SMITHSONIAN INSTITUTION Contributions from the United States National Herbarium Volume 42: 1-144 Identifying Harmful Marine Dinoflagellates ee saat ciel ene . wt HS ON) MAY Ud 2UU2 LIBRARIES 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. Gulledge. Identifying Harmful Marine Dinoflagellates. Smithsonian Contributions from the United States National Herbarium, volume 42: 144 pages (including 48 plates, | figure and | table). — A taxonomic identification and reference guide of 48 harmful marine dinoflagellate species present in the world’s oceans. This guidebook illustrates the morphology and taxonomy of harmful marine dinoflagellates of the following genera: Alexandrium, Dinophysis, Gymnodinium, 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, dinophysistoxins, and ichthyotoxins. Detailed taxonomic descriptions of plate and thecal morphology, 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, biogeography, distribution, and habitat and locality. Species illustrations presented as scanning electron micrographs, differential interference contrast and epifluorescence 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 dinoflagellate 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 Prorocentrum hoffmannianum Faust, back Prorocentrum ruetzlerianum Faust. Contributions of the United States National Herbarium (ISSN 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 History, Smithsonian Institution, Washington, DC, 20560-0166, USA. The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials 739.48--1984. Contributions from the U.S. National Herbarium was first published in 1890 by The United States Department of Agriculture. From July 1, 1902 forward it was published as a Bulletin of the United States 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 . catenella . minutum . monilatum . ostenfeldii . pseudogonyaulax . tamarense . tamiyavanichi Cochlodinium polykrikoides Coolia monotis Dinophysis acuminata D. acuta D. caudata D. fortii . mitra . Horvegica . rotundata . sacculus . (IPOS Gambierdiscus toxicus Gonyaulax polygramma Gymnodinium breve G. catenatum G. mikimotoi G. pulchellum G. sanguineum G. veneficum Gyrodinium galatheanum Lingulodinium polyedrum Noctiluca scintillans Ostreopsis heptagona O. lenticularis O. mascarenensis O. ovata O. siamensis Pfiesteria piscicida Prorocentrum arenarium P. balticum P. belizeanum P. concavum P. faustiae pe Se eee Se See SS SCOHOHSD 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 4}, 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 P. lima P. maculosum P. mexicanum P. micans P. minimum P. ruetzlerianum GLOSSARY BIBLIOGRAPHY 59, 138 60, 139 61, 140 62, 141 63, 142 65, 143 66, 144 67 73 Harmful Marine Dinoflagellates 5 Identifying Harmful Marine Dinoflagellates Maria A. Faust and Rose A. Gulledge INTRODUCTION Interest in identifying harmful dinoflagellate species has received worldwide recognition in recent years due to the increase in red tides, fish kills, and shellfish poisoning events reported from coastal marine ecosystems (Hallegraetf 1991). The publication, /dentifving Harmful Marine Dinoflagellates, is an effort by the authors to present a fully illustrated identification guide for harmful dinoflagellate taxa. The user will recognize general information on dinoflagellate 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 dinoflagellate 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 aquaculture, 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 dinoflagellate 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). Dinoflagellate 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 6 Harmful Marine Dinoflagellates extended period of time, and then germinate to initiate blooms (Spector 1984). Dinoflagellate *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 fish, 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 fish and shellfish consumption, sometimes resulting in gastrointestinal disorders, permanent neurological damage, or even death. While harmful dinoflagellate 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 1. Table 1. Toxin production and red tide events of harmful marine dinoflagellates Species Red Produced Toxin Reference Tide Alexandrium acatenella YES PSP toxins Prakash & Taylor 1966 A. catenella YES Ichthyotoxins; PSP toxins: C1-4, GTX, Prakash et al. 1971, Fukuyo 1985, Fukuyo SXT et al. 1985, Ogata & Kodama 1986 A, minutum YES PSP toxins: GTX1-4 Oshima et al. 1989 A, monilatum YES Ichthyotoxins PSP toxins: GTX1, SXT — Gates & Wilson 1960, Ray & Aldrich 1967, Schmidt & Loeblich 1979 A. ostenfeldii NO Mild PSP toxins; Spirilides Cembella et al. 1987, 1988, 2000 A, pseudogonyaulax NO Goniodomin A Murakami et al. 1988 A, tamarense YES — Strong PSP toxins: GTX1-5, NSXT, SXT Larsen & Moestrup 1989, Shimizu et al. 1975, Oshima et al. 1977 A, tamiyavanichi NO Strong PSP toxins: GPX, SXT Fukuyo et al. 1989, Kodama et