherical (Figs. 1-4). The ventral groove is deep and wide, and houses a flagellum, a tooth and a tentacle (Figs. 1,2,4). Only one tlagellum is present in this species and is equivalent to the transverse flagellum in other dinoflagellates (Fig. 1). The tooth is a specialized extension of the cell wall (Fig. 4). The prominent tentacle is striated and extends posteriorly (Fig. 4). Cells have a wide range in size: from 200-2000 \xm in diameter (Zingmark 1970; Dodge 1973; Dodge 1982; Lucas "1982; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Harmful Marine Dinoflagellates 47 Nomenclatural Types: Holotype: Medusa scintillim Macartney, 1810: 264-265, pi. 15, figs. 9-12 Type Locality: North Sea: Heme Bay, Kent, England Synonyms: Medusa scintillins Macartney, 1810 Noctiluca miiiaris Suriray, 1836 Morphology and Structure: Noctiluca scintillans is a nonphotosynthetic heterotrophic and phagotrophic dinoflagellate species; chloroplasts are absent and the cytoplasm is mostly colorless (Figs. 1,2). The presence of photosynthetic symbionts can cause the cytoplasm to appear pink or green in color (Sweeney 1978). A number of food vacuoles are present within the cytoplasm. A large eukaryotic nucleus is located near the ventral groove with cytoplasmic strands extending from it to the edge of the cell (Fig. 2)(Zingmark 1970; Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Steidinger &Tangen 1996). Reproduction: Noctiluca scintillans reproduces asexually by binary fission (Fig. 3) and also sexually via formation of isogametes. This species has a diplontic life cycle: the vegetative cell is diploid while the gametes are haploid. The gametes are gymnodinioid with dinokaryotic nuclei (Zingmark 1970). Ecology: Noctiluca scintillans is a strongly buoyant planktonic species common in neritic and coastal regions of the world. It is also bioluminescent in some parts of the world. This bloom-forming species is associated with fish and marine invertebrate mortality events. A^. scintillans red tides frequently form in spring to summer in many parts of the world often resulting in a strong pinkish red or orange discoloration of the water (tomato-soup). Blooms have been reported from Australia (Hallegraeff 1991), Japan, Hong Kong and China (Huang & Qi 1997) where the water is discolored red. Recent blooms in New Zealand were reported pink with cell concentrations as high as 1.9 X 10^ cells/L (Chang 2000). In Indonesia, Malaysia, and Thailand (tropical regions), however, the watercolor is green due to the presence of green prasinophyte endosymbionts (Sweeney 1978; Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger cS: Tangen 1996). This large cosmopolitan species is phagotrophic, feeding on phytoplankton (mainly diatoms and other dinoflagellates), protozoans, detritus, and fish eggs (Fig. 2)(Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Toxicity: Toxic blooms of N. scintillans have been linked to massive fish and marine invertebrate kills. Although this species does not produce a toxin, it has been found to accumulate toxic levels of ammonia which is then excreted into the surrounding waters possibly acting as the killing agent in blooms (Okaichi & Nishio 1976; Fukuyo et al. 1990). Extensive toxic blooms have been reported off the east and west coasts of India, where it has been implicated in the decline of fisheries (Aiyar 1936; Bhimachar & George 1950). Habitat and Locality: Noctiluca scintillans is a cosmopolitan species distributed world wide in cold and warm waters. Populations are commonly found in coastal areas and embayments of tropical and subtropical regions (Dodge 1982; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Remarks: This species is frequently referred to as A', miiiaris although Macartney's specific name has priority. Taylor (1976) suggests that the simplest solution to the problem of nomenclature is to accept the priority of the 'scintillans' especially as this has been used by two major works (Kofoid & Swezy 1921; Lebour 1925). Ostreopsis heptagona Norris, Bomber et Balech, 1985 Plate 31, Figs. 1-6 Species Overview: Ostreopsis heptagona is an armoured, marine, benthic dinoflagellate species. It was discovered in the Florida Keys. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The 48 Harmful Marine Dinoflagellates epitheca and hypotheca are not noticeably different in size. Unique features of this genus are on the cingulum. In ventral view the cinguluni reveals two prominent structures: a ventral plate (Vp) with a ventral pore (Vo), and an adjacent curved rigid plate (Rp). The distinguishing feature at the species level is the shape of the first apical plate (T) on the epitheca (Fig. IXFaustetal. 1996). Cells of Ostreopsis heptagona are large, broadly oval, oblong and pointed (Figs. 1-2). Thecal surface is smooth with scattered small round pores (diam.=0.3 |im) that can only be observed at the SEM level (Figs. 1,2). Cells have a dorsoventral diameter of 80-108 (im, and a transdiameter of 46-59 jim (Faust et al. 1996). Nomenclatural Types: Iconotype: Ostreopsis heptagona Norris, Bomber andBalech, 1985: fig. I Type Locality: Gulf of Mexico: Knight Key, Florida, USA Thecal Plate Description: The plate formula of Ostreopsis heptagona is: Po, 3', 7", 6c, 6s?, Vp, Rp, 5'", !p, 2""(Fig. 5). The epitheca contains 1 1 plates. The apical pore plate (Po) is 15 |im long, narrow and curved (Figs. 1,3), situated between apical plates I', 2' and 3', with a long, slit-like apical pore. The 1' plate, the distinguishing plate for this species, is large and irregularly heptagonal (seven-sided)(Figs. 1,5). The hypotheca has eight plates. The posterior intercalary plate (Ip) is one of the most characteristic plates of O. heptagona; it is long and narrows dorsally, extending along the dorso- ventral axis (Figs. 2,5)(Faust et al. 1996; Norris etal. 1985). The cingulum is equatorial and narrow (Figs. 1-3). Within the cingulum the Vo is situated on the Vp, adjacent to the Rp (Fig. 4)(Faust et al. 1996). Norris et al. (1985) identified 5 sulcal plates and a transitional plate (t) in this species. Morphology and Structure: Ostreopsis heptagona is a photosynthetic species. Mixotrophy has been documented in other specis of this genus with the Vo as the proposed feeding apparatus (Faust et al. 1996). Reproduction: Cells of O. heptagona reproduce asexual ly by binary Fission. Ecology: Cells of O. heptagona are frequently found as epiphytes on macroalgae in the Caribbean (Morton & Faust 1997). Live cells exhibit an unusual jerky swimming motion and a strong positive geotropic tendency. Cells almost immediately attach to the nearest substrate. Cells attach tenaciously by a network of mucilage strands (Fig. 3) which are expelled by thecal pores (Norris et al. 1985). Toxicity: This species was determined to be toxic (J. Babinchak, according to Norris et al. 1985). Species Comparisons: Ostreopsis heptagona is distinguished by two major features: a) an irregulary-shaped asymmetric heptagonal (seven- sided) r plate that occupies the left center of the epitheca (this plate is hexagonal, six-sided, in all other species of this genus) (Faust et ai. 1996; Steidinger & Tangen 1996); and b) the pentagonal and dorso-ventrally elongate I p plate in the hypotheca (Faust et al. 1996). In O. heptagona plate 5" is pentagonal as it contacts plates I', 3' and 6", and plate 6" is quadrangular and does not touch 3'. In both O, siamensis and O. ovata plate 5" is quadrangular and does not touch 1', while 6" is hexagonal and contacts two apical plates, 1' and 3'. Plate Ip in O. heptagona is rather narrow, and is always curved, concave to the left and gradually narrows dorsally (Faust et al. 1996). Plate Ip in O. siamensis is also narrow, but maintains nearly the same width throughout its length. This plate is different in O. ovata: Ip is comparatively wider and shorter, and widens dorsally (Norris et al. 1985). Etymology: The name 'heptagona' refers to the distinct seven-sided shape of the first apical plate of this species. Habitat and Locality: Populations of O. heptagona have been reported as epiphytic on macroalgae in the Caribbean Sea (Morton & Faust 1997), and found in the plankton in the Florida Keys (Steidinger & Tangen 1996). Maximum densities were reported for O. heptagona associated with Dictyota dichotoma (Bomber 1985) and Acanthophora spicifera (Morton & Faust 1997). Harmful Marine Dinoflagellates 49 OStreOpsis lenticiitaris Fukuyo, 1981 Plate 32, Figs. 1-8 Species Overview: Ostreopsis lenticularis is an armoured, marine, benthic dinoflagellate species. It was discovered as an epiphyte on macroalgae in the Gambler and Society Islands of French Polynesia, and New Caledonia, Pacific Ocean. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The epitheca and hypotheca are not noticeably different in size. Unique features of this genus are on the cingulum. In ventral view the cingulum reveals two prominent structures: a ventral plate (Vp) with a ventral pore (Vo), and an adjacent curved rigid plate (Rp). The distinguishing feature at the species level is the shape of the first apical plate (1') on the epitheca (Fig. l)(Faustetal. 1996). Cells of Ostreopsis lenticidaris are lenticulate to broadly oval (Figs. 1,2). The celt surface is smooth and covered with randomly spaced pores (0.4 \.ivc\ diameter) with smooth raised edges (Figs. 1-4); the pores are large and round (Fig. 3). Cells have a dorso-ventral diameter of 65-75 pm and a transdiameter of 57-63 pm (Faust et al. 1996;Fukuyo 1981). Nomenclatural Types: Holotype: Ostreopsis lenticidaris Fukuyo, 1981: figs. 30-34 Type Locality: South Pacific Ocean: Gambler and Society Islands, and New Caledonia Thecal Plate Description: The plate formula of Ostreopsis lenticidaris is: Po, 3', 7", 6c, 6s?, Vp, Rp, 5'", Ip, 2""(Fig. 6). The epitheca contains 1 1 plates. The narrow apical pore plate (Po) is 16 um long (average) with a slit-like apical pore, and is situated adjacent to apical plate 2' (Figs. 1,5). The r plate is large, irregularly pentagonal-shaped, and situated in the center (Figs. I,5)(Faust et al. 1996). The hypotheca is composed of eight plates. Plate Ip, situated centrally, is a narrow, asymmetric, pentagonal plate (Figs. 2,5). Plate 1"" contacts the sulcal region (Fig. 6)(Faust et al. 1996). The lipped cingulum is narrow and shallow with a smooth edge (Figs. 1,2,4). Within the cingulum is the Vo located on the Vp, and adjacent to a Rp (Figs. 4,5). The shape of the Vp varies from oblong to circular. The sulcus is small and hidden (Faust et al. 1996). Morphology and Structure: Ostreopsis lenticidaris is a photosynthetic species with many golden-brown chloroplasts. A large nucleus is located posteriorly (Fukuyo 1981). There is evidence of mixotrophy in this species: prey organisms are engulfed via the Vo, the proposed feeding apparatus (Faust et al. 1996). Reproduction: Ostreopsis lenticidaris reproduces asexually by binary fission. Ecology: O. lenticidaris can be benthic, epiphytic or tycoplanktonic (Steidinger & Tangen 1996) commonly associated with macroalgae. in the plankton, attached to soft coral and between sand grains. Engulfed cells were often observed in this species collected from Belizean waters (Faust et al. 1996). Toxicity: This is a known toxic species; it produces ostreotoxin (OTX), a water-soluble toxin (Tindall et al. 1990), and an unnamed toxin (Ballantineetal. 1988). Species Comparisons: Ostreopsis lenticidaris differs from other species in the genus by its lentil-like cell shape, medium size and randomly spaced round pores. The size and location of plates 2'", 3'" and 4'" are also distinguishing features (Faust et al. 1996). This species closely resembles Gambierdiscus toxicus in size, shape and color, but O. lenticidaris has a slightly pointed ventral area while G. toxicus has a round and indented one (Fukuyo 1981). O. lenticidaris is also similar to O. siamensis in shape and thecal plate configuration (Fukuyo 1981). Habitat and Locality: Populations of O. lenticidaris were originally found in the Gambler and Society Islands and New Caledonia, Pacific Ocean, associated with macroalgae (Fukuyo 1981). Populations can be found from tropical shallow waters to offshore reefs (Steidinger & Tangen 1996). Cells have been observed 50 Harmful Marine Dinoflagellates epiphytic on macroalgae {Dictyota sp. and Acanthophora spicifera) in the Caribbean region (Carlson & Tindall 1985; Ballantine et al. 1988; Morton & Faust 1997) and the SW Indian Ocean (Quod 1994). In the Caribbean, this species has been observed in the plankton (Faust 1995), attached to soft corals (Ballantine et al. 1985; Carslon & Tindall 1985) and between sand grains (Ballantine et al. 1985; Carslon & Tindal! 1985; Faust 1995). Ostreopsis mascarenensis Quod, 1994 Plate 33, Figs. 1-8 Species Overview: Ostreopsis mascarenensis is an armoured, marine, benthic dinoflagellate species. It was discovered in shallow barrier reef environments and coral reefs in the Mascareignes Archipelago, SW Indian Ocean. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The epitheca and hypotheca are not noticeably different in size. Unique features of this genus are on the cingulum. in ventral view the cingulum reveals two prominent structures: a ventral plate (Vp) with a ventral pore (Vo), and an adjacent curved rigid plate (Rp). The distinguishing feature at the species level is the shape of the tlrst apical plate (1') on the epitheca (Fig. l)(Faustetal. 1996). Cells of O. mascarenensis are very large and broadly oval (Figs. 1,2,7). This is the largest species in the genus. Cells have a dorsoventral diameter of 155-178 ^m and a transdiameter of 1 18-134 jim. The thecal surface is smooth with small evenly distributed pores (Figs. 1-4) that often contain ejected trichocysts (Fig. 6). The pores are round with two small openings (diam.