al. 1988 Cochlodinium polykrikoides YES Coolia monotis NO Dinophysis acuminata YES D. acuta YES D. caudata YES D. fortii NO Ichthyotoxins Cooliatoxin DSP toxins: OA DSP toxins: DTX1, OA Ichthyotoxins DSP toxins: DTX1-2, OA Yuki & Yoshimatsu 1989, Kim 1998, Ho & Zubkotf 1979 Nakajima et al. 1981, Holmes et al. 1995 Cembella 1989, Lee et al. 1989, Kat 1985 Lee et al, 1989, Yasumoto 1990 Okaichi 1967 Lee et al. 1989, Yasumoto 1990 Harmful Marine Dinoflagellates Species Red Produced Toxin Reference Tide D. mitra NO DSP toxins: DTX1, OA Lee et al. 1989 D. norvegica YES DSP toxins: DTX1, OA Cembella 1989, Lee et al. 1989, Yasumoto 1990 D. rotundata NO DSP toxins: DTX1 Lee et al. 1989 D. sacculus YES DSP toxins: OA Masselin et al. 1992, Giacobbe et al. 1995, Delgado et al. 1996 D. tripos NO DSP toxins: DTX1 Lee et al. 1989 Gambierdiscus toxicus NO Ciguatoxin, Gambieric acid, Maitotoxin Murata et al.1990, Yasumoto et al. 1977, 1987, 1993, Yokoyama et al. 1988 Gonyaulax polygramma YES _ Fish and shellfish kills due to anoxia after Hallegraeff 1991, Koizumi et al. 1996 red tide Gymnodinium breve YES NSP toxins: Brevetoxins Baden 1983, Baden et al. 1982, Hughes 1979 G. catenatum YES PSP toxins Morey-Gaines 1982, Mee et al. 1986 G. mikimotoi YES NSP toxin: Gymnodimine; [chthyotoxins Hallegraeff 1991, Seki et al. 1996 G, pulchellum YES Ichthyotoxins Onoue et al. 1985, Onoue & Nozawa 1989, Steidinger et al. 1998 G. sanguineum YES Ichthyotoxins Cardwell et al. 1979, Tindall et al. 1984, Carlson & Tindall 1985 G. veneficum NO [chthyotoxins Ballantine 1956, Abbott & Ballantine 1957, Dodge 1982 Gyrodinium galatheanum YES Ichthyotoxins Braarud 1957, Steemann Nielsen & Aabye Jensen 1957, Pieterse & Van Der Post 1967 Lingulodinium polyedra YES PSP toxins: SXT Bruno et al. 1990 Noctiluca scintillans YES _ Fish and shellfish kills due to high levels Okaichi & Nishio 1976 of ammonia after red tide Ostreopsis heptagona NO Unnamed toxin J. Babinchak (according to Norris et al. 1985) O. lenticularis NO OTX, Unnamed toxin Tindall et al. 1990, Ballantine et al. 1988 O. mascarenensis NO Ciguatera toxin? Quod 1994, Morton, S.L. (personal communication) O. ovata NO Unnamed toxin Nakajima et al. 1981 Harmful Marine Dinoflagellates Species Red Produced Toxin Reference Tide O. siamensis NO Unnamed toxin Nakajima et al. 1981, Usami et al, 1995 Pfiesteria piscicida NO Ichthyotoxins Burkholder et al. 1995, Noga et al. 1996, Burkholder & Glasgow 1997 Prorocentrum arenarium NO DSP toxins: OA Ten-Hage et al. 2000 P. balticum YES Unknown toxin Silva 1956, 1963, Numann 1957 P. belizeanum NO DSP toxins: DTX1, OA Morton et al. 1998 P. concavum NO DSP toxins: OA, FAT, Unnamed toxin — Tindall et al. 1984, Tindall et al. 1989, Dickey et al. 1990, Hu et al. 1993 P_ faustiae NO DSP toxins: DTX1, OA Morton 1998 P. hoffmannianum NO DSP toxins: OA, FAT Aikman et al. 1993 P. lima NO DSP toxins: DTX1,2,4, OA, FAT, Murakami et al. 1982, Yasumoto et al. prorocentrolide 1987, Torigoe et al. 1988, Tindall et al. 1989, Lee et al. 1989, Marr et al. 1992, Hu et al. 1993, 1995 P. maculosum NO DSP toxins: OA, Prorocentrolide B Dickey et al. 1990, Hu et al. 1996 P. mexicanum NO FAT Steidinger 1983, Carlson 1984, Tindall et al. 1984 P. micans YES Shellfish kills Pinot & Silva 1956, Horstman 1981 P. minimum NO DSP toxin: Venerupin Nakazima 1965, 1968, Smith 1975, Okaichi & Imatomi 1979, Tangen 1983, Shimizu 1987 P. ruetzlerianum NO Unnamed toxin Quod (personal communication) ABBREVIATIONS: DSP = diarthetic shellfish poisoning, DTX1, DTX2, DTX3, DTX4 = dinophysistoxins,; FAT = fast acting toxin; GTXI, 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 etc.). possess ornamentation (pores, areolae, spines, ridges, Armored or thecate species, those that a multi-layered cell wall, can be Harmful Marine Dinoflagellates 9 distinguished from unarmored or _ athecate 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). Fig. 1. Identifying dinoflagellates: A. lateral view of a desmokont cell type (two dissimilar flagella apically inserted); B. ventral view of a dinokont cell type (two dissimilar flagella ventrally inserted); C. ventral view of a thecate peridinioid cell; D. ventral view of an athecate gymnodinoid cell; E. apical view of epithecal plates; F. antapical view of hypothecal plates. 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 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. | 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 flagella arise from is the ventral side, the opposite side is the dorsal. Ventral view (Fig. | 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. | 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 et al. 1995). In thecated species the plate pattern, or tabulation, is crucial (see Balech & Tangen 1985) (Fig. | 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 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. Brawley, 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 dinoflagellate 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 et Kofoid) Balech, 1985 Plate 1, Figs. 1-4 Species Overview: Alexandrium acatenella is an armoured, marine, planktonic dinoflagellate. It is associated with toxic PSP blooms in Pacific coastal regions. Taxonomical 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 um in length and 26-35 um in transdiameter width (Whedon & Kofoid 1936; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gonyaulax 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'™, 2". 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 broadly 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 | time its width without overhanging. Narrow lists are present on the cingulum (Figs. I- 3). The deeply excavated sulcus widens Harmful Marine Dinoflagellates I 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 al. 1995; Steidinger & Tangen 1996). Morphology and Structure: 4. acatenella is a photosynthetic 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: 4. acatenella reproduces asexually by binary fission (Whedon & Kofoid 1936). Ecology: 4. acatenella is a planktonic species associated with paralytic shellfish poisoning (PSP) events and red tides. Populations are most abundant in neritic waters at 15°C. 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 4. 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 I', 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). Alexandrium catenella (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 um in length and 18-32 um 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, I’, 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: 4. catenella is a photosynthetic species with numerous yellow- green to orange-brown chloroplasts. The nucleus is large and U-shaped (Whedon & Kofoid 1936). Reproduction: 4. 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 dinoflagellate 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 4. 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 um in length to 23-32 um in width (Fukuyo 1985; Hallegraeff 1991; Meksumpun et al. 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: cl-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 A. 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; 198 1a; 1981b), California (Sharpe 1981) and most of the Pacific coast of the U.S.A. (Nishitani & Chew 1988). Species Comparison: 4. catenella is very similar morphologically (size, shape and thecal plate formula) to 4. 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 4. catenella from Japan and 4. tamarense from Japan and the U.S.A. revealed a close genetic relationship between the two species, however they remain distinct (Adachi et al. 1995), Chains of this species are quite distinctive, but can resemble 4. 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: A/exandrium 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 Dinoflagellates 13 Alexandrium minutum Halim, 1960 Plate 3, Figs. 1-6 Species Overview: Alexandrium minutum is an armoured, marine, planktonic dinoflagellate. It is a widely distributed species associated with toxic PSP blooms in coastal regions. Taxonomical 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, |' (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 pm in length and 13-24 um 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, 1960: 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 (1') (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 1' plate (Figs. 2-4). The distinctive sixth precingular plate (6") is long and narrow (Fig. 1)(Balech 1989; 1995; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996; Hwang et al. 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 flat 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. 1,2)(Balech 1989; 1995; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996; Hwang et al. 1999). Morphology and Structure: 4. minutum is a photosynthetic species with an elliptical nucleus (Balech 1989; 1995). Reproduction: 4. minutum reproduces asexually by binary fission. This species also has a sexual cycle that produces a characteristic resting cyst (Fig. 6)(Bolch et al. 1991). Ecology: 4. minutum is a__ planktonic 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 10’ cells/L (Hwang et al. 1999). Another red tide of 4. 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 um in diameter) (Fig. 6); kidney-shaped in lateral view (15-19 pm long). The cyst wall is covered with mucilage (Bolch et al. 1991). Toxicity: A/exandrium minutum is a_ strong producer of PSP gonyautoxins (GTX): GTAXI, 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 al. 