=0.6 |im) with smooth edges (Fig. 3)(Quod 1994; Faust etal. 1996). Nomenclatural Types: Holotype: Ostreopsis mascarenensis Quod, 1994: fig. 1 Type Locality: West Indian Ocean: Saint Leu, Reunion Island, Mascareignes Archipelago Thecal Plate Description: O. mascarenensis is a large cell with very large plates (Fig. 1). The plate formula for this species is: Po, 3', 7", 6c, 6s?, Vp, Rp, 5'", Ip, 2"". On the epitheca, the apical pore plate (Po) bears a long curved slit- like apical pore (26 |Lim) with an array of minute openings (Fig. 4). The 1' plate is large, long and hexagonal, 102 |im long and 40 jum wide (Fig. I). In the hypotheca, the posterior intercalary plate (Ip) is long and wide (Fig. 2). Plate 1'" is large compared to other species in the genus (Fig. 8)(Quod 1994; Faust etal. 1996). The lipped cingulum is narrow with a smooth edge (Figs. 1,2,5). It houses the Vo situated on the Vp, and the Rp (Fig. 6). The sulcus is recessed and hidden (Fig. 5)(Quod 1994; Faust et al. 1996). Morphology and Structure: Cells of Ostreopsis mascarenensis are photosynthetic with light golden-colored chloroplasts. This species has two pusules in the sulcus and one dorsal red pyrenoid (Quod 1994). There is evidence of mixotrophy in this species: prey organisms are engulfed via the Vo, the proposed feeding apparatus (Faust et al. 1996). Reproduction: O. mascarenensis reproduces asexually by binary fission. Ecology: Cells of O. mascarenensis are commonly associated with dead corals and sediments and as epiphytes on macroalgae (Quod 1994; Faust et al. 1996). Cells exhibit geotropic swimming. Cells may form blooms, reaching a density of > 10,000 cells.g fresh weight of algal tissue (Quod 1994). Toxicity: This species produces an unnamed toxin which may cause ciguatera (Quod 1994). This toxin has not been detected in fish (Morton, S.L., personal communication 1998). Species Comparisons: O. mascarenensis differs from other species of the genus by its large size, thecal morphology, geotropic swimming behaviour and dissimilar plates, in particular, plates r,2',3M"'and lp(Quod 1994). Habitat and Locality: Populations of O. mascarenensis can be commonly found in Harmful Marine Dinotlagellates 51 shallow (2-5tTi) barrier reef environments and coral reefs in the SW Indian Ocean. This species has been observed as an epiphyte on Turbinaria sp., Galaxcmra sp., dead corals and sediments at Mayotte, Reunion and Rodriguez Islands (Quod 1994). Cells were also discovered from the lagoonal island. Tobacco Cay, Belize, in the Caribbean Sea (Faust et al. 1996). Ostreopsis ovata Fukuyo, 1981 Plate 34, Figs. 1-7 Species Overview: Ostreopsis ovata is an armoured, marine, benthic dinotlagellate species. It was discovered from French Polynesia, New Caledonia and the Ryukyu Islands, Pacific Ocean. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The epitheca and hypotheca are not noticeably different in size. Unique features of this genus are on the cingulum. In ventral view the cingulum reveals two prominent structures: a ventral plate (Vp) with a ventral pore (Vo), and an adjacent curved rigid plate (Rp). The distinguishing feature at the species level is the shape of the first apical plate (T) on the epitheca (Fig. l)(Faustetal. 1996). Cells of O. ovata are tear-shaped, ovate and ventrally slender (Figs. 1,2,6). It is the smallest species in the genus. Thecal surface is smooth, ornamented with minute, evenly distributed pores (0.07 |am diameter)(Figs. 1-4). Cells have a dorsoventral diameter of 47-55 |im and transdiameter of 27-35 ^m (Faust et al. 1996). Nomenclatural Types: Holotype: Ostreopsis ovata Fukuyo, 1981: figs. 35-38 Type Locality: Pacific Ocean: French Polynesia, New Caledonia and the Ryukyu Islands Thecal Plate Description: Thecal plates of Ostreopsis ovata are very thin and delicate, and their morphology is very difficult to preserve. The plate formula for this species is: Po, 3', 7", 6c, 6s?, Vp, Rp, 5'", Ip, 2"". In the epitheca, the r plate is long and hexagonal, and occupies the left center of the cell (Fig. 1). The apical pore plate (Po) features a short asymmetrical slit-like apical pore, and is associated with narrow apical plate 2' (Figs. 1,4). In the hypotheca, the posterior intercalary plate (Ip) is long and narrow (9 X 27 ^im) (Fig. 2) (Faust et al. 1 996). Cingulum is equatorial, relatively wide, and bordered by narrow lists (Figs. 1,2). Within the cingulum, the Vo is situated on the Vp surrounded by the Rp (Fig. 5)(Faust et al. 1996). The sulcus contains eight plates (Steidinger & Tangen 1996). Morphology and Structure: Cells of Ostreopsis ovata are photosynthetic containing many golden chloroplasts. Large ovate nucleus is posterior (Fig. 6)(Fukuyo 1981). There is evidence of mixotrophy in this species: prey organisms are engulfed via the Vo, the proposed feeding apparatus (Faust et al. 1996). Reproduction: O. ovata reproduces asexually by binary fission. Ecology: O. ovata can be tycoplanktonic, benthic or epiphytic (Steidinger & Tangen 1996). Engulfed cells were often observed in this species collected from Belizean waters (Faust et al. 1996). Toxicity: This species produces an unnamed toxin (Nakajimaet al. 1981). Species Comparisons: O. ovata differs from the other species in the genus by its small size, very delicate thecal plates and a short, straight Po. It is readily identifiable from O. siamensis and O. lenticularis by its ovoidal, tear-shaped body (Fukuyo 1981). Habitat and Locality: Ostreopsis ovata is infrequently observed in the field. Populations are usually found in protected, inshore regions from the tropical Pacific Ocean (Fukuyo 1981; Yasumoto et al. 1987; Quod 1994), the Caribbean Sea (Besada et al. 1982; Carlson & Tindall 1985) and the Tyrrhenian Sea (Tognetto et al. 1995). Substrate specificity for this species needs to be determined. 52 Harmful Marine Dinoflagellates Ostreopsis siamensis Schmidt, 1902 Plate 35, Figs. 1-8 Species Overview: Ostreopsis siamensis is an armoured, marine, benthic dinotlagellate species. It was first identitled from plankton samples from the Gulf of Siam (Thailand). Taxonomic Description: Species in this genus are anterio-posteriorly compressed and are observed in apical or antapical view. The epitheca and hypotheca are not noticeably different in size. Unique features of this genus are on the cingulum. In ventral view the cingulum reveals two prominent structures: a ventral plate (Vp) with a ventral pore (Vo), and an adjacent curved rigid plate (Rp). The distinguishing feature at the species level is the shape of the tlrst apical plate (!') on the epitheca (Fig. l)(Faustetal. 1996). Cells of O. siamensis are ovate and tear- shaped (Figs. 1,2,7,8). The thecal surface is smooth with evenly scattered round pores (Figs. 1-3). Large (0.5 j.mi diameter) and small (0.1 |.uii diameter) pores are present (Fig. 4). Cells have a dorsoventral diameter of 108-123 j.im and a transdiameter of 76-86 ^im (Faust et al. 1996). INomenclatural Types: Holotype: Ostreopsis siamensis Schmidt, 1902: i1gs. 5-7 Type Locality: Gulf of Thailand: Thailand Thecal Plate Descri|)tion: The plate formula for Ostreopsis siamensis is: Po, 3', 7", 6c, 6s?, Vp, Rp, 5'", Ip, 2"" (Fig. 8). On the epitheca, a narrow curved apical pore plate (Po) (Fig. 1) is closely associated with the narrow apical plate 2' (Fig. 3). The apical pore appears as a curved slit 2 [xm long (Fig. 3). The I' plate is large, narrow and pentagonal (Fig. 1). The hypotheca is composed of eight plates (Fig. 2). The posterior intercalary plate (Ip) is large, elongated (26 X 55 |am), and pentagonal (Fig. 2). Plate 1"" contacts the sulcal region (Figs. 2,5)(Faust et al. 1996). The narrow cingulum is deep with a smooth edge (Figs. 1-3) and is composed of six plates. In the cingulum the Vo is situated on the Vp next to the Rp (Figs. 5,6). The Vo may be open or closed. The sulcus is small, recessed and hidden below plates 1"" and 2""(Faust et al. 1996). Morphology and Structure: Cells of O. siamensis are photosynthetic and contain numerous golden-brown chloroplasts. A large nucleus is posterior. There is evidence of mixotrophy in this species: prey organisms are engulfed via the Vo, the proposed feeding apparatus (Faust et al. 1996). Reproduction: O. siamensis reproduces asexually by binary fission. Ecology: O. siamensis are benthic, epiphytic, and can be tycoplanktonic (Steidinger & Tangen 1996). They have been observed in plankton samples, but it is most frequently associated with sand and as epiphytes on macroalgae. These cells swim very slowly and spin around the dorso-ventral axis (Fukuyo 1981). Engulfed cells were often observed in this species collected from Belizean waters (Faust et al. 1996). Toxicity: This species is a known toxin producer; It produces an analog of palytoxin (Nakajimaetal. 1981; Usami et al 1995). Species comparison: O. siamensis differs from other species of the genus by a number of features: a. a tear-drop shape; b. large cell size; and c. small round evenly distributed thecal pores (Faust et al. 1996). Habitat and Locality: Ostreopsis siamensis has been observed in various tropical regions of the world. Populations were originally discovered in plankton samples collected from the Gulf of Siam (Thailand) (Schmidt 1902, figs. 5-7) and then seldom observed again for over 70 years. Cells were later found as epiphytes on macroalgae in the Pacific Ocean (Taylor 1979; Yasumoto et al. 1980; Fukuyo 1981; Nakajima et al. 1981; Holmes et al. 1988), the SW Indian Ocean (Quod 1994), the Florida Keys (Bomber 1985), and the Caribbean region (Carlson 1984; Tindall et al. 1984; Ballanline et al. 1988; Faust 1995; Faust & Morton 1995). They have also Harmful Marine Dinoflagellates 53 been associated with sand in the Caribbean (Faust etal. 1996). Pfiesteria piscicida Steidinger et Burkholder, 1996 Plate 36, Figs. 1-9 Species Overview: Pfiesteria piscicida is a putatively toxic dinotlagellate species with flagellated and cyst stages. This species, dubbed the 'ambush predator', was first observed in the Pamlico Sound, North Carolina, USA, in 1991 after a massive fish kill. Pfiesteria piscicida has been associated with fish kills, and then feeds on the dead prey (Burkholder et al. 1992; 1995; Steidinger el al. 1996). Taxonomic Description: Pfiesteria piscicida is a polymorphic and multiphasic dinotlagellate species with a number of unicellular stages throughout its life cycle: bi- and tri flagellated zoospores, and nonmotile cyst stages. Within the different life stage forms there is a wide range in size and morphology (Steidinger et al. 1996). The flagellated stages are small, oblong thecate cells that resemble gymnodinioid cells, although they are actually small cryptic peridinioid cells (Figs. 1-4). The bitlagellated stages, zoospores, have two size groups: 5-8 i^m (gametes) and 10-18 ^m (Fig. 3). The larger triflagellated stage, 25-60 |im, is a planozygote with the features of a vegetative eel! along with one transverse and two longitudinal flagella (Fig. 4). Cyst stages, with highly resistant cell walls, range in size from 25-33 ^im (Fig. 5). The flagellated forms are typically planktonic and ephemeral, whereas the cyst stages are benthic (Steidinger etal. 1996). Nomenclatiiral Types: Holotype: Pfiesteria piscicida Steidinger, Burkholder, Glasgow, Hobbs, Garrett, Truby, Noga and Smith, 1996: 160, fig. 2 Type Locality: North Atlantic Ocean: Pamlico River Estuary, North Carolina, USA Synonyms: Pfiesteria piscimorte Burkholder et al., 1993 Pfiesteria piscimortids Burkholder et al., 1995 "phantom dinotlagellate" Burkholder et al., 1992 Etymology: The genus 'Pfiesteria' is named in honor of Dr. Lois A. Pfiester, a pioneer in describing and unravelling the sexual life cycles of freshwater dinoflagellates. The species name 'piscicida ' is taken from the Latin words 'pisces ' for fish, and 'cida' for killer (Steidinger et al. 1996). Thecal Plate Description: The biflagellated stages off*, piscicida have thin thecal plates with a plate formula unique to the Dinophyceae: Po, cp, X, 4', la, 5", 6c, 4s, 5"', 2"" (Figs. 6-9). Raised sutures designate plate tabulation (Figs. 1.4). Thecal nodules border plate sutures (Fig. 6). Theca is smooth with scattered pores; trichocysts are present. The epitheca is equal to or exceeds the hypotheca in height (Fig. 1). The apical pore complex (APC) houses a broadly ovate apical pore plate (Po) and closing plate (cp)(Figs. 6-8). The elongate canal plate (x plate) is at a slight angle to the APC (Figs. 7,8). The first apical plate (!') is rhomboid in shape (Fig. 6). The broad and shallow cingulum is without lists, and descends almost 1 time its width. The sulcus is excavated, without lists, descends to the right, and slightly invades the epitheca via the anterior sulcal plate (s.a.)(Figs. 1,9) (Steidinger etal. 1996). Morphology and Structure: P. piscicida exhibits a number of different life cycle stages. This species uses both heterotrophic and mixotrophic nutritional modes depending on the life stage. Flagellated stages are mixotrophic: they use a peduncle (Figs. 1,2) to capture and ingest prey (myzocytosis), and kleptochloroplasts (chloroplasts retained from ingested algai prey) to photosynthesize when prey supply is low. Large food vacuoles are often found in the epitheca, the mesokaryotic nucleus is located in the hypotheca (Schnepf et al. 1989: Elbrachter 1991;' Fields & Rhodes 1991; Stoecker 1991; Steidinger etal. 1996; Lewitus et al. 1999). Reproduction: Biflagellated zoospores reproduce asexually via temporary cysts. Sexual reproduction has also been documented for this species: bitlagellated zoospores produce anisogamous gametes (Fig. 3), which fuse to produce triflagellated planozygotes (two longitudinal tlagella and one transverse) (Fig. 4). Sexual and asexual reproduction can occur on 54 Harmful Marine Dinoflagellates either a fish or algal diet (Tester, P.. personal communication). Species Coiiiparisoiis: P. piscicida is a distinct free-living estuarine dinoflagellate (Feiisome et al. 1993, Burkholder & Glasgow 1995; 1997). Ecology and Toxicity: P. piscicida is an estuarine species with a wide temperature and salinity tolerance. A cryptic heterotrophic species, it is a prey generalist that feeds on bacteria, algae, microtauna, fmfish and shellfish, and may well represent a significant estuarine microbial predator. Feeding mode is governed by the presence or absence of fish and fish material. Life cycle stage is governed by the presence of live or dead fish (Burkholder 1995; Burkholder & Glasgow 1997). In the absence of fish, bifiagellated stages feed myzocytotically on bacteria, algae and microfauna; i.e. prey is suctioned into a food vacuole via a feeding tube or peduncle (Fig. 2), and then digested (Burkholder & Glasgow 1995; Glasgow et al. 1998). Similar to other heterotrophic dinofiagellate species, a large food vacuole allows P. piscicida to phagocytize large prey items (Gaines & Elbrachter 1987; Schnepf & Elbrachter 1992; Burkholder et al. 1998). PJiesieria piscicida is a strong ichthyotoxic dinofiagellate species: in the presence of live fish, P. piscicida 's behavior is stimulated by a chemosensory cue, an unknown substance in fish secreta/excreta. Benthic stages (Fig. 5) then rapidly emerge as flagellated forms that swarm, immobilize, and kill the prey. Some prey experience ulcerative fish disease (open skin lesions) before dying. P. piscicida is lethal to fish at relatively low concentrations (> 250-300 cells/ml). At lower levels (-100-250 cells/ml) ulcerative fish disease results. Similar ulcers have been reported from shellfish as well. After a kill benthic stages form which inconspicuously descend back to the sediments (Burkholder & Glasgow 1995; 1997; Burkholder et al. 1995; 1998;Nogaetal. 