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 1960), Italy (Montresor et al. 1990), northern Adriatic waters (Mediterranean Sea)(Honsell 1993), Turkey (Koray & Buyukisik 1988), Spain and Portugal (as A. ibericum)(Balech 1985b), France (Nezan et al. 1989), South Australia (Hallegraeff et al. 1988), and the east coast of the United States (Steidinger & Tangen 1996). Alexandrium monilatum (Howell) Balech, 1995 Plate 4, Figs. 1-6 Species Overview: Alexandrium monilatum is an armoured, marine, planktonic dinoflagellate. It is a coastal warm water species associated with toxic red tides and massive fish and shellfish kills. Taxonomical Description: A very distinctive chain-forming species, A. monilatum typically occurs in long chains of 16 or more cells. Single cells are medium to large, wider than long, and flattened —anterio-posteriorly (Figs. 1,2). Epithecal shoulders are occasionally observed. Thecal plates are thin with many delicate pores. Cells range in size between 28-52 um in length and 33-60 um in transdiameter width (Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996), Nomenclatural Types: Holotype: Gonyaulax monilata Howell, 1953: 153, figs. 1-5 Type Locality: North Atlantic Ocean: Indian River, Florida, USA Synonyms: Gonyaulax monilata Howell, 1953 Gessnerium mochimaensis Halim, 1967 G. monilata (Howell) Loeblich, 1970 Pyrodinium monilatum (Howell) Taylor, 1976 Thecal Plate Description: The plate formula for A. monilatum is: Po, 4', 6", 6c, 10s, 5", 2"". The large apical pore complex (APC) is broadly triangular and slightly curving posteriorly. The large apical pore plate (Po) is ovate with a small comma-shaped foramen (Fig. 3). The anterior attachment pore (aap) is large and round (Fig. 3). Small pores are present along the margin of the Po. The characteristic first apical plate (1') is not connected to the Po; it is short and broadly pentagonal (Figs. 2,3). The 1' plate is typically without a ventral pore, however, specimens from Florida reveal a pore at the juncture where the 1’, 2' and 4' plates meet (Fig. 2)(Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca and hypotheca are nearly equal. The antapex is slightly concave. The median cingulum is deeply excavated, devoid of lists, and is displaced in a descending fashion one time its width (Fig. 2). The sulcus bears a diagnostic feature: a large and rhomboid-shaped posterior Ssulcal plate (s.p.)(Fig. 4). The s.p. is concave and recessed with radial markings, and contains a large central posterior attachment pore (pap)(Fig. 4)(Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). Morphology and Structure: 4. monilatum is a photosynthetic species with central radiating brownish chloroplasts. The quarter-moon shaped nucleus is equatorial (Balech 1995). Reproduction: 4. = monilatum reproduces asexually by binary fission; plane of fission is oblique. This species also has a sexual cycle with armoured isogamous gametes that fuse at oblique angles (Fig. 5). Gametes range in size from 36 X 36 um to 47 X 56 um. After fusion, a planozygote forms which then encysts into a characteristic resting cyst (Fig. 6) (Walker & Steidinger 1979). Ecology: A. monilatum is a planktonic estuarine dinoflagellate species associated with toxic red tides and massive fish mortality events in warm coastal waters off Florida, Texas and Venezuela (Howell 1953; Ray & Aldrich 1967). Offshore coastal water blooms have also been reported in Florida and Texas (Williams & Ingle 1972; Wardle et al. 1975). One reported red tide from Texas had cell concentrations ranging from 5 X 10° cells/L to 10 X 10° cells/L (Gates & Wilson 1960). This species produces a dark colored resting cyst as part of its life cycle. The cyst is smooth and round to ovoid. Cysts range in size from 60 to 87 um in diameter (Fig. 6) (Walker & Steidinger 1979). Harmful Marine Dinoflagellates 15 Toxicity: Alexandrium monilatum produces a strong ichthyotoxin resulting in a paralyzing effect (Gates & Wilson 1960, Ray & Aldrich 1967). From laboratory culture studies, Schmidt and Loeblich (1979) report production of paralytic shellfish poison (PSP) toxins: saxitoxin (STX) and gonyautoxins (GTX1); the toxins are hemolytic and neurotoxic (Bass & Kuvshinoff 1982; Clemons et al. 1980). The toxins produced from this species do not accumulate in shellfish (molluscs do not feed on this species) and it is not toxic to birds (Ray & Aldrich 1967). Massive fish kills have been reported from Texas bays in the Gulf of Mexico (Gunter 1942; Connell & Cross 1950; Ray & Aldrich 1967) and on the east coast of Florida in the Atlantic Ocean (Howell 1953). Habitat and Locality: Alexandrium monilatum is a warm water species known from subtropical and tropical regions of the Atlantic Ocean: east coast of Florida (Howell 1953), Venezuela in the Caribbean Sea (Halim 1967), and Texas in the Gulf of Mexico (Gunter 1942; Connell & Cross 1950; Ray & Aldrich 1967). Populations have also been reported from the tropical Pacific Ocean off Ecuador (Balech 1995), and surprisingly in the Chesapeake Bay (Morse 1947). Alexandrium ostenfeldti (Paulsen) Balech et Tangen, 1985 Plate 