1996; Steidinger et al. 1996). P. piscicida and possibly other PJies(eria-\ikQ species are suspected to be responsible for a number of major fish and shellfish kills in the North Carolina Albemarle-Pamlico estuary, and in the Maryland Chesapeake Bay (Burkholder et al. 1995; Burkholder & Glasgow 1997). The ever changing morphology of this species may give answers to a number of mysterious fish kills along the southeast coast of the United States (Steidinger etal. 1996). This species was initially linked to serious health problems in humans who had come in direct contact with it (narcosis, respiratory distress, epidermal lesions, and short-term memory loss); however, a study sponsored by the Centers for Disease Control (CDC) has revealed no such relationship (Swinker et al. 2001). Other CDC-funded studies are currently addressing possible associated human health problems with PJiesieria and P/iesteria-\\kc species in several states, including Marvland and North Carolina (P. Tester, personal communication). Habitat and Locality: PJiesieria piscicida was first identified from the Pamlico Sound in North Carolina. Since its emergence; however, P. piscicida and Pfiesteria-WkQ species have been reported from other eutrophic, temperate to subtropical estuarine systems in the eastern United States: from Delaware inland bays to Mobile Bay, Alabama (Burkholder et al. 1993; Burkholder etal. 1995; Lewitus et al. 1995). This natural range is expected to expand, considering the warming trend in global climate, and the increased human impact on coastal areas resulting in decreased water quality (Smayda 1992; Adler et al. 1993: Epstein et al. 1993; Hallegraeff 1993; Burkliolder & Glasgow 1997). Prorocentrum arenarium Faust, 1 994 Plate 37, Figs. 1-6 Species Overview: Prorocentrum arenarium is an armoured, marine, sand-dwelling, benthic dinofiagellate species. This toxic species is associated with coral rubble and colored sand in tropical cmbayments of the Caribbean Sea. Taxonomic Description: Prorocentrum arenarium is a bivalvate species often observed in valve view. Cells are round to slightly oval in valve view (Figs. 1,2,6); cell size ranges between 30 to 32 fjm in diameter. Both valves are concave in the center. The thecal surface is smooth (Figs. 1-3) with distinct randomly distributed valve poroids (65-73 per valve). The valve centers are devoid of pores. The poroids Harmful Marine Dinotlagellates 55 vary from kidney-shaped to oblong (Figs. 1-5), with an average size of 0.62 |im long and 0.36 ^m wide. Spacing between poroids is 2-3 fim. Valve margins exhibit evenly spaced marginal poroids, 50-57 per valve, and are similar in size to valve poroids (Figs. 1-5). These poroids are useful diagnostic features of this species and are easily viewed under the light microscope. The intercalary band is smooth and wide (Figs. 2,3)(Faust 1994). The peritlagellar area, which lacks ornamentation, is a broad triangle on the right valve at the anterior end of the cell (Figs. 1,3,5). The anterior region of the right valve is excavated; the left valve margin is flattened (Fig. 2). The flagellar and auxiliary pores are unequal in size (Fig. 5). The longitudinal flagellum is short (average length of 11 jam) (Fig. l)(Faust 1994). Nomenclatural Types: Holotype: Prorocentrum arenarium Faust, 1994: figs. 14, 15 Type Locality: Caribbean Sea: Carrie Bow Cay, Belize, Central America Morphology and Structure: Prorocentnim arenarium is a photosynilietic species with a prominent central pyrenoid and a posterior nucleus (Fig. 6). A small (2-3 fim), narrow, tubular, peduncle-like structure in the peritlagellar area has been observed in this species. This structure originates and emerges from the flagellar pore (Faust 1994). Reproduction: Prorocentrum arenarium reproduces asexually by binary fission. Ecology: Prorocentrum arenarium is a benthic and epiphytic species. Cells are motile, propelled by two flagella, or are attached to sand or coral rubble. This species can be a significant component of benthic Prorocentrum assemblages in colored sand patches in the Caribbean (1200- 6000 cells/g sand) (Faust 1994). The presence of a peduncle-like structure may indicate mixotrophic feeding within the sand (Faust 1994). Toxicity: This is a known diarrhetic shellfish poison (DSP) toxin-producing species, producing okadaic acid (OA)(Ten-Hage et al. 2000). Species Comparison: Only a few round to nearly round Prorocentrum species are known: P. arenarium (Faust, 1994) is smaller than P. emarginatum (cell diameter 35-40 nin)(Faust 1990b), but larger than P. ruetzlerianum (cell diameter 28-35 ^m) (Faust 1990b) and P. compressum (cell diameter 36 (im)(Matzenauer 1933; Bohm 1936; Schiller 1937; Tafall 1942; Dodge 1975). The valve poroids of P. arenarium are distinct from similarly known benthic Prorocentrum species: P. lima has approximately 58-86 round pores per valve and 55-72 marginal pores with a diameter of 0.3-0.7 |im (Faust 1991); P. maculosum has about 85-90 valve poroids and 65-75 marginal poroids with a diameter of 0.6 |im (Faust 1993b). The architecture of the periflagellar area off. arenarium, with no ornamentation (Faust 1994), is similar to that of P. concavum, P. ruetzlerianum (Faust 1990b), P. foraminosum (Faust 1993b), and P. tropicalis (Faust 1997). P. arenarium has a smooth intercalary band. This feature is also characteristic of other benthic Prorocentrum species: P. lima (Faust 1991), P. hoffmannianum (Faust 1990), and P. foraminosum (Faust 1993b). The peduncle-like organelle in P. arenarium is similar in structure to the peduncle observed in P. norrisianum (Faust 1997). Habitat and Locality: Prorocentrum arenarium is associated with coral rubble and colored sand in tropical embayments of the Caribbean Sea and the SW Indian Ocean (Faust 1994; Ten-Hage et al. 2000). Prorocentrum balticiim (Lohmann) Loeblich III, 1970 Plate 38, Figs. 1-4 Species Overview: Prorocentrum balticum is an armoured, marine, planktonic, bloom-forming dinotlagellate species. This cosmopolitan species is commonly found in cold temperate to tropical waters world-wide. Taxonomic Description: P. balticum is a bivalvate species often observed in valve view. Cells are small (< 20 |im in diameter), and round to ovoid in valve view (Figs. 1,2,4), with two 56 Harmful Marine Dinoflagellates minute and distinct apical projections (Figs. !,3,4). Although cells are nearly spherical, some have broad shoulders. Thecal valves are covered with many tiny interconnected spines (Figs. 1-4) which form narrow transverse rows on the intercalary band (Fig. 1). Many scattered rimmed pores are present on the cell surface (Fig. 2)(Dodge 1975; 1982; Toriumi 1980; Steidinger & Tangen 1996; Faust et al. 1999). Two minute apical spines (Figs. 1,3,4) border the relatively small periflagellar area. The peritlagellar pores are different sized (Fig. 3)(Dodge 1975; Toriumi 1980; Steidinger & Tangen 1996; Faust et al. 1999). Nomcnclatural Types: Holotype: ExuviaeUa baltica Lohmann, 1908: 265, plate 17, fig. la,b Type Locality: unknown Synonyms: Prorocentrum pomoideum Bursa, 1959 ExuviaeUa aequatorialis Hasle, 1960 Morphology and Structure: Prorocentrum bahicum is a photosynthetic species with a round nucleus situated posteriorly (Dodge 1975; Dodge 1982; Toriumi 1980). Reproduction: P. balticum reproduces asexually by binary fission. Ecology: P. balticum is a planktonic species. It is a neritic and oceanic species with world-wide distribution (Dodge 1975; Dodge 1982; Steidinger & Tangen 1996). Cells are active swimmers. This species has been reported to form red tides in many parts of the world (see Lassus 1988). Many blooms have occurred in brackish water habitats (Tangen 1980; Zotter 1979; Edler et al. 1984) confirming Braarud's (1951) earlier growth experiments that revealed P. balticum'?, highest growth rates were at low salinities (10-15 o/oo). Toxicity: Although toxicity in P. bahicum has never been confirmed, it has been associated with toxic red tides (Silva 1956; Silva 1963; Numann 1957). Steidinger (1979) regards it as a questionable toxic species. Species Comparison: P. balticum is not easily distinguished from P. minimum and a critical assessment of its taxonomic status is still needed. Both are small species, valves covered with small spines, and periflagellar areas are relatively small with two pores. P. balticum is distinguished by its small size, its almost spherical shape (Toriumi 1980), and its two minute apical projections (Faust etal. 1999). Because of its small size, records of P. bahicum may actually include closely related, but undescribed species (Steidinger & Tangen 1996). Habitat and Locality: Prorocentrum balticum is commonly found in marine waters all over the world: cosmopolitan in cold temperate to tropical waters (Dodge 1975; 1982; Steidinger & Tangen 1996). Prorocentrum belizeanitm Faust, 1993 Plate 39, Figs. 1-9 Species Overview: Prorocentrum belizeanum is an armoured, marine, benthic dinotlagellate species. This species is associated with floating detritus and sediment in tropical embayments of the Caribbean Sea. Taxonomic Description: Prorocentrum belizeanum is a bivalvate species often observed in valve view. Cells are round to slightly oval (Figs. 1,2,4,7,8). Cells measure between 55-60 ^m in length and 50-55 pm in width. Valves are concave in the center (Figs. 2,4) (Faust 1 993a). Thecal surface is heavily areolated; approximately 853-1024 areola are present on each valve (Figs. 1-5). The areolae are round to oval (0.66-0.83 ^im in diameter) (Figs. 1-6). Some bear trichocyst pores at their base. Ejected trichocysts are often observed. The intercalary band is smooth; however, marginal areolae give the appearance of a transversely striated intercalary band (Figs. 7,8)(Faust 1993a). The peritlagellar area is a V-shaped triangle located apically on the right valve (Figs. 1,4,6,8). Both the left and right valves are excavated (Figs. 1,4), Two peritlagellar pores, flagellar and auxiliary, are equal in size. The auxiliary pore is surrounded by a flared periflagellar collar (Fig. Harmful Marine Dinoflagellates 57 6). Accessory pores are also present. The left valve anterior margin bears a large rounded and flared curved apical collar that borders the periflagellar area (Figs. 1-4,6,8). In lateral and apical view, the curved apical collar resembles a rounded lip (Figs. 3,4)(Faust 1993a). Nomenclatural Types: Holotype: Prorocentrum belizeanum Faust, 1993: tigs. 1,2 Type Locality: Caribbean Sea: Twin Cays, Belize, Central America Morphology and Structure: Prorocentrum belizeanum is a photosynthetic species with a centrally located pyrenoid and a large kidney- shaped posterior nucleus (Fig. 7)(Faust 1993a). Reproduction: Prorocentrum belizeanum reproduce asexually by binar>' fission. Ecology: P. belizeanum is a benthic species that can be a major component (1200 cells/mL) of benthic Prorocentrum assemblages in floating detritus and sediments in tropical coastal waters of the Caribbean, Cells are motile or are often attached to sediments and detrital particles (Faust 1993a). Toxicity: This is a known diarrhetic shellfish poison (DSP) toxin-producing species producing okadaic acid (OA) and small amounts of Dinophysistoxin-1 (DTXl)(Morton et al. !998). Species Comparison: Only a few round or near- round Prorocentrum species are known: P. belizeanum is larger then P. hojfmanniamim (45- 55 ^m long and 40-45 jim wide)(Faust 1990b) and larger than P. compressum (36 |im in diameter)(Matzenauer 1933; Bohm 1936; Schiller 1937; Tafall 1942; Dodge 1975). The areolae of P. belizeanum are distinct from similar known benthic Prorocentrum species (Faust 1993a): P. hojfmannianum has approximately 670 areola per valve (diam.= 1.0- 1.15 |im), and P. ruetzlerianum has about 550 pentagonal-shaped areola per valve (diam.= 1.0 ^m)(Faust 1990b). The architecture of the periflagellar area of P. belizeanum is similar to P. lima (Taylor 1980) and the planktonic species P. playfairi (Croome & Tyler 1987). P. hojfmannianum (Faust 1990b), however, has a more complex platelet configuration (Faust 1993a). The periflagellar area of P. belizeanum lacks an apical spine (Faust 1993a), which is similar to P. hoffmannianum (Faust 1990b) and P. lima (Faust 1991), but different from P. compressum, which has two apical spines (Tafall 1942; Dodge 1975). P. reticulatum (Faust 1997), P. sabulosum (Faust 1994), P. belizeanum (Faust 1993a) and P. hojfmannianum (Faust 1990b) share a distinct feature in the periflagellar area: three small accessory pores adjacent to a periflagellar pore (Faust 1997). The flared curved apical collar (or 'raised anterior ridge') on the left anterior margin of P. belizeanum is similar to the curved apical collar of P. hoffmannianum. However, P. belizeanum has a more prominent and rounder collar than P. hojfmannianum, which is broader (Faust 1990b; Faust 1993a; Steidinger & Tangen 1996). Habitat and Locality: Cells of P. belizeanum are common in tropical coastal waters (Steidinger & Tangen 1996) associated with floating detritus (Faust 1993a). Prorocentrum concavum Fukuyo, 1981 Plate 40, Figs. 1-7 Species Overview: Prorocentrum concavum is an armoured, marine, benthic dinollagellate. This toxic species is often associated with floating detritus and sediments in tropical and neritic waters. Taxonomic Description: P. concavum is a bivalvate species often observed in valve view. Cells are broadly ovoid. Valve centers are concave and flattened (Figs. 1,2.5-7). Cells measure 50-55 |am in length and 38-45 |im in width. The valve surface is covered with 1000- 1 100 prominent shallow areolae. The areolae are round to oval with smooth edges (Figs. 1,3) and often observed with a central pore (0.8 jim diameter) (Fig. 3). No marginal pores are present and the cell center is devoid of areolae (Fig. 5). The intercalary band is granulated and horizontally striated (Figs. l,2)(Fukuyo 1981; Faust 1990b). 58 Harmful Marine Dinoflagellates The periflagellar area is a narrow, rimmed, V- shaped depression on the right valve (Figs. 1,4,5,7). It is composed of eight apical plates, without ornamentation, fitted with a large flagellar pore, and a much smaller auxiliary pore (Fig. 4). The left valve is slightly indented anteriorly with a thickened apical ridge (raised margin) bordering the periflagellar area (Fig. l)(Fukuyo 1981; Faust 1990b). ^ P. concavttm and P. tropicalis (Faust 1997) have similar intercalary bands: granulated and horizontally striated. Habitat and Locality: P. concavum populations are often associated with floating mangrove detritus and sediments in tropical and neritic waters (Faust 1990b; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Prurocentnmi concavum Fukuyo, 1981: figs. 13-19,49 Type Locality: Pacific Ocean: French Polynesia, New Caledonia and the Ryukyu Islands Morphology and Structure: Prorocentrum concavum is a photosynthetic species with golden-brown chloroplasts (Faust 1990b). Two cup-shaped pyrenoids are also present (Fukuyo 1981). Reproduction: Prorocentrum concavum reproduces asexually by binary fission. Ecology: P. concavum is a benthic species that can also be tycoplanktonic. Cells can be either motile or embedded in mucus attached to detritus (Faust 1990b; Steidinger & Tangen 1996). Toxicity: This species is known to be toxic, producing the following toxins: fast-acting toxin (FAT)(Tindall et al. 1984), diarrhetic shellfish poison (DSP) toxins (Hu et al. 1993), okadaic acid (OA)(Dickey et al. 1990), and an unnamed toxin (Tindalletal. 