5, Figs. 1-6 Species Overview: Alexandrium ostenfeldii is an armoured, marine, planktonic dinoflagellate. Generally, it is a cold-water coastal species found in low numbers mainly along the west coast of Europe. Taxonomical Description: A distinctive species, cells of A. ostenfeldii are large and nearly spherical (Fig. 1). Cells are single, but are often found in two-celled colonies. Epitheca and hypotheca equal in height (Figs. 1). This species has thin thecal plates and a characteristic large ventral pore on the first apical plate (1')(Fig. 2). Faint surface pores are numerous and unevenly distributed. Cells range in size between 40-56 um in length and 40-50 um in transdiameter width (Balech 1995; Balech & Tangen 1985; Konovalova 1993; Larsen & Moestrup 1989; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Goniodoma ostenfeldii Paulsen, 1904: 20, fig. 2 Type Locality: Iceland Synonyms: Goniodoma ostenfeldii Paulsen, 1904 Goniaulax tamarensis Lebour var. globosa Braarud, 1945 Goniaulax ostenfeldii (Paulsen) Paulsen, 1949 Heteraulacus ostenfeldii (Paulsen) Loeblich, 1970 Gonyaulax globosa (Braarud) Balech, 1971b Gonyaulax trygvei Parke, 1976 Protogonyaulax globosa (Braarud) Taylor, 1979 Gessnerium ostenfeldii (Paulsen) Loeblich and Loeblich, 1979 Pyrodinium phoneus Woloszynska and Conrad, 1939 Triadinium ostenfeldii (Paulsen) Dodge, 1981 Thecal Plate Description: The plate formula for A. ostenfeldii is: Po, 4', 6", 6c, 10s, 5", 2"". The apical pore complex (APC) is triangular or rectangular in shape. The apical pore plate (Po) is relatively large with a large comma-shaped foramen (Figs. 2,4). It can be either in direct contact with the first apical plate (1')(Fig. 4a) or indirectly connected via a thin suture (thread-like process)(Fig. 4b). The most distinctive plate of this species is the I' plate: a) it bears a large characteristic ventral pore; and b) a 90 degree angle is formed at the point where the ventral pore and the 4' plate come in contact (Figs. 2,3). The distinctive sixth precingular plate (6") is wider than high (Figs. 2,3)(Balech 1995; Balech & Tangen 1985; Larsen & Moestrup 1989; Taylor et al. 1995). The broad epitheca is convex-conical, while the hypotheca is hemispherical with an obliquely flattened antapex (Figs. 1,5). The slightly excavated cingulum is equatorial and displaced in a descending fashion less than one time its width: it has narrow lists (Figs. 1,3). The sulcus is slightly depressed and inconspicuous (Balech 1995; Balech & Tangen 1985; Konovalova 1993; Larsen & Moestrup 1989; Taylor et al. 1995). Morphology and Structure: A. ostenfeldii is a photosynthetic species with radiating 16 Harmful Marine Dinoflagellates chloroplasts. The nucleus is U-shaped and equatorial (Fig. 5)(Balech & Tangen 1985). Reproduction: A. ostenfeldii reproduces asexually by binary fission. This species also has a sexual cycle with isogamous mating types; a planozygote is formed (Jensen & Moestrup 1997), Ecology: A. ostenfeldii is a planktonic estuarine dinoflagellate species found in low numbers, mainly along the west coast of Europe, and recently along the southeast coast of Nova Scotia, Canada (Cembella et al. 2000). To date, no blooms have been reported (except in Belgium as Pyrodinium phoneus (Woloszynska & Conrad 1939; Hansen et al. 1992), This species produces temporary resting cysts (Fig. 6). Cysts are large and spherical, ranging in size from 35 to 40 um in diameter. Cysts are pale in color with a reddish-brown granule, and a well-defined cingular groove. The smooth and clear cell wall is covered with mucilage (Mackenzie et al. 1996; Jensen & Moestrup 1997). Toxicity: There has long been some doubt as to the toxic potential of this species (Balech 1995; Hansen et al. 1992). Because A. ostenfeldii does not form monospecific blooms, it has been difficult to determine this species’ toxin producing potential. A. ostenfeldii, however, is capable of producing paralytic shellfish poison (PSP) toxins; albeit, it is the least toxic of all the Alexandrium species tested for PSP toxins (Cembella et al. 1987; 1988). This species has been associated with shellfish poisoning in Scandinavia (Jensen & Moestrup 1997), and one report of mussel toxicity (as Pyrodinium Phoneus) has been reported from Belgium (Woloszynska & Conrad 1939), Recently, a study of aquaculture shellfish from Nova Scotia, Canada, revealed the presence of spirilides, fast-acting neurotoxins, primarily produced by western Atlantic strains of A. ostenfeldii (Cembella et al. 2000). Hansen et al. (1992) conducted studies with a tintinnid ciliate exposed to high concentrations of A. ostenfeldii: results were erratic swimming behavior (backwards) followed by swelling and lysis of the ciliates. Species Comparison: A. ostenfeldii is easily misidentified as other A/lexandrium species; detailed thecal plate observation is often necessary for proper identification (Balech 1995; Larsen & Moestrup 1989). A. ostenfeldii and A. tamarense are often confused for each other since they overlap in size and often co-occur; however, 4. ostenfeldii is slightly larger and is more widely distributed (has a wider salinity range) than the latter species (Moestrup & Hansen 1988). Other differences between these two species include: A. ostenfeldii has a much larger ventral pore on the first apical plate 1'; and the 6" plate is wider than high, whereas the width and height of the 6" plate in A. famarense are equal (Balech 1995; Hansen et al. 1992). This species also closely resembles another Alexandrium species, A. peruvianum, — Both species are large cells with distinctive large ventral pores on the I|' plate; however, morphological differences are evident in the I' plate and the APC. Moreover, A. ostenfeldii is a larger cell and produces PSP toxins (Balech 1995; Steidinger & Tangen 1996; Taylor et al. 1995). Habitat and Locality: A cold-water estuarine species, A. ostenfeldii was, until recently, believed to be confined to the western european coast: Iceland and Norway (Paulsen 1904; Braarud 1945; Balech & Tangen 1985), Denmark (Moestrup & Hansen 1988), Belgium (as Pyrodinium phoneus (Woloszynska & Conrad 1939), and Spain (Fraga & Sanchez 1985). Recently, Balech (1995) collected cells of A. ostenfeldii from Alexandria Harbor, Egypt, and also from the NW Pacific Ocean, off of Washington State, U.S.A. Populations have also been observed from British Columbia and the Kamchatka Peninsula in the Pacific Ocean (Konovalova 1993; Steidinger & Tangen 1996; Taylor et al. 1995). In the northwest Atlantic Ocean, cells have been reported from Canada: in the Gulf of St. Lawrence (Levasseur et al. 1998), and southeastern Nova Scotia (Cembella et al. 2000). Remarks: Belonging to the Alexandrium complex, A. ostenfeldii has a long and complex taxonomic history. Harmful Marine Dinoflagellates 17 Alexandrium pseudogonyaulax (Biecheler) Horiguchi ex Yuki et Fukuyo, 1992 Plate 6, Figs. 1-9 Alexandrium armoured, marine, It is a toxic species Species Overview: pseudogonyaulax 1s an planktonic dinoflagellate. found in coastal regions and __ brackish environments. Taxonomical Description: Cells of A. pseudogonyaulax are medium to large, irregularly pentagonal-shaped with moderate dorso-ventral flattening. Cells are wider than long; the epitheca is slightly shorter than the hypotheca (Figs. 1,2). The first apical plate (1') is characteristically displaced with a large ventral pore on the anterior margin (Figs. 3-5). The thecal plates are smooth and thin with scattered minute pores. Cells range in size between 34-60 um in length and 39-69 um in width (Balech 1995: Montresor et al. 1993; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Goniodoma pseudogoniaulax Biecheler, 1952: 55, figs. XXX-XXXI] Type Locality: Mediterranean Sea: Thau Lagoon, France Synonyms: Goniodoma pseudogoniaulax Biecheler, 1952 Alexandrium — pseudogonyaulax _ (Biecheler) Horiguchi, 1983 Thecal Plate Description: The plate formula for A. pseudogonyaulax is: Po, 4', 6", 6c, 10s, 5", 2". The apical pore plate (Po) is oval shaped, contains a large comma-shaped foramen and a number of irregular pores, and is positioned longitudinally on the apex (Figs. 3,4,6). The distintive 1' plate does not come in contact with the Po (Figs. 3,4,6); it is roughly pentagonal and wider anteriorly (Figs. 3,6). The sloped anterior margin bears a large ventral pore that is wider than long (Figs. 3,4,6). The ventral pore does not penetrate the 4' plate (Balech 1995; Montresor et al. 1993; Yuki & Fukuyo 1992). The short, convex epitheca is dome-shaped (Figs. 1,2). The hypotheca is slightly longer with an obliquely concave antapex (Figs. 1,2). The shallow cingulum is displaced in a descending fashion less than one time its width (Fig. 5). The sulcus lacks lateral lists. It slightly penetrates the epitheca obliquely on the right (Balech 1995). Morphology and Structure: A, pseudogonyaulax is a photosynthetic species with central radiating yellow-brown chloroplasts. The transversely elongated nucleus is large and curved, and centrally located (Balech 1995; Montresor 1995). Reproduction: 4. pseudogonyaulax reproduces asexually by binary fission. This species also has a sexual cycle with isogamous mating types. The smaller rounder gametes (Fig. 7) fuse (one gamete engulfs the other), produce a planozygote which then encysts into a characteristic resting cyst (Fig. 8)(Montresor et al. 1993; Montresor 1995). Ecology: 4. pseudogonyaulax is a coastal and brackish water dinoflagellate species. Blooms of this species are commonly reported in the Strait of Georgia, British Columbia (North Pacific Ocean)(Taylor & Haigh 1993). This species produces a characteristic and unusual resting cyst: a non-smooth cyst. The cysts are round and dark, and are often covered with a mucilaginous layer (Fig. 