1989). Species Comparisons: Prorocentrum concavum, at the LM level, is difficult to differentiate from a number of other Prorocentrum species due to their similar size and shape; e.g. P. concavum is often confused with P. lima (Fukuyo 1981; Faust 1990b), but P. lima is not areolate and bears marginal pores (Faust 1990b). The location and arrangement of areolae on the surface of P. concavum closely resembles that of P. hoffmannianum (about 670/valve)(Faust 1990b) and P. belizeanum (about 950/valve) (Faust 1993a); however, the latter two species have fewer areolae per valve and also have marginal pores, while P. concavum does not (Faust 1990b). Prorocentrumfaustiae Morton, 1998 Plate 41. Figs. 1-4 Species Overview: Prorocentrum faustiae is an armoured, marine, benthic dinofiagellate species. This species is associated with macroalge from the Australian Barrier Reef. Taxonomic Description: Prorocentum faustiae is a bivalvate species often observed in valve view. Cells are broadly ovate to rotundate with a rugose appearance (Figs. 1-3). Valve centers are concave (Figs. 1-3). Cells are 43-49 ^m long and 38-42 |.im wide. Small pores (0.1 fim in diameter), probably containing trichocysts, are dense on the valve surface and along the valve perifery (Figs. 1-3). The intercalary band is transversely striated (Fig. 3) (Morton 1998). The perifiagellar area is a wide triangular, V- shaped region located apically on the right valve (Figs. 1,4). Sixteen apical platelets make up the periflagellar area. Included also are two pores: a large flagellar pore, and a smaller auxiliary pore (Fig. 4)(Morton 1998). Nomenclatural Types: Holotype: Prorocentrum faustiae Morton, 567, figs. 1-4 1998: Type Locality: Australia Coral Sea: Heron Island, Morphology and Structure: Prorocentrum faustiae is a photosynthetic species containing numerous golden-brown chloroplasts and a centrally located pyrenoid (Figs. 1,2). A large kidney-shaped nucleus is situated posteriorly (Morton 1998). Reproduction: Prorocentrum reproduces asexually by binary fission. faustiae Harmful Marine Dinoflagellates 59 Ecology: Prorocentrum fansliae is a benthic species epiphytic on macroatgae (Morton 1998). Toxicity: P. faustiae is a diarrhetic sheilfisli poison (DSP) toxin-producing species producing okadaic acid (OA) and Dinophysistoxin-1 (DTXl)(Morton 1998). Species Comparison: Prorocentrum faustiae is similar in shape and size to P. hoffmannianum (45-55 |Lim long and 40-45 |im wide); however, the former lacks thecal areolae, which are very abundant on the latter. P. faustiae lacks a distinct ridge along the valve perifery which distinguishes this species from P. maculosum (Morton 1998). Etymology: The species 'faustiae' is named in honor of Dr. Maria Faust, Smithsonian Institution, for her advancements in the taxonomy o'i non-planktonic dinoflagellates (Morton 1998). Hubitiit and Locality: Populations of P. faustiae are associated with macroalgae from Heron Island, Australia (Morton 1998). Prorocentrum hoffmannianum Faust, 1990 Plate 42, Figs. 1-6 Species Overview: Prorocentrum hoffmannianum is an armoured, marine, bentliic dinotlagellate species. This species is associated with floating detritus and sediment in tropical embayments of the Caribbean Sea. Taxonomic Description: Prorocentrum hoffmannianum is a bivalvate species often observed in valve view. Cells are ovoid, broadest in mid-region, tapering slightly apically (Figs. 1,2,5,6). Cells are 45-55 |im long and 40- 45 |im wide. Both valves are slightly concave in the center. The intercalary band is smooth and appears as a tlared ridge around the cell (Figs. 1,2,5). Observed under LM, the marginal areolae can give the appearance of a striated intercalary band (Fig. 5)(Faust 1990b). The valve surface is deeply areolate; areolae are dense, large, and round to oblong (Figs. 1-4). Small round to ovoid pores are found within deep areolae; these pores have smooth margins, are 1.0-1.5 ^m in diameter, and many bear trichocyst pores (Fig. 3). There are approximately 650-700 areolae on each valve (Faust 1990b). The periflagellar area is a wide triangle situated apically on the right valve (Figs. 1,4). It houses eight periflagellar platelets and two periflagellar pores: a flagellar pore and auxiliary pore (equal in size); accessory pores are also present. The flagellar pore is surrounded by a small flared periflagellar collar (Fig. 4). Both left and right valves are apically excavated (Figs. 1,4). The left valve exhibits a flared and flattened curved apical collar that borders the periflagellar area (Figs. l,2)(Faust 1990b). Nomenclatural Types: Holotype: Prorocentrum hoffmannianum Faust, 1990: figs. 13,14 Type Locality: Caribbean Sea: Twin Cays, Belize, Central America Synonyms: Exuviaella hoffmannianum (Faust) McLachlan, Boalch and Jahn, 1997 Morphology and Structure: Prorocentrum hoffmannianum is a photosynthetic species containing golden-brown chloroplasts, a centrally located pyrenoid, and a large posterior nucleus (Fig. 5)(Faust 1990b). Reproduction: Prorocentrum hoffmannianum reproduces asexually by binary fission. Ecology: Prorocentrum hoffmannianum is a benthic species. Cells are motile or attached to detritus by mucilage (Faust 1990b). Toxicity: This species is considered toxic producing fast-acting toxin (FAT) and diarrhetic shellfish poison (DSP) toxin: okadaic acid (OA)(Aikman et al. 1993). Species Comparison: Prorocentrum hoffmannianum is similar in shape to P. lima, but larger and broader with dense areolae. P. hoffmannianum is often misidentified as P. concavum, but can be distinguished by its ovoid shape and presence of areolae in the center of the valve (Fukuyo 1981; Faust 1990b; 1991). 60 Harmful Marine Dinoflagellates The architecture of the periflagellar area of P. hojfmanniamim is similar to P. lima, P. concavum (Fukuyo 1981) and P. playfairi (Croome & Tyler 1987); however, P. hoffmannianum has a more complex platelet configuration (Faust 1990b). P. reticulatitm (Faust 1997), P. sabulosum (Faust 1994), P. belizeanum (Faust 1993a) and P. hoffmannianum (Faust 1990b) share a distinct feature in the peritlagellar area: three small accessory pores adjacent to a peritlagellar pore (Faust 1997). Both P. hqjfnuinnianitm and P. belizeanum have a prominent flared curved apical collar on the left valve bordering the peritlagellar area, although the curved apical collar of the latter species is rounder, whereas that of the former is flatter (Faust 1993a). Etymology: This species is named in honor of Dr. Robert S. Hoffmann, Assistant Secretary for Research, Smithsonian Institution, for his encouragement, support and scientific leadership (Faust 1990b). Habitat and Locality: Populations of P. hoffnianniainim are often associated with floating detritus in tropical coastal regions of the Caribbean Sea (Faust 1990b). Remarks: In Carlson (1984), P. concavum identified on Plate 5, figs, n-s, is P. hoffmannianum based on thecal surface morphology, periflagellar area and intercalary band characteristics. In addition, the illustration of P. concavum (fig. 17) by Steidinger (1983) is neither P. concavum nor P. hoffmannianum, but is an unidentified species (Faust 1990b}. Prorocentruni lima (Ehrenberg) Dodge, 1975 Plate 43, Figs. 1-7 Species Overview: Prorocentrum lima is an armoured, marine, benthic dinoflagellate species with world-wide distribution. Taxoiioinic Description: P. lima is a bivalvate species often observed in valve view. Cells are oblong to ovate, small to medium-sized, broadest in the mid-region, and narrow toward the anterior end (Figs. 1,2,4-6). Cell size ranges between 32- 50 |im in length and 20-28 \xm in width. Thecal valves are thick and smooth with scattered surface pores (Figs. 1-4). Each valve contains about 50-80 small round marginal pores evenly spaced around the perifery of the valve (0.6 |im in diameter)(Figs. 1,3), and about 60-100 larger round to oblong unevenly distributed valve pores with trichocysts (0.48 |.im in diameter) (Figs. 1,2,4). All pores have smooth edges (Figs. 3,4). The center is devoid of pores (Figs. 1,2,4). Marginal pores is a useful diagnostic feature of this species distinguishing it from other Prorocentrum species. Occasionally P. lima can be found without marginal pores or with partially filled pores. In older cells, the thecal surface can become vermiculate. The intercalary band appears as a thick, smooth, and well-defined margin at the periphery of the valve giving the appearance of a flared ridge (Figs. 1,2,4-6) (von Stosch 1980; Dodge 1975; Faust 1990b; Faust 1991; Steidinger & Tangen 1996). The periflagellar area is a shallow V-shaped depression on the right valve (Fig. 3) made up of eight platelets and two pores: a larger flagellar pore and a smaller auxiliary pore (Figs. 1,3-5). A protruding periflagellar collar surrounds the auxiliary pore (Fig. 3). Both valves are anteriorly indented; the left valve margin has a flattened apical ridge that borders the periflagellar area (Figs. l,2,6)(Faust 1991; Steidinger & Tangen 1996). Nomenclatural Types: Holot\pe: Prorocentrum lima (Ehrenberg) Dodge, 1975: 109, figs. 1E,F, plate IB.C Type Locality: unknown Synonyms; Exuviaella marina Cienkowski, 1 88 1 Exuviaella lima (Ehrenberg) Butschli, 1885 Exuviaella marina var. lima (Ehrenberg) Schiller 1933 Basionym: Cryptomonas lima Ehrenberg, 1 860 Morphology and Structure: Prorocentrum lima is a photosynthetic species containing two chloroplasts, a central pyrenoid and a large posterior nucleus (Figs. 5,6)(Dodge 1975). Reproduction: P. lima reproduces asexually by binary fission. This species also exhibits an alternate form of asexual reproduction in which a chain of cell pairs is enclosed within a thin- Harmful Marine Dinollaszellates 61 walled cyst. In this mode multiple vegetative divisions occur within a hyaline envelope (a division cyst) which may contain a chain of 4 to 32 cells (Faust 1993d). Sexual reproduction has also been documented: isogamous gametes form, conjugation takes place, and a large hypnozygote (resting cyst) is produced (Fig. 7)(Faust 1993c). Ecology: P. lima is a benthic and epiphytic species that can be tycoplanktonic. Cultured cells readily adhere to the culturing vessel via mucous strands and rarely swim freely (Fukuyo 1981; Steidinger & Tangen 1996). This species produces a pale colored resting cyst as part of its life cycle. Cysts are large (TO- TS jam diameter) and round with a smooth triple- layered wall (Faust 1993c). Toxicity: Prorocentrum lima is a toxic dinoflagellate species known to produce a number of toxic substances: fast-acting toxin (FAT)(Tindall et al. 1989); prorocentrolide (Torigoe et al. 1988); and diarrhetic shellfish poison (DSP) toxins (Yasumoto et al. 1987): okadaic acid (OA)(Murakami et al. 1982; Lee et al. 1989; Marr et al. 1992); Dinophysistoxin-I (DTXl)(Marr et al. 1992); Dinophysistoxin-2 (DTX2)(Hu et al. 1993); and Dinophysistoxin-4 (DTX4)(Hu et al. 1995). Species Comparison: P. lima is difficult to identify due to its similar morphology to several other Prorocentrum species with a triangular periflagellar area and an oval or ovoid shape (e.g. P. foraminosum, P. concavum and P. hoffmanniamim). P. lima can be distinguished by its size, shape, narrow periflagellar area and the presence of valve and marginal pores. P. concavum, however, is larger, broader, has more valve pores and does not have marginal pores. P. foraminosum and P. hoffmanniamim are also similar in shape to P. lima, though both are larger species with very different valve pore numbers and arrangements. P. hoffmannianum, moreover, is much broader and its valve surface is deeply areolated (Steidinger 1983; Steidinger & Tangen 1985; 1996; Fukuyo 1981; Faust 1990b; 1991; 1993b). Steidinger (1983) recognized that the marginal pores of P. lima can be used to differentiate this species at the light microscope level from completely areolated species such as P. concavum or P. compressum which are similar in shape. Habitat and Locality: Prorocentrum lima is a neritic, estuarine species with world-wide distribution (Steidinger & Tangen 1996). Cells can be found in temperate (Lebour 1925; Schiller 1933; Carter 1938) as well as tropical oceans (Fukuyo 1981; Steidinger 1983; Carlson 1984; Faust 1990b). This species occurs in sand (Lebour 1925; Drebes 1974; Dodge 1985), attached to the surface of red and brown algae and benthic debris (Fukuyo 1981; Steidinger 1983; Carlson 1984), associated with coral reefs (Yasumoto et al. 1980; Fukuyo 1981; Bomber et al. 1985; Carlson & Tindall 1985), or can be found attached to floating detritus in mangrove habitats (Faust 1991). Prorocentnim nuiciilosiim Faust, 1993 Plate 44, Figs. 1-6 Species Overview: Prorocentrum maculosum is an armoured, marine, benthic dinoflagellate species. This toxic species is often associated with floating detritus in tropical coastal regions of the Caribbean Sea. Taxonomic Description: Prorocentrum maculosum is a bivalvate species often observed in valve view. Cells are 40-50 jim long and 30- 40 |am wide, broadly ovate with the maximum width behind the middle region, and slightly tapered at the anterior end (Figs. 1,2). The thecal surface is rugose with distinct scattered valve poroids (85-90 per valve)(Figs. 1-3). The poroids vary from kidney-shaped to circular or oblong (average diam.=6.0 ^m), 2-4 \im apart (Fig. 3). Valve center is devoid of poroids (Figs. l,2,6)(Faust 1993b). The valve margins form a distinct ridge which appears as a flange around the cell (Figs. 1,2). Marginal pores are equally spaced (65-75 per valve), and appear larger and more uniform than the valve poroids (Figs. l,2)(Faust 1993b). The periflagellar area is a broad triangle on the anterior end of the right valve (Figs. 1,4) made up of 8 platelets and 2 pores (Fig. 4). A thin apical ridge (raised margin) on the left valve surrounds the periflagellar area (Figs. 2,4). The 62 Harmful Marine Dinoflagellates flagellar and auxiliary pores are about equal in size, both surrounded by a curved and flared periilagellar collar (Fig. 4)(Faust 1993b). Noinenclatural Types: Holotype: Prorocentrum maculosum Faust, 1993: figs. 1,2 Type Locality; Caribbean Sea: Twin Cays, Belize, Central America Synonyms: Exuviaella maculosum (Faust) McLachlan, Boalch and Jahn, 1997 Morphology and Structure: Prorocentrum maculosum is a photosynthetic species containing golden-brown chloroplasts and a centrally located pyrenoid. A large posterior nucleus is situated adjacent to the pyrenoid (Fig. 5)(Faust 1993b). Reproduction; Prorocentrum maculosum reproduces asexually by binary fission. Ecology: P. maculosum is a benthic species. Cells are motile or attach to detritus or sediment by mucous strands (Faust 1993b). Toxicity: This is a known toxic species that produces prorocentrolide B, a fast-acting toxin (Hu et al. 1996). A diarrhetic shellfish poison (DSP) toxin, okadaic acid (OA), has also been reported from one Caribbean clone previously identified as P. concavum (Dickey et al. 1990), but reassigned to P. maculosum (Faust 1996b; Zhou & Fritz 1996). Species Comparison: The use of scanning electron microscopy has revealed major differences in the micromorphology of benthic species within the genus Prorocentrum (Faust 1990a; Faust 1993b). Under LM P. maculosum may be confused with P. lima (Faust 1991) which has round valve pores and a smooth thecal surface. Dodge (1975), when revising the taxonomy of the genus Prorocentrum, described P. lima to be a morphologically variable species. However, the architecture of the flagellar pore area was not considered. P. maculosum and P. lima can be distinguished by a number of surface features. The thecal surface of P. maculosum is rugose, covered with large kidney-shaped poroids; a periflagellar collar surrounds both equally-sized flagellar and auxiliary pores (Faust 1993b). \n P. lima the thecal surface is smooth with round pores; only the larger flagellar pore is surrounded by a curved periflagellar collar (Faust 1 99 1 ). The valve margins of P. tropicalis form a distinct ridge that appears as a flange around the cell, similar to P. maculosum (Faust 1993b), The periflagellar architecture of P. maculosum is similar to P. hojfmannianum (Faust 1990b), P. compressum (Abe 1967; Dodge 1975), P. playfairi and P. foveolata (Croome & Tyler 1987). Etymology: The name 'maculosum' originates from Latin and refers to 'speckled, spotted', which describes the thecal surface of this species (Faust 1993b). Habitat and Locality; Populations of P. maculosum are often associated with floating detritus in tropical coastal regions of the Caribbean Sea (Faust 1993b). Prorocentrum mexicanum Tafall, 1942 Plate 45, Figs. 1-7 Species Overview: Prorocentrum mexicanum is an armoured, marine, benthic dinoflagellate species. This toxic species is commonly found in tropical shallow embayments. Taxonomic Description: Prorocentrum mexicanum is a bivalvate species often observed in valve view. Cells are ovate to oblong with straight sides (30-38 \xm long and 20-25 |am wide) (Figs. 1,2,6). The valve surface of young cells is smooth (Fig. 2), but in older cells it may appear rugose (Figs. 1,3,5). Both valves have many large trichocyst pores (100 per valve) radially arranged in furrowed depressions (Figs. 1-5), and 80 marginal pores (Fig. 3). Trichocyst pores are round with a smooth edge (0.5 |um in diameter) and even in size (Fig. 4). Ejected trichocysts are common. Valve center devoid of pores. The intercalary band is broad and transversely striated (Figs. 3,5)(Faust 1990b). The periflagellar area, located apically and off-center on the right valve, is a relatively small, V-shaped, shallow depression (Figs. 1,5). It Harmful Marine Dinotlagellates 63 houses a prominent curved periflagellar collar adjacent to the auxiliary pore (Figs. 1,2,5). Opposite is a smaller peritlagellar plate adjacent to the flagellar pore {Fig. 5). The large peritlagellar collar (2X6 jim) may appear as an apical spine, and has been reported as such (Fukuyo 1981; Carlson 1984). Both valves are excavated (Figs. l,2)(Faust 1990b). Nomenclatural Types: Holotype: Prorocentrum mexicanum Tafall, 1942:plate34, figs. 3,8 Type Locality: North Pacific Ocean: Mexico Synonyms: Prorocentrum maximum Schiller, 1937 Prorocentrum rhathymum Loeblich, Sherley and Schmidt, 1979 Morphology and Structure: P. mexicanum is a photosynthetic species with a posterior nucleus (Faust 1990b). Reproduction: Prorocentrum mexicanum reproduces asexually by binary fission. Sexual reproduction has also been observed in natural cell populations (Faust, M.A., pers. com.). Ecology: P. mexicanum is a benthic species that can be tycoplanktonic (Steidinger & Tangen 1996). Cells swim freely or attach to floating detritus with mucous strands. Cells are often found embedded in mucilage (Faust 1990b). Toxicity: P. mexicanum is a known toxin- producing species (Steidinger 1983; Carlson 1984; Tindall et al. 1984) producing fast-acting toxin (FAT)(Tindall et al. 1984). Species Comparison: With its prominent peritlagellar collar, P. mexicanum most resembles P. caribbaeum in general cell shape; however, P. caribbaeum is a larger species, is broader and heart-shaped, and broadest in the anterior region (Dodge 1975; Faust 1993a). Trichocyst pore morphology is also similar in these two species; however, significant differences lie in the number of trichocyst pores: P. caribbaeum has a greater number of pores per valve (145-203) than P. mexicanum (100 per valve). Ejected trichocysts are often observed in cells of both species (Faust 1990b; 1993a). P. mexicanum, P. emarginatum and P. caribbaeum all have radially arranged valve pores and display two different sized pores (Loeblich et al. 1979; Fukuyo 1981; Steidinger 1983; Faust 1990b; 1993a). The peritlagellar area and platelet architecture of P. caribbaeum is similar to that of P, mexicanum (Carlson 1984; Faust 1993a). The intercalary band of P. mexicanum is transversely striated. This is similar to P. caribbaeum and P. emarginatum (Faust 1990b; 1993a). Habitat and Locality: Prorocentrum mexicanum is a common species found in tropical and subtropical benthic communities (Steidinger & Tangen 1996) of shallow protected areas of the Pacific and Atlantic Oceans (Faust 1990b). Prorocentrum micans Ehrenberg, 1833 Plate 46, Figs. 1-6 Species Overview: Prorocentrum micans is an armoured, marine, planktonic, bloom-forming dinotlagellate. This is a cosmopolitan species in cold temperate to tropical waters. Taxononiic Description: P. micans is a bivalvate species often observed in valve view. Cells of this species are highly variable in shape and size (Figs. l-5)(see Bursa 1959; Dodge 1975). Cells are tear-drop to heart shaped, rounded anteriorly, pointed posteriorly, and broadest around the middle (Figs. 1,2,4-6). This species is strongly flattened with a well- developed winged apical spine (10 (jm long) on the left valve (Figs. 1,3). Cells are medium-sized (35-70 jim long, 20-50 ^m wide) with a length:width ratio usually less than two. The cell surface is rugose, covered with shallow minute depressions (Figs. 1,2). Numerous tubular trichocyst pores are also present in short rows arranged radially (Figs. 1,5,6). Intercalary band is smooth and wide (Figs. l,4-6)(Wood 1954; Toriumi 1980; Dodge 1975; 1982; 1985; Fukuyo et al. 1990; Steidinger & Tangen 1996; Faust et al. 1999). The peritlagellar area is a relatively small, shallow, broad triangular depression situated 64 Harmful Marine Dinoflagellates apically on the right valve off-center (Fig. 3). Two perifiageilar pores are present: one large flagellar pore and one smaller auxiliary' pore (Fig. 3). Adjacent to the flagellar pore is a small, slightly curved peritlagellar plate (Fig, 3). The large pointed apical spine lies adjacent to the peritlagellar area, directly opposite the peritlagellar plate (Fig. 3)(Taylor 1980; Toriumi 1980). Nomenclatural Types: Holotype: Prorocentnim micans Ehrenberg, 1834:307 Type Locality: North Sea: near Kiel, Berlin, Germany Synonyms: Cerc^na sp. Michaeiis, 1830 Prorocentnim schilleri Bohm in Schiller, 1933 Prorocentnim levantinoides Bursa, 1959 Prorocentnim pacificiim Wood, 1 963 IVlorphology and Structure: P. micans is a photosynthetic species with two golden-brown chloroplasts situated peripherally. A large kidney-shaped nucleus is situated posteriorly. Two anterior vacuoles are usually present (Dodge 1975; 1982; Toriumi 1980; Fukuyo et al. 1990). Reproduction: P. micans reproduces asexually by binary fission. Ecology: P. micans is one of the most common and diversified species in the genus Prorocentnim. It is a planktonic species commonly found in neritic and estuarine waters, but it is also found in oceanic environments; it is cosmopolitan in cold temperate to tropical waters. This species is also known to tolerate very high salinity: populations have been reported from hypersaline salt lagoons (>90 o/oo) in the Caribbean islands (Steidinger & Tangen 1996). Cells are active swimmers (Dodge 1982; Steidinger & Tangen 1996). This species forms extensive red tides in many parts of the world (Fukuyo et al. 1990; 1999). Toxicity: Although P. micans is capable of forming extensive blooms, it is usually considered harmless (see Taylor & Seliger 1979; Anderson et al. 1985; Graneli et al. 1990). It may excrete substances that inhibit diatom growth, but apparently these substances do not enter the food chain or affect organisms at higher trophic levels (Uchida 1977). There are only a few reports of P. micans having caused problems: shellfish kills in Portugal (Pinto & Silva 1956) and South Africa (Horstman 1981). Claims for toxicity of this species need confirmation. Early reports on P. micans being a paralytic shellfish poison (PSP) producer (Pinto & Silva 1956) are unconfirmed, and recent incidents involving shellfish mortality have been attributed to oxygen depletion (Lassus & Benhome 1988). Species Comparison: This species varies considerably in shape and size and may be confused with closely related species; e.g. P. gracile, P. sciitelliim and P. caribbaeiim. P. gracile has a very strong winged apical spine, is not as broad, and has a length:width ratio usually larger than 2; P. scutellum is in the same size range as P. micans, but bears a shorter and broader apical spine (Dodge 1975; 1982). P. caribbaeum is also in the same size range, but is heart-shaped and broadest around the anterior end, whereas P. micans is more tear-drop shaped and broadest around the middle (Dodge 1985; Faust 1993a). P. gracile and P. micans share two distinct features: a.) similar trichocyst pore pattern (Steidinger & Williams 1970; Steidinger & Tangen 1996); and b.) similar arrangement of apical spine: the spines lie adjacent to the peritlagellar area (Toriumi 1980). Trichocyst pore number is highly variable in this species (Dodge 1985): 83 pores per valve were illustrated for one P. micans specimen (Dodge 1965), 101 pores per valve for another specimen (Dodge 1985), and 139 pores per valve in yet another specimen (Sournia 1986). Trichocyst pore morphology of this species resembles that of P. caribbaeum; however, the latter species has a much greater number of pores per valve: 145-203 (Faust 1993a). Habitat and Locality: P. micans is commonly found in marine waters all over the world (Dodge 1975). Harmful Marine Dinoflagellates 65 Prorocentritm minimum (Pavillard) Schiller, 1933 Plate 47, Figs. 1-7 Species Overview: Prorocentrum minimum is an armoured, marine, planktonic, bloom-forming dinoflagellate. It is a toxic cosmopolitan species common in cold temperate brackish waters to tropical regions. Taxonomic Description: Prorocentrum minimum is a bivalvate species often observed in valve view. Cells are small (14-22 jim long to 10-15 ^m wide) and shape is variable: cells range from triangular (Fig. 1), to oval (Figs. 3,5,7), to heart-shaped (Fig. 6). Cells are laterally flattened (Fig. 3). A short apical spine is sometimes observable (Figs. 1-4,7). Valves with short, evenly shaped broad-based spines (about 600-700 per valve) arranged in a regular pattern (Figs. 1-4). These can appear as rounded papillae depending on angle of view. There are two sized pores present: smaller pores are scattered (Figs. 1,4), while larger pores are located at the base of some peripheral spines. The intercalary band is transversely striated (Figs. 2.5,6) (Parke & Ballantine 1957; Faust 1974; Dodge 1982; Steidinger & Tangen 1996). The broad anterior end is truncate with a relatively small, shallow, broadly V-shaped depressed peritlagellar area located apically on the right valve, slightly off-center (Figs. 1-7). The periflagellar area bears eight apical platelets and two pores of unequal size: a large flagellar pore and a smaller auxiliary pore (Fig. 2). Adjacent to the flagellar pore is a small apical spine (Figs. 2,7). Adjacent to the auxiliary pore is a small, curved and forked peritlagellar collar (Figs. 1,2) (Parke & Ballantine 1957; Dodge & Bibby 1973; Faust 1974). Nomenclatural Types: Holotype: Exuviaella minima Schiller, 1933; figs. 33a,b Type Locality: Mediterranean Sea: Gulf of Lion, France Synonyms: Exuviaella minima Pavillard, 1916 Prorocentrum trianguiatum Martin, 1929 Exuviaella marie-lebouriae Parke and Ballantine, 1957 Prorocentrum cordiformis Bursa, 1959 Prorocentrum mariae-lebouriae (Parke and Ballantine, 1957) Loeblich III, 1970 Morphology and Structure: Prorocentrum minimum is a photosynthetic species with golden-brown chloroplasts, one large pyrenoid and two pusules. The nucleus is broadly ellipsoidal and posteriorly situated (Parke & Ballamine 1957; Faust 1974; Dodge 1982). Reproduction: P. minimum reproduces asexually by binary fission. Ecology: P. minimum is a bloom-forming planktonic species. Cosmopolitan in cold temperate brackish waters to tropical regions; mostly estuarine, but also neritic (Steidinger & Tangen 1996; Faust et al. 1999). Due to its small size, this species is probably often lost or overlooked in field samples (Dodge. 1982). Ceils are active swimmers (Parke & Ballantine 1957). Recently, Stoecker et al. (1997) reported mixotrophy in this species; ingested cryptophytes were observed in cells of P. minimum. Toxicity: P. minimum is a toxic species; it produces venerupin (hepatotoxin) which has caused shellfish poisoning resulting in gastrointestinal illnesses in humans and a number of deaths. This species is also responsible for shellfish kills in Japan and the Gulf of Mexico, Florida (Nakazima 1965; Nakazima 1968; Smith 1975; Okaichi & Imatomi 1979; Tangen 1983; Shimizu 1987; Steidinger & Tangen 1996). Species Comparisons: P. minimum can be confused with P. balticum; however, the former species differs by its larger size and different shape, and by having only one apical spine and a forked perifiagellar collar (Faust et al. 1999). Habitat and Locality: P. minimum is commonly found along the west coast of the USA, Japan, Gulf of Mexico, Caspian, Adriatic, Mediterranean and Black Seas, and Scandinavian waters; often in large numbers (Dodge 1982; Tangen 1980; 1983; Marasovic et al. 1990). 