8). They contain a reddish-orange accumulation body. — Size ranges from 40 to 55 ym in diameter. The cyst wall consists of two layers: a smooth inner layer and a paratabular outer layer (Fig. 9). The cyst paratabulation equals the tabulation of a vegetative cell. This is the only reported species in the genus Alexandrium to produce a non- smooth cyst (Montresor et al. 1993; Nichetto et al. 1995). Toxicity: 4. pseudogonyaulax produces a unique phycotoxin, goniodomin A (GA), that has an antifungal effect (Murakami et al. 1988). The toxin GA targets the liver and thymus (Terao et al. 1989; 1990). Species Comparison: 4. pseudogonyaulax closely resembles two other Alexandrium species: 4. hiranoi and A. satoanum. Common features include general shape and size, and lack of contact of the first apical plate, 1', with the Po. 18 Harmful Marine Dinoflagellates Distinguishing features lie in the cell outline, the ventral pore, the |' plate, cyst morphology and habitat: a) A. Airanoi has a round shape, A. pseudogonyaulax is wider than long, A. satoanum ts also wider than long with the general outline resembling a top: the epitheca and hypotheca have straighter sides; b) the ventral pore of A. hiranoi is circular and invades the 4' plate, in A. pseudogonyaulax the ventral pore is semi-circular and does not invade the 4', and in A, satoanum, no ventral pore is present (has a.a.p. and p.a.p); c) the I’ plate in 4. hiranoi is Slender and rectangular, whereas in 4. pseudogonyaulax the |' is almost pentagonal; d) the cyst of 4. hiranoi is smooth, while the cyst of A. pseudogonyaulax is paratabulate with thick sutures; and e) A. hiranoi is found in rockpools, A, pseudogonyaulax is found in coastal brackish habitats (Kita & Fukuyo 1988; Montresor et al. 1993; Steidinger & Tangen 1996), This species roughly resembles 4. tamarense, however the latter species is not as round, and has a broader APC (Taylor et al. 1995). Habitat and Locality: 4. psewdogonyaulax is a coastal species which has been reported from several localities in Europe: France along the Mediterranean coast (Biecheler 1952), Italy in the Gulf of Trieste, North Adriatic Sea (Honsell et al. 1992; Montresor et al. 1993; Nichetto et al. 1995), Portugal and Norwegian fjords (Balech 1995). In the Pacific Ocean this species is a common bloom former in the Gulf of Georgia in British Columbia (Taylor & Haigh 1993), and populations have been observed in coastal waters of Japan (Inoue, in Kita & Fukuyo 1988). Alexandrium tamarense (Lebour) Balech, 1985 Plate 7, Figs. 1-6 Species Overview: Alexandrium tamarense is an armoured, marine, planktonic dinoflagellate. It is associated with toxic PSP blooms in cold water coastal regions. Taxonomical Description: Cells of 4. famarense are small to medium in size, nearly spherical, and slightly longer than wide (Fig. 1). The first apical plate bears a ventral pore (Figs. 3,5). Cells are commonly found single or in pairs (Figs. 1-3), and less commonly in fours. Paired cells may contain an anterior attachment pore (aap) and a posterior attachment pore (pap)(Fig. 4). Thecal plates are smooth and thin (Fig. 3). The size and shape of this species is highly variable: cells range in size between 22-51 um in length and 17-44 um in transdiameter width (Lebour 1925; Fukuyo et al. 1990; Hallegraeff 1991; Hallegraeff et al. 1991; Larsen & Moestrup 1989; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gonyaulax tamarensis Lebour, 1925: 92, plate XIV, figs. la-Id Type Locality: English Channel: River Tamar Estuary, near Plymouth, United Kingdom Synonyms: Gonyvaulax tamarensis Lebour, 1925 Gonvaulax tamarensis var. excavata Braarud, 1945 Gonyaulax excavata (Braarud) Balech, 1971 Gessnerium tamarensis (Lebour) Loeblich and Loeblich, 1979 Protogonyaulax tamarensis (Lebour) Taylor, 1979 Alexandrium excavatum (Braarud) Balech and Tangen, 1985 Thecal Plate Description: The plate formula for A. tamarense is: Po, 4', 6", 6c, 8s, 5", 2". The apical pore complex (APC) is rectangular and narrows ventrally (Fig. 3). The apical pore plate (Po) houses a large fishhook shaped foramen and a small round aap (Figs. 3,4). The first apical plate (I') is variable in shape: from a broad triangle to a narrow rectangle, and bears a small ventral pore (Figs. 3,5). The 1' plate comes in direct contact with the Po (Fig. 3)(Lebour 1925; Fukuyo et al. 1985; 1990; Larsen & Moestrup 1989; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca and hypotheca are nearly equal in height (Figs. 1,2,5). The epitheca is broadly conical, while the hypotheca is roughly trapezoidal (Figs. 1,2,5). The posterior end is slightly indented resulting in two hypothecal lobes; the left lobe is slightly larger than the right (Figs. 1,2). The deeply excavated cingulum is displaced in a descending fashion one time its width with narrow lists (Figs. 2,5). The deep sulcus, with lists, widens posteriorly (Figs. 2,5). Harmful