66 Hamillil Marine Dinoflagellates Prorocentrum ruetzlerinnum Faust, 1990 Plate 48, Figs. 1-6 Species Overview: Prorocentrum ruetzlerianuin is an armoured, marine, benthic dinotlagellate species. This species is associated with floating detritus and sediment in tropical embayments of the Caribbean Sea. Taxoiiomic Description: P. ruetzleriamun is a bivalvate species often observed in valve view. Cells are round to ovoid (Figs. 1,4-6) with an average diameter of 28-35 |im. Valve centers are slightly concave {Fig. 1). The entire valve surface is deeply areolate; the areolae are ovate to pentagonal deep depressions (Figs. 1,2,6). Each areola houses a central round pore (1 |.uti diameter) (Fig. 2). Approximately 500-550 areolae are present on each theca, along with 70- 80 evenly spaced marginal areolae. The intercalary band is broad and transversely rugose with long sinuous rugae (Figs. 1,2). Viewed with LM, the valve margins have a distinct striated pattern (Figs. 4,5). This type of intercalary band is unique to this species (Faust 1990b). The peritlagellar area is relatively small, without ornamentation, and set into a shallow, V- shaped depression on the right valve (Figs. 1-3). The flagellar pore is much larger than the auxiliary pore (Fig. 3)(Faust 1990b). Nomenclatiiral Types: Holotype: Prorocentrum ruetzlerianuin Faust, 1990: figs. 21-23 Type Locality: Caribbean Sea: Twin Cays, Belize, Central America Morphology and Structure: Prorocentrum ruetzlerianum is a photosynthetic species with golden chloroplasts, a centrally located pyrenoid (Figs. 4,5), 1990b). and a posterior nucleus (Faust Reproduction: Prorocentrum ruetzlerianum reproduces asexually by binary fission. Ecology: P. ruetzlerianum is a benthic species associated with floating detritus and sediment. This is not a common species and is often in low numbers when present. Cells are motile or attach to detrital particles (Faust 1990b). Toxicity: Quod (1996, pers. com.) has shown that this species is a toxin producer; however, the toxin principals have yet to be determined. Species Comparison; There are several deeply areolated Prorocentrum species all with varying amounts of areolae per valve: P. ho/fmannianum has approximately 670 round to oval areolae per valve (1.1 |im diameter)( Faust 1990b); P. bellzeanum has about 853-1024 round to oval areolae per valve (0.73 |im diameter)(Faust 1993a); and P. sabulosum has about 391 round to oval areolae per valve (1.3 |im diameter)(Faust 1994). Etymology: This species was named after Dr. Klaus Ruetzler, Invertebrate Zoologist, National Museum of Natural History, Smithsonian Institution, for his extensive investigations on Twin Cays mangrove ecology, his patience, advice, encouragement, and generous support of microbial ecology investigations. Habitat and Locality: Populations of P. ruetzlerianum are often associated with floating detritus and sediments in tropical coastal regions of the Caribbean Sea (Faust 1990b). Harmful Marine Dinoflagellates 67 GLOSSARY anisogamous - Sexual reproduction in which the gametes differ from each other morphologically (Taylor 1987); e.g. Alexandrium tamaren.se. amphitrophy - Nutrition mode of photosynthetic dinoflagellates in which either heterotrophy or autotrophy alone can support cell functions. antapex - The posterior-most part of the cell. antapical - In dinokonts, the posterior pole of the cell. antapical plates - In thecated dinokont species, the plates covering the posterior end of the cell (designated with "") not in contact with the cingulum. anterior - In desmokonts, the top part of the cell. apex - The anterior-most part of the cell. apical - In dinokonts, the anterior pole of the cell. apical collar - A topographic feature of some prorocentroids (desmokonts). It is an extension of the intercalary band on the left valve along the anterior margin bordering the periflagellar area (e.g. P. belizeanum and P. hoffmanniamtm). This feature can be curved, flared, rounded or flattened. Oftentimes, this feature can only be viewed via SEM. apical horn - A prominent apical extention of the cell formed by apical plates; it is a feature only found on thecate species. In these species, the apical horn constitutes the apex of the cell (Steidinger & Tangen 1996). apical plates - In thecated dinokont species, the thecal plates that surround and are in contact with the apex of the cell (designated with ' ) not in contact with the cingulum. In those species with an apical pore complex (APC), the plates that touch the APC. apical pore (ap) - Pore located on the Po plate. This feature is not always a round or oval hole, but can be long and narrow and/or curved, or even fishhook shaped. In Alexandrium spp. the ap is referred to as a foramen. If the ap is a hole, then it may have a closing/cover plate (cp) or canopy. apical pore complex (APC) - This feature is located on the epitheca of many marine, armoured (thecated) dinokont species. It includes an apical pore plate (Po), which bears an apical pore (ap), and often times, small periferal pores. In addition, there can be a ventral apical plate or canal plate (X plate). The X plate is always posterior and ventral to the Po. apical pore plate (Po) - Part of the apical pore complex (APC); a feature located on the epitheca of many marine, armoured (thecated) dinokont species. The Po houses an apical pore (ap), and often times, small periferal pores. The Po can be long and narrow, as in Ostreopsis spp., or wide and triangular, as in Gamhierdisciis spp. areolae - Surface ornamentation on thecal plates that approximates deep depressions with or without raised sides. The sides may be round to polygonal and are closely appressed. Areolae can contain pores, even double pores. armoured - Dinoflagellate species that have thecal plates of varying thickness and orientation in identifiable tabulation series. Often the plates are thickened or ornamented with reticulations, spines, grooves, etc... which are often characteristic to a species. asexual reproduction - A method of reproduction where a IN cell produces two to four cells with the same chromosome number. This can be by binary fission of a motile stage or a nonmotile stage. In many armoured dinokonts the original cell divides along predetermined sutures and then each half produces a new half with new thin plates. autotrophy - Photosynthetic nutritional mode in which inorganic compounds (CO2 and carbonates) are utilized for growth, metabolism and reproduction. auxotrophy - Heterotrophic nutritional mode in which specific external organic compounds (usually the vitamins B12, biotin and thiamine) are required in small amounts by most photosynthetic dinoflagellates. benthic - Occuring at the bottom of the water column. binary fission - A method of asexual reproduction in which the parent cell divides into two equal, or nearly equal, parts, each of which develops to parental size and form. 68 Harmful Marine Dinollagellates bioluminescence - The emission of light from certain species of dinoflagellates by either mechanical or chemical stimulation. bloom - High concentrations of planktonic organisms due to enhanced cell division (growth) rates. Seasonal blooms are often related to periodical increase in nutrient and light conditions (e.g. spring bloom). Exceptional blooms are often dominated by one or a few species and may discolor the water a reddish-brown color, hence the name 'red tide'. canal plate (X-plate) - A narrow elongated plate found on the epitheca of some thecate dinokont species ventral to the APC. catenate - Cells connected in a series; cells in chain formation. chlorophyll - Plant pigments found in chloroplasts which function as photoreceptors of light energy for photosynthesis. chloroplast - Membrane-bound organelle found in the cytoplasm of various eukaryotic organisms that contain the chlorophyll pigments and the enzyme systems for photosynthesis. chromosomes - Complex, helical structures in plant and animal nuclei that carry the linearly arranged genetic units, DNA and RNA. ciguatera - A human intoxication caused by ingestion of tropical piscivorous reef fishes contaminated with toxin-producing benthic/epiphytic dinoflagellates. These fish accumulate biotoxins through the food chain (Steidinger 1993). More than 175 separate gastrointestinal, neurotoxic, or cardiovascular symptoms may be associated with this poisoning (Becker & Sanders 1991). In extreme cases death can result from respiratory failure. Although incidence is high, human mortality is low (Hallegraeff 1995). cin<»ular plates - In thecate dinokont species, the plates that make up the cingulum (designated with a 'c'). cingulum - In dinokont species, this structure is usually a furrow (girdle) encircling the cell once or several times, and it can be displaced. In thecated species, the cingulum is made up of plates. This structure is missing in some desmokont-type cells (e.g. Prorocenirum). closing plate (cp) - A small plate located in the apical pore complex (APC) of some marine. armoured dinokont species, and associated with the apical pore (ap). concave - Hollowed or rounded inward resembling the inside of a bowl. convex - Curved or rounded resembling the exterior of a sphere or circle. costae - A rib or rib-like structure, often located in the apical pore complex of thecated species; e.g. Coolia monotis. cyst - Any dormant or resting nonmotile cell possessing a distinct cell wall. cytoplasm - Protoplasm within a plant or animal cell external to the nucleur membrane. DAPl (4',6-Diamidino-2-Phenylindole) - A highly specific and sensitive tluoresceing DNA stain used in epitluorescent microscopy to observe structures containing DNA. DAPl specifically binds to double stranded DNA, and when excited with light the DAPI-DNA complex fluoresces a bright blue (Porter & Feig 1980). desmokont - A dinoflagellate ceil type in which two dissimilar flagella emerge from the anterior part of the cell; e.g. Prorocentrum sp. This morphological type does not have a cingulum or a sulcus. diameter - With the exception of the Prorocentroids, the maximum cell width measured between the lateral extremes of the cingular flanges (Balech 1995). Diarrhetic Shellfish Poisoning (DSP) - A human gastrointestinal disease caused by the ingestion of toxic marine shellfish (filter- feeding bivalves) from cold and warm temperate regions of the Atlantic and Pacific Oceans (Steidinger 1993). Shellfish can accumulate and store large quantities of red tide dinoflagellate toxins without apparent harm to themselves (Steidinger & Baden 1984). Symptoms include diarrhea, nausea and vomiting lasting a few days. No human deaths have been reported (Hallegraeff 1995). dinoflagellate - Biflagellated unicellular alga member in the Phylum Pyrrhophyta. dinokont - A dinoflagellate cell type in which two flagella are inserted ventral ly; one flagelluin is transverse and housed in a cingulum and the other is longitudinal and housed in a sulcus. A dinokont dinoflagellate can be a thecate species (with thecal plates) or an athecate species (without thecal plates)(Steidinger & Tangen 1996). Harmful Marine Dinotlagellates 69 dinophysoid - Group of dinokont type dinoflagellales. Members of this group are the only thecate dinoHagellates fundamentally divisible into two lateral halves and have an anterior cingulum and a narrow sulcus. They are laterally compressed, and their shapes in lateral view are essential for genus and species identification (Taylor etal. 1995). diploid - A cell that has a nucleus with two sets of chromosomes (2N). dorsal - Of or relating to the back side of an organism. In dinokonts, opposite the ventral side (front side)(Steidinger & Tangen 1996). dorsoventral - Extending along the axis joining the dorsal and ventral sides. encystment - To form or become enclosed in a cyst (resting spore). There are several types of encystment. Stressed cells can 'round up' and settle out of the water column and yet be viable if the stress conditions are removed. Others involve temporary cysts for asexual reproduction, flotation, and other functional aspects of individual survival. Yet another type of encystment involves sexual reproduction and the production of thick- walled hypnozygotes which can remain encysted for months, even years (Steidinger & Tangen 1996). epibenthic - Attached to the bottom. epifluoresceiice microscopy - A method of microscopy used to view light-excited regions of an organism stained with a fluorochrome dye. epiph\te - An alga which attaches itself and lives nonparasitically on another plant or on some nonliving object. Cell can attach via a mucoid holdfast or thread(s). epitheca - The anterior part of the dinokont-type cell above the cingulum. eukaryote - A cell with a membrane-bound nucleus. excystment - When the hypnozygote matures and is ready to produce a motile cell from the resting cell, a naked cell will emerge from an opening in the cyst wall. This emerging cell will either be flagellated or amoeboid. Typically, this cell will undergo meiosis and produce four vegetative IN cells that are motile in the water column (Steidinger & Tangen 1996). flagellar pore - in desmokont dinoflagellates, the pore in which tlagella emerge located in the peritlagellar area (flagellar pore area). flagellum - Whip-like structures arising from the cell and responsible for propelling cells in a watery fluid. All dinoflagellates at some time in their life cycle have two dissimilar flagella: a transverse flagellum (provides propulsion) and a longitudinal flagellum (provides direction). They either emerge through one pore or two separate pores. fluorescence - Emission of energy as visible light. foramen - A relatively large comma-shaped cavity (apical pore) on the Po plate of Alexandrium spp. (sometimes fishhook shaped as in Alexandrium catenella and A. tamarense). It is a diagnostic feature of the APC. gametes - In armoured and unannoured species a IN cell that fuses with another IN cell to produce a zygote (2N). geotropic - Oriented by gravity. growth - Increase of body volume, and proliferation of a cell. haploid - Vegetative or gametic cells that have one set of chromosomes (IN). hepatotoxic - Toxic to the liver. heterothallism - Sexual cycle in dinoflagellates which involves opposite mating types; e.g. Gymnodinium catenatum. heterotrophy - Nutritional mode in which absorption of organic matter is required for growth, metabolism and reproduction; e.g. auxotrophy, mixotrophy, myzocytosis, phagotrophy and organotrophy. horn - In armoured cells an extension of the apical or antapical plates. hypnozygote - A thick-walled zygote formed following fusion of two motile gametes (dipIoid-2N). hypotheca - The posterior part of a dinokont- type cell below the cingulum. hystrichosphere - A fossilized dinoflagellate cyst. intercalary band - Marginal growth zones between thecal plates; usually straited horizontally or transversely. isogamous - Sexual reproduction in which the fusing gametes differ morphologically from the vegetative cells, but are morphologically identical to each other (Taylor 1987); e.g. Alexandrium mondatum. 70 Harmful Marine Dinoflagellates lacerate - With a deeply and irregularly incised margin. lanceolate - Tapering at both ends. lateral - In desniokonts oriented toward the left or right side of the cell. lenticulate - Shaped like a double convex lens; shaped like a lentil. life cycle - A continuum of phases and cell types in the reproduction and growth of a species. The life cycle usually contains at least an asexual phase in which a cell can divide by binary fission and produce two similar cells (IN). It may also contain a sexual phase in which gametes fuse to form zygotes (2N) and these zygotes produce IN cells. list - Membranous thecal extensions of armoured dinoflagellates (often associated with the cingulum and sulcus); some extensions are curved or ribbed. lobe - A rounded projection on a structure. inegacytic growth zone - The cell growth that occurs at the suture between the two valves of the Prorocentrales or the fissure halves of the Dinophysiales. When this usually horizontally striated zone is at its maximum extent, the cell will be at its greatest depth or width, respectively (Sleidinger & Tangen 1996). niesokaryotic - Dinotlagellate nucleus which possesses characteristics of both prokaryotes and eukaryotes. mixotrophy - Heterotrophic nutritional mode of some photosynthetic dinotlagellates in which ingestion of food panicles is required for existence. niucocyst - An ejectile organelle; a minute structure that emerges through pores in the theca of armoured dinotlagellates that releases mucous or mucous threads when discharged. myzocytosis - Heterotrophic nutritional mode in which prey is suctioned into a food vacuole via a feeding tube or peduncle, and then digested (Schnepf & Deichgraber 1983). neritic - The region of shallow water adjoining the seacoast; e.g. bays, lagoons, mangroves, salt marshes, etc. Neurotoxic Shclltlsh Poisoning (NSP) - A human neurological disease caused by the ingestion of toxic marine shelltlsh (Ulter- feeding bivalves). Symptoms are similar to those of ciguatera poisoning and include temperature reversal sensations, as well as headache, chills, and muscle and joint pain (Hallegraeff 1995, Steidinger 1993). Cases have been reported from the southeast US and eastern Mexico (Steidinger 1993). nucleus - A membrane-bound organelle in eucaryotic cells which contains a large percentage of the genetic material in the cell. In dinoflagellates, it is most often referred to as a mesokaryon or a dinokaryon due to its unique feature: chromosomes are permanently condensed. organelle - A specialized subcellular structure having a special function; e.g. mitochondria. organotrophy - Heterotrophic nutritional mode of dinotlagellates without chloroplasts; i.e. total nutrition and growth is derived exclusively from organic compounds. osmotrophy - Active uptake of dissolved organic substances for nutrition. ovate - Shaped like an egg; one end broader than the other. Paralytic Shellfish Poisoning (PSP) - A human neurological disease caused by the ingestion of toxic marine shellfish (filter-feeding bivalves) as well as other harvested seafood. PSP has been reported from cold and warm seas (Steidinger 1993). Shellfish can accumulate and store large quantities of bloom or red tide dinotlagellate toxins without apparent harm to themselves (Steidinger & Baden 1984). Symptoms include: tingling sensation around lips gradually spreading to face and neck; prickly sensation in fingertips and toes; headache, dizziness, nausea, vomiting, diarrhea. In extreme cases, muscular paralysis occurs resulting in death from respiratory paralysis (Hallegraeff 1995). peduncle - A small, flexible, fmger-like appendage located near the flagellar pores in some photosynthetic as well as nonphotosynthetic species. Its functions are not fully understood, but it has been associated with feeding behavior (phagotrophy). pellicle - A retaining envelope which may be found around certain dinoflagellates or which can develop at a certain stage of the life history. periflagellar area - In prorocentroids (desniokonts) this region is located on the anterior end of the right valve within a notched area (V-shaped triangular Harmful Marine Dinoflagellates 71 depression). It consists of several plates or platelets around one or two peritlagellar pores, the auxilliary pore (A) and the flagellar pore (F). Accessory pores, apical spine(s), periflagellar collars and/or peritlagellar plates may also be present. periflagellar collar - A topographic feature of the periflagellar area of some prorocentroids (desmokonts). It is a thecal extension of a periflagellar plate that can lie adjacent to the periflagellar pores (flagellar and auxiliary) and/or surround one or both pores (e.g. Prorocentrum macidosum). A periflagellar collar can be flared and/or protuberant, or appear as a winged spine (e.g. Prorocentrum mexicamim). Oftentimes, this feature can only be viewed via SEM. periflagellar plates - A topographic feature of the periflagellar area of some prorocentroids (desmokonts); platelets situated around the periflagellar pores. periflagellar pores - In prorocentroids (desmokonts), large apical pores found in the periflagellar area: auxiliary pore (A) and/or the flagellar pore (F). A periflagellar collar can surround these pores; e.g. P. maculosum. phagotrophy - Heterotrophic feeding mode of generally non-photosynthetic dinoflagellates in which whole prey (or parts of) are ingested or engulfed, with digestion occurring in phagocytic vacuoles. photosynthesis - The use of only inorganic compounds for growth, metabolism and reproductionin presence of light. phytoplankton - Planktonic plant life. pigments - Any coloring matter in plant or animal cells. plankton - Refers to free-living organisms in aquatic environments that have little or no self-motility and therefore float and drift under the action of water movement. plastids - Cytoplasmic organelles of photosynthetic cells that serve as centers of specialized metabolic activities. pore - Openings or channels in the theca of dinoflagellates that can be involved in extrusion of trichocysts or mucocysts and other active processes. Pore number and location are variable within a species, but in many groups, the pattern is a reliable, but variable character for identification of species (Steidinger & Tangen 1996). poroid - Shallow surface depressions on the valve surface. post-cingular plates - In thecated dinokont species, the plates touching the cingulum in the hypotheca (designated with "' ). posterior - In desmokonts, the bottom end of the cell. pre-cingular plates - In thecated dinokont species, the plates touching the cingulum in the epilheca (designated with " ). premedian cingulum - In dinokont-type cells when the cingulum is above the midpoint of the cell. prokaryote - A cell that contains a primitive nucleus where the DNA-containing region lacks a definitive membrane; e.g. bacteria and cyanobacteria. prorocentroids - Group of desmokont type dinoflagellate. Two anteriorly inserted flagella and two large laterally flattened valves easily distinguish the species of this group. The right valve has a small indentation, the periflagellar area, that houses the flagellar pore(s) (Taylor et al. 1995). pustule - A small elevation on the valve surface resembling a blister or pimple. pyrenoid - Cytoplasmic structure made of protein or appended to the chloroplasts in most phytoflagellates. They are associated with the formation or storage of polysaccharide food reserves (usually starch)(Steidinger & Tangen 1996). reticulae - Surface ornamentation on thecal plates where raised straight lines or ridges cross one another creating a complex pattern resembling a network of fibers, veins or lines. rugose - Uneven surface covered with shallow minute depressions. Scanning Electron Microscopy (SEM) - Instrumentation used to observe minute surface details of small organisms/objects at high magnification by means of electron lenses. SEM techniques are often employed and needed to correctly identify a dinoflagellate species. sexual reproduction - A method of reproduction that involves two gametes (IN) that fuse to produce a zygote (2N). striae - Surface ornamentation on unarmoured or armoured dinoflagellates that appear as longitudinal lines, ridges or grooves; on armoured species the striae can be interrupted 72 Harmful Marine Dinotlagellates by pores and may be associated with other markings, such as reticulations. sulcus - Longitudinal area on the ventral surface of dinokont-type cells that forms a pronounced furrow or depression that houses the longitudinal flagellum. In thecated species, the sulcus is made up of sulcal platelets (designated by 's'). This feature is not present in some desmokont-type cells (Steidinger & Tangen 1996). sutures - in armored species, visible linear boundaries between thecal plates (Steidinger & Tangen 1996). theca - Dinoflagellate membrane system encompassing the whole cell consisting of a complex of three to six membranes. thecal plates - Plates of armoured (thecated) species which are composed of cellulose or polysaccharide microfibrils. Their particular size, shape and arrangement on the cell are characteristic to a species (Steidinger & Tangen 1996). traiisdiameter - With the exception of the Prorocentrales, the cell width measured between the lateral extremes of the cingulum bottom; i.e. the flanges are excluded. Minimum cingular width; a measurement of width used in armoured dinokonts (Balech 1995). trichocysts - A cytoplasmic ejectile organelle; a minute structure that emerges through pores in the theca of armoured dinotlagellates that releases tllamentous or fibrillar threads when discharged. tycoplaiiktonic - Benthic dinoflagellate species found at some time in the water column. unarmoured - Dinokont-type cells that do not have an identifiable plate series and do not have apical pore complexes. vacuole - A cytoplasmic membrane-bound cavity within a cell that may function in digestion, storage, secretion or excretion. valves - In the thick-walled desmokonts, two opposing halves of the theca are called valves (right and left). The right valve is the one most indented anteriorly by the periflagellar plates. ventral - The front side of an organism (opposite dorsal side): in dinokonts, side of sulcus and juncture of the cingulum-sulcus; in dinokonts, the side of flagellar insertion (Steidinger & Tangen 1996). ventral pore (vp) - In some armored species, a ventral pore may be present at the juncture of the first apical plate (T) and an anterior intercalary or another apical plate on the epitheca. Sometimes the pore is in one of the apical plates. The presence of a ventral pore or its placement may be diagnostic for certain species (Steidinger & Tangen 1 996). ventral ridge - In dinokonts, an identifiable ridge on the right side of the sulcal intrusion onto the epitheca. verniiculate - Surface ornamentation on thecal plates in which the plates are marked with irregular fine lines or with wavy impressed lines. zygote - A cell (2N) formed by the union of two gametes (IN) during sexual reproduction. 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Mesoscale features of summer (1975-1977) dinoflagellate blooms in the York River, Virginia (Chesapeake Bay Estuary). In: D.L. Taylor & H.H. Seliger (eds.). Toxic Dinoflagellate Blooms, Elsevier/North-Holland, New York: 279-286. Harmful Marine Dinoflagellates 97 PLATE 1 APC [redrawn (ram Balech 1995) (redrawn from Balech 1995) Alexandrium acatenella. Figs. 1-2. LM: ventral view of empty thecae. Cell small to medium, longer than wide, angular to round. Conical epitheca with shoulders; larger than hypotheca. Figs. 3-4. Line drawings. Fig. 3. Ventral view: T plate bears ventral pore (vp). H\potheca with two antapical spines (arrows). Fig. 4. Po comes in direct contact with 1' plate. APC: comma-shaped foramen (arrow). 98 Harmful Marine Dinoflagellates PLATE 2 (Haltegrasffetal. 1991) (Hallegraeffetal. 1991) Alexandrium catenella. Fig. 1. SEM: ventral view. Two cell chain. Cells round; wider than long. Rounded apex and slightly concave antape.x. Cingulum deep and lipped; sulcus deeply impressed and widens posteriorly. H\potheca with prominent sulcal lists (arrows). Fig. 2. LM: Ibur cell chain. Cells anlerior- posteriorly compressed. Fig. 3. LM; apical view. First apical plate (V) comes in direct contact with apical pore plate (Po). Ventral pore absent from 1" plate. Fig. 4. SEM; apical pore complex (APC). ioramcn fishhook shaped; anterior attachment pore (aap) adjacent. Fig. 5. Line drawing: thecal plates depicted. Fig. 6. LM: resting c>st elliptical with rounded ends. Harmful Marine Dinoflagellates 99 PLATE 3 (redrawn from Balech 1 995 Alexandhiim mimitum. Fig, 1. SEM: ventral view. Cell small and ellipsoidal. Epitheca conical, larger than hypotheca. Hypotheca short and wide: antapex obliquely flattened. Intercalary bands present, Cingulum deep, lipped: displaced IX its width. Sulcus shallow (sa=anterior sulcal plate). Apical pore plate (Po) in direct contact with 1 " plate. Fig. 2. LM: ventral view. Ventral pore (vp) present on 1' plate. Fig. 3. SEM: apical view. Po large, narrow and oval: indirectly connected to 1' plate, Vp present (arrow). Figs. 4-5. Line drawing. Fig. 4. Ventral view. 1' plate slender and rhomboidal. Fig. 5. Po connection to 1" plate: a. direct; b. indirect via thin suture. Fig. 6. LM: cyst circular in apical view. 100 Harmful Marine Dinoflauellates PLATE 4 aap. foramen (redrawn from Balech 1995) O (VVfelker S Steitjlnoer 1979^'*' (redrawn from Balech 1995) (Walker & Steidinger 1979} Ale.xandrium nionilainm. Fig. 1. LM: four-cell chain. Cells large, wider than long, flattened anterio- posteriorly. Antapex slightly concave (arrow). Figs. 2- 4. Line drawings. Fig. 2. Ventral pore (vp) depicted {Florida specimens) at anterior margin of 1' plate where it comes in contact with plates 2' and 4', Cingulum (C) deeply excavated, wide, descending; displaced one time its width. Fig. 3. Apical pore plate (Po) does not come in contact with 1' plate. Anterior attachment pore (aap) large, round and dorsally situated in the APC. Foramen comma-shaped. Fig. 4. Antapical view: posterior sulcal plate (sp) large, rhomboid and concave with radial markings. Posterior attachment pore (pap) large and centrally located. Figs. 5-6. LM. Fig. 5. Two isogamous gametes fusing at oblique angles. Fig. 6. Mature resting cysts: dark and round, with a triple layered wall. Harmful Marine Dinotlagellates 101 PLATE 5 (redrawn from Balech 1995) (Fukuyo et al.) Alexandrium ostenfeldii. Figs. 1-3. LM. Fig. \. Ventral view. Cell large and nearly spherical. Cingulum deeply excavated. Epitheca broad and convex-conical. Hypotheca hemispherical with an obliquely flattened antapex. Fig. 2. Epitheca: apical view. Ventral pore (vp) large and distinct. First apical plate (F) forms a 90 degree angle at the point where vp and 4' plate come in contact. Apical pore complex (AFC) with comma-shaped foramen. Figs. 3-4. Line drawings. Fig. 3. Ventral view: 6" plate wider than high. Cingulum (C) slightly excavated. Fig. 4. AFC and 1' plate: a. Po in direct contact with T; b. Po in indirect contact with V via thin suture. Fig. 5. LM: vegetative cell. Small equatorial nucleus (n). Fig. 6. LM: temporary cyst large and spherical, covered in mucilage. Nucleus visible (arrowhead)(Mackenzie et al. 1996). 102 Harmful Marine Dinoflagellates PLATE 6 AU'xaiulriuin pseudogonyaulax. Figs. 1-4. LM Fig. I. VciUral view. Cell broadl\ pentagonal: wider than long. Fpitheca short and dome-shaped. Hypolheca longer than epitheea. Cingulum shallow and barely displaced. Fig. 2, Dorsal view, Antapex obliquely concave. Fig. 3. Epitheea: ventral view. Apical pore plate (Po) with comma-shaped foramen. 1' plate pentagonal with large wide ventral pore (vp) on 4' plate margin. Fig, 4. Epitheea: apical view, t' plate does not come in contact with Po. Po oval and longitudinal on apex. I'igs. 5-6. Fine drawings. Fig. 6. Po and 1' plate not in contact. Fig. 7. LM: isogamous gametes smaller and rounder than vegetative cells. Fig. 8. LM: round resting cyst. Fig. 9. SEM: paratabulate cyst. Harmful Marine Dinoflagellates 103 PLATE 7 (Fukuyoetal.) Atexandrium tamarense. Fi ceils small to medium: si nearly sptierical, Cingulum lipped. Left h\pothcal lobe Nucleus (n) visible. Figs. 2 chain: cingulum displaced (s) widens posteriorly. Fig. Apical pore plate (Po g. 1. LM. Two cell chain: ightly longer than wide, (C) deeply escavated and slightly larger than right. -4. SEM. Fig. 2. Two cell IX its width. Deep sulcus 3. Epitheca: apical view. ) rectangular: narrows ventrally. Po and first apica! plate (T) in direct contact. Small ventral pore present on 1' plate. Fig. 4. Apical pore complex (APC): foramen large and fishhook shaped. Small round anterior attachment pore (aap) present (HallegraetT 199!). Fig. 5. Line drawing. Fig. 6. LM. Oblong resting cyst with rounded ends, reddish lipid bodies; covered in mucilase. 104 Harmful Marine Dinotlagellates PLATE 8 (redrawn from Balech 1 995) Alexandrium tamiyavanichi. Figs. 1-3. LM. Fig. 1. Two cell chain: ceils medium-sized; round to slightly wider than long. Epitheca with shoulders. Fig. 2. Cells stained with calcofluor white: cingulum displaced IX its width: sulcus widens posteriorly. Fig. 3. Apical \iew: apical pore plate (Po) houses comma-shaped tbramen. First apical plate (!') with ventral pore (vp). Figs. 4-5. Line drawings. Fig. 4. 1' plate in direct contact with Po. Po with large central foramen surrounded by small pores. Anterior sulcal plate (s.a.) invades epitheca; an anterior projection ofs.a. fits into a notch in the 1' plate (arrows). Fig. 5. Ventral view: sulcal lists project anteriorly (arrows). Fig. 6. Posterior sulcal plate (s.p.) with round posterior attachment plate (pap) in center (arrow). Harmful Marine Dinoflagellates 105 PLATE 9 Cochlodinium polykrikoides. Figs. 1-7. LM. Fig. 1. Four cell chain. Single cell small and ellipsoid. Epitheca (E) rounded and conical. Hypotheca (H) divided into two posterior lobes (arrows). Numerous rod-shaped chloroplasts. Fig. 2. Cingulum (c) deeply excavated; circles cell 1.8-1.9 times. Fig. 3. Colony of single and chained cells. Fig. 4. Large nucleus (n) in epitheca. Figs. 5-7. Cysts. (Figs. 3,6,7 by Matsuoka & Fukuyo) 106 Harmful Marine Dinoflagellates PLATE 10 Coolia monotis: Figs. 1-5, SEM. Fig. 1. Ventral view: spherical shape. Cingulum lipped and equatorial. Sulcus with tlcxihle lists (arrowheads). Ventral pore present (arrow). Fig. 2. Dorsal view: apical pore plate (arrow), Po, located otT-center on epitheca. Fig. 3. Antapical view: h\pothecal plates. Fig. 4. Smooth edged thecal pores unevenl\ distributed. Fig. 5. Po about 12 ^m long, slightly curved and narrow with a slit-like apical pore. Two supporting rib-like costae (arrows) and evenly spaced round pores surround the pore. Figs. 6,7. LM. Fig. 6. Ventral view of lipped cingulum and sulcus. Fig. 7. Planozygote with two longitudinal tlagella (arrows). Fig. 8. Line drawing: thecal plate arrangement. Harmful Marine Dinoflagellates 107 PLATE 11 (Zingone et al. 1998) (Redrawn from Steidinger & Tangen 1996) Dinophysis acuminata. Figs. 1-5. SEM: lateral view. Fig. I, Cell oval and rotund; thecal surface with shallow depressions and scattered pores. Left sulcal list (LSL) extends beyond midpoint of cell. Weil- developed cingular lists: anterior cingular list (ACL); posterior cingular list (PCL). C=cinguium. Fig. 2. Long and narrow cell with prominent surface areolae, each with a pore. Antapex tapered and ventrally off- center. Small posterior protrusion present (arrow). Fig. 3. Long and narrow cell. Thecal surface smooth with small scattered pores. Megacvlic zone (M) void of pores. Posterior protrusions on antapex (arrow). Figs. 4-5. LM: lateral view. Fig. 4. Surface areolae and tapered antapex (from Larsen & Moestrup 1992: fig. Id). Fig. 5. Large dorsal nucleus (N). Small, blunt projections on tapered antapex (arrow). Fig. 6. Line drawing. 108 Harmful Marine Dinoflagellates PLATE 12 CINGULAR LISTS (Redrawn from Steidinger & Tangen 1996) Dinophysis acuta. Fig. 1. SRM: lateral view. Cell oblong and robust; tlicca heavily areolated. Well developed ciiigular lists (CL) and left suleal list (i.SLK Pointed antapex. Figs. 2-3. VM: lateral view (from Larsen & Moestrup 1992: i'lgs. 2a.d; scale bars=20 ^im). Fig. 2. Large areolae, each with a pore (arrows). Fig. 3. Widest point below mid-section (dashed line) aligned with third suleal rib (arrow). Fig. 4. Line drawing. Harmful Marine Dinoflaeellates 109 PLATE 13 LEFT SULCAL LIST (Redrawn from Steidinger & Tanger 1 996) Dinophysis caiidata. Figs. t-2. SEM. Fig. 1. Large. long and distinctive cell with extended ventral hypothecal process. Cingulum narrow; lists supported by ribs (arrowhead). Strong left sulcal list (double arrows). Right sulcal list present (single arrow). Fig. 2. Ventral view: cell compressed laterally. Figs. 3-4. LM. Fig. 3. Large posterior nucleus (n). Fig. 4. Left sulcal list with three supporting ribs (arrowheads); posterior projection with small knob-like spines (arrows). Surface areolae evident. Fig. 5. SEM. Paired cells joined at dorsal expansion (arrow). Fig. 6. Line drawing. no Harmful Marine Dinoflagellates PLATE 14 LSL {Larsen & Moestrup 1992) 2 (Larsen & Moestrup 1992)^ 3 (redrawn from Steldlnger & Tangen 1996) Dinophysis Jorni. I'ig. 1. SliM: lateral view. Left sulcul list (LSL) long and well-developed. Right sulcal list (I three short ribs (arrows). Ventral hypothecal margin concave below LSL (arrowheads). Fig. 3. Dorsal view; hypothecal margin smoothly convex. Short anterior cingular list (ACL) and posterior cingular list (PCL) supported b\ numerous ribs. Fig. 4. Ventral view: dividing cell. Megacytic zone expanding (arrows). Epitheca, sulcus, RSL and LSL visible. Fig. 5. LM: large nucleus (n). Fig. 6. Line drawing (Phalacroma mitra). 112 Harmful Marine Dinoflagellates PLATE 16 (Larsen& Moestrup 1992) 2 (Larsen & Moestrup 1992) 3 LSL (Larsen & Moestrup 1992) (Larsen & M'^Strup 1 992) 5 ^'^"^'^^^ f^^-" Ste,dinger a Tangen 1996) 6 Dinophysis norvegica. Fig. 1. SEM: lateral view. Cell heavily arcolated with pointed antapcx and posterior protrusions (arrowlicuds). Ventral margin concave below left sulcal list (LSL)(arrow). Well developed cingular lists (CL) and LSL. Figs. 2-5. LM: lateral view. Fig. 2. Cell less robust than in Fig. I; pointed antapex. Fig. 3. Robust cell with rounded antapex. Heavily areolated. Ventral margin straight below LSL (arrows). Fig. 4. Deepest point of cell through mid- point (dashed line), just above third rib of LSL. I"ig. 5. Large posterior nucleus (n). Pointed antapex with posterior projections (arrows). Fig, 6. Line drawing. Right sulcal list depicted (RSL). Harmful Marine Dinoflagellates 113 PLATE 17 O (redrawn from Sleidingef & Tangen 1996) Dinophysis rotundata. Figs. 1-2. SEM: lateral view. Fig. t. Cell broadly rounded. Small cap-like epitheca (c) not obscured by cingular lists. Right sulcal list (arrow). Fig. 2. Left sulcal list (LSL) (large arrow), over 1/2 the cell length, widens posteriorly. Surface pores present (small arrows). Figs. 3-4. LM (from Larsen & Moestrup 1992; figs. 8b.c). Fig. 3. Large food vacuoles (fv). LSL supported by three ribs (arrows). Widest width of cell between second and third rib. Fig. 4. Posterior nucleus (n). Fig. 5. Line drawing (as Phalacroma rotundata). 114 Harmful Marine Dinotlagellates PLATE 18 liLiiiMuiitsciai. I990J (Zjfigoneetai.1998) (after Slein 1883) Dinophysis sacculns. Figs. 1-3. SEM: lateral view. Fig. 1. Cell oblong with rounded posterior, llypotheca long, margins undulate. Thecal surface coarsely areolaled. Short left sulcal list (LSL). Cingulum with two well developed lists. Small blunt posterior projections (arrow). Fig. 2. Cingulum lined with pores. Right sulcal list (RSL) visible. Fig. 3. Smooth thecal surface with pores. Metacytic zone (M) devoid of pores. Figs. 4-5. LM: lateral view. Fig. 4. llypotheca sack-like with deep thecal pores. Posterior end with two blunt projections (arrows). Fig. 5. Large posterior nucleus (n). Fig. 6. Line drawing; morphotype from Stein (1883). Harmful Marine Dinotlagellates 15 PLATE 19 (Larsen & Moestrup 1992) 3 I lO (redrawn (rom Sleidmger & Tangen 1996) Dinophysis tripos. Fig. 1. SEM: lateral view. Cell large, oblong and heavily areolated. Hypothecal projections with toothed posterior ends (arrows). Lett sulcal list (LSL) large, wide and reticulated. Figs. 2,3. LM; lateral view. Fig. 2. Anterior cinguiar list (ACL) projected anteriorly obscuring low epitheca (arrowheads). Narrow cingulum, Chloroplasts visible (arrows). Fig. 3. Paired cells. Hypothecal projection on dorsal margin sometimes seen with a narrow list (arrow) connecting two daughter cells during cell division. Fig. 4. Line drawing. 116 Harmful Marine Dinoflagellates PLATE 20 Gamhienliscus foxiciis. Figs. 1-3. SEM. Fig. 1. Fpithcca; cell round lo ellipsoid: anterior-postcriorly compressed. Cell surface smooth with small scattered pores. Apical pore complex located at the apex (arrow). Fig. 2. Hypothcca: Ip plate large and pentagonal. Sulcal region deeply excavated (arrow). Fig. 3. Apical pore plate with characteristic fishhook shaped apical pore. Fig. 4. LM. Fpitheca: cingulum and sulcal region in focus. Fig. 5. LM. Hypotheca: sulcal ridge (arrow); large nucleus (n). Fig. 6. iJne drawing: thecal plate arrangement. Harmful Marine Dinoflagellates 117 PLATE 21 (redrawn Irom Steidmger * Tanger 1996) Gonyaulax polygramma. Figs. 1-3. SEM. Fig. 1. Ventral view: cell large, elongate and quadrilateral. Epitheca with prominent apical horn (arrow). Cingulum left-handed, displaced 1.5 X its width; sulcus widens posteriorly. Longitudinal ridges on thecal surface with reticulations in between. Fig. 2. Lateral ventral view: transverse (TF) and longitudinal (LF) tlagella present. One antapical spine (arrow). Fig. 3. Dorsal view: hypotheca truncate with straight sides. Three antapical spines (arrows): one large and two small. Figs. 4-5. LM. Fig. 4. Ventral view: reticulations evident; one long antapical spine (arrow). Fig. 5. Dorsal view: prominent longitudinal ridges. Fig. 6. Line drawing. 118 Harmful Marine Dinoflagellates PLATE 22 (after Steidinger et al. 1978) (redrawn Irom Steidmger & Joyce 1973) Gymnodinium breve. Fig. I. SEM: ventral view. Cell small, wider than long, dorso-ventrally flattened. Cell ncarK square in outline; prominent apical process (APj directed ventrally. Apical groove (AG) present on apical process, adjacent to sulcus. Figs. 2-3. LM. Fig. 2. Dorsal view; large nucleus (N) in hypotheca. Transverse (TF) and longitudinal (LF) flagella present. Hypotheca bilobed (arrow). Fig. 3. Ventral view: displaced cingulum (large arrow) and lipid globule (small arrow). Fig. 4. Line drawing. Cingulum (C) displaced, descending. Long sulcus (S) extends to apex of cell, Harmful Marine Dinoflagellates 119 PLATE 23 (YUKI & YOSHiMATSU 1 987) from Sleidinger 4 ranqen 1 9961 Gymnodinium catenatiim. Figs. 1-3. SEM: ventra! view. Fig. 1. Cell small, elongate-ovoid with slight dorso-ventral compression. Conical apex; rounded and notched antapex. Cingulum (C) excavated; sulcus (S) long. Distinctive horse-shoe shaped apical groove (AG) encircles apex. Fig. 2. Two cell chain; attachment point visible (arrow). Premedian cingulum displaced 2X its width. Longitudinal (LP) and transverse (TF) tlagella visible. Fig. 3. Chain cells with anterior-posterior compression. Terminal cell slightly longer. Thecal surface rugose to smooth (Blackburn et al. 1989). Figs. 4-5. LM. Fig. 4. Chain- formation (Yuki and Yoshimatsu 1987). Fig. 5. Single cell. Conical epitheca with concave to flat apex. Bilobed hypotheca (arrow). Fig. 6. Line drawing. Fig. 7. SEM: cyst with microreticulations. ag=apical groove; c=cingulum 120 Harmful Marine Dinotlageltates PLATE 24 '^'W^>:^**^*«-' Gymnodinnim mikimoloi. Figs. 1-4. SEM. Tig. 1. Ventral view: cell small, broadly oval to almost round. Epitheca slightly smaller than hypotheca. Characteristic straight apical groove (AG). Cingulum (C) deep, displaced 2 times its width. Sulcus (S) slightly invades epitheca (arrowheads). Hypotheca notched b\ widening sulcus (arrow). Fig. 2, Dorsal view: apical groove extends to dorsal side of epitheca creating slight indentation at the apex (arrowhead). Hypotheca bilobed (arrow). Fig. 3. Apical view of apical groove (arro\v)(arter Fukuyo et al.). Fig. 4. Cell compressed dorso-ventrall) (after Fukuyo et al.). Figs. 5-7. LM. Fig. 5. Cingulum displaced 2 times its width (arrows)(from Larsen & Moestrup 1989: tig. 16g). Fig. 6. Large nucleus (N) in left lobe of hypotheca. Fig. 7. Vegetative division. Division plane oblique. Harmful Marine Dinotlagellates 12 PLATE 25 (after Larsen 1994} (Larsen 1994S Gymnodiniiim pulchellum. Figs. 1-2. SEM: ventral view. Fig. 1. Cell . .lall and broadly oval. Cingulum wide, displaced 1-1.5 X its width. Deeply excavated sulcus creates lobed hypotheca. Conspicuous undulating apical groove (AG). Fig. 2. Well- deseloped apical groove: reverse S-shape. Transverse tlagellum (TF) housed in cingulum. Sulcus slightly invades epitheca with finger-like projection (arrow). Figs. 3-5. LM: ventral view. Figs. 3-4. Apical groove distinguishable (small arrows). Chloroplasts and pyrenoids present, Lobed hypotheca (large arrow). Fig. 5. Large elliptical nucleus (N) in left central part of cell. Fig. 6. Line drawing. C=cingulum 22 Harmful Marine Dinotlagellates PLATE 26 (Hallegraeff 1991) Gymnodinium sangitineum. Figs. 1-3. LM. Cell large, pentagonal, and slightly dorso-ventrally flattened. Cells vary in shape and size. Fig. 1. Ventral view. Epitheca and hypotheea nearly equal in size: epitheca conical, hypotheca bilobed (arrows). Fig. 2. Ventral view. Deep cingulum median, displaced 1-2 times its width. Sulcus deeply notches hypotheca. Apical groove present (arrow). Fig. 3. Cell deeph pigmented; central nucleus (n). Fig. 4. Line drawing. Spindle- shaped chloroplasts radially arranged. Harmful Marine Dinoflagellates 123 PLATE 27 A) I CD ^^^ Ol 0) » 4) fl) n T7 a) Q 01 £ 3 CD ^ to Ul O) Gymnodinhim veneficum. Figs. 1-3. Line drawings. Fig. 1. Ventral view: cell small and ovoid. Epitheca slightly pointed, without apical groove. Cingulum deep and displaced 1-2 times its width. Fig. 2. Dorsal view: large central nucleus (N). Two to eight irregular chloroplasts present (C). Fig. 3. Sigmoid sulcus slightly invades epitheca (arrow). 124 Harmful Marine Dinoflagellates PLATE 28 (redrawn from Steidinger & Tangen 1996) AG {after Taylor etal. 1995) Gyrodinium gaiatheamtm. Figs. 1-2. SEM: ventral view. rig. 1. Cell small, oval to round, with distinct apical groove (AG). Cingulum (C) displaced 3 times its width. Short and narrow sulcus (S) slightly invades epitheca. Fig. 2. Epitheca and hypotheca round. Cingulum wide, houses transverse nagellum (single arrow). Longitudinal tlagella present (double arrow). Fig. 3. LM: ventral view. Cingulum deepi) excavated (arrows). Nucleus (N) large and central. Fig. 4. Line drawing. Harmful Marine Dinoflagellates 125 PLATE 29 Lingulodinium polyednim. Figs. 1-3. SEM. Fig. 1. Ventral view: cells angular and polyhedral-shaped. Thick plates well defined and coarsely areolate. Epitheca with shoulders and nearly flattened apex. Hypotheca with straight sides and flattened antapex (arrow). Cingulum deep and displaced 1-2 X its width. Sulcus widens posteriorly. Fig. 2. Apical view; first apical plate (F) long and narrow. Apical pore plate (Po) with raised inner elliptical ridge. Cingulum with lists (arrowheads). Strong ridges along sutures outline thecal plates. Fig. 3. Thecal areolae with large trichocysts (arrow)(Lewis and Burton 1988). Fig. 4. Line drawing. Figs. 5-6. SEM: resting cysts. Fig. 5. Cyst sperical with numerous tapering spines. Fig. 6. Cyst theca after excystment. 126 Harmful Marine Dinotlagellates PLATE 30 "JfttT' c o to