Marine Dinoflagellates 19 The posterior attachment pore (pap), if present, is small and located in the right half of the posterior sulcal plate (Lebour 1925; Fukuyo et al. 1985; 1990; Larsen & Moestrup 1989; Balech 1995; Taylor et al. 1995; Steidinger & Tangen 1996). Morphology and Structure: A. tamarense is a photosynthetic species with a number of orange- brown chloroplasts. A lunar-shaped nucleus Is situated ventrally just inside the cingulum (Fig. 1)(Fukuyo 1985; Larsen & Moestrup 1989). Reproduction: 4. famarense — reproduces asexually by binary fission; plane of fission is oblique. This species also has a sexual cycle with anisogamous mating types. The gametes join laterally for sexual fusion, produce a planozygote which then encysts into a characteristic resting cyst (Fig. 6)(Loeblich & Loeblich 1975; Turpin et al. 1978; Silva 1962). Ecology: 4. tamarense is a planktonic dinoflagellate species associated with toxic paralytic shellfish poisoning (PSP) events around the world. Toxic blooms are commonly reported in Japan (Fukuyo et al. 1985; Ogata et al. 1982; Oshima et al. 1982). Red tide blooms of 4A. tamarense have been reported in Europe (Mortensen 1985; Moestrup & Hansen 1988), and are common along the NE coast of North America (New England and Canada)(Bicknell & Walsh 1975; Hurst 1975; Loeblich & Loeblich 1975). During a red tide event reported in the Faroe Islands, Norway, in 1984, population levels of 4. tamarense were estimated at 1 X 10’ cells/L and completely dominated the plankton (Mortensen 1985; Moestrup & Hansen 1988). This species produces a ellipsoidal resting cyst that cannot be distinguished from the cyst produced by 4. catenella. This cyst has rounded ends with a thick cell wall, and is covered in mucilage (Fig. 6). Cysts often contain colorless granules and distinct reddish lipid bodies. Size ranges from 36-56 um in length and 23-32 um in width (Turpin et al. 1978; Fukuyo 1985; Bolch & Hallegraeff 1990; Hallegraeff 1991; Hallegraeff et al. 1991). Toxicity: A/exandrium tamarense is a known toxin-producing dinoflagellate species. This species produces very potent paralytic shellfish poison (PSP) neurotoxins which can affect humans, other mammals, fish and birds (Larsen & Moestrup 1989): gonyautoxins (GTX1, GTX2, GTX3, GTX4 and GTXS), neosaxitoxin (NSTX) and saxitoxin (SXT)(Shimizu et al. 1975; Oshima et al. 1977). This species is responsible for numerous human illnesses and several deaths after consumption of infected shellfish: ten deaths in Venezuela in 1977 (Reyes-Vasquez et al. 1979), and one death in Thailand in 1984 (Tamiyavanich et al. 1985). Resting cysts of 4. famarense can also harbor PSP toxins. Dale et al. (1978) demonstrated that cysts were more than ten times as toxic as their motile stage counterparts. Not all strains of A. tamarense are toxic: both toxic and nontoxic strains have been reported in New England within the same red tide event (Yentsch et al. 1978). Strains in Australia (Hallegraeff 1991), River Tamar estuary, Britain (type locality)(Moestrup & Hansen 1988) and the Gulf of Thailand (Fukuyo et al. 1988) are all non-toxic. The usual route of PSP toxin transmission is via contaminated shellfish; however, bloom events of A. tamarense have been linked to several massive fish kills: Atlantic herring in the Bay of Fundy, Canada (White 1980); and rainbow trout and salmon in the Faroe Islands, Norway (Mortensen 1985). Hayashi et al. (1982) attribute the fish kills to dinoflagellate toxins accumulated in the food chain; i.e. fish feed on zooplankton infected with PSP poisons and die. However, Ogata and Kodama (1986) report production of ichthyotoxins in cultured media of this species. Species Comparison: A. famarense can resemble a number of other species within the genus, but it can be distinguished by its cell shape and size, presence of a ventral pore (vp) on the |' plate, and shape of the thecal plates (Balech 1995; Hallegraeff 1991; Larsen & Moestrup 1989; Steidinger & Tangen 1996). A. tamarense is very similar morphologically (size, shape and thecal plate formula) to 4. catenella; both also produce deadly PSP toxins. Morphological differences lie in the shape of the Po, and presence or absence of a vp: the Po in 4. catenella is slightly smaller than that in 4. tamarense, and the vp is absent (Fukuyo 1985). Molecular testing conducted on A. catenella from Japan and 4. tamarense from Japan and the 20 Harmful Marine Dinoflagellates U.S.A. revealed a close genetic relationship between the two species, however they remain distinct (Adachi et al. 1995). Morphologically, A. fundyense is nearly identical to A. tamarense except for the missing ventral pore on the |' plate. 4. minutum can also be misidentified as 4. tamarense; however, A. tamarense is a smaller species, is always longer than wide, and is found in colder waters than A. minutum (Balech 1995; Hallegraeff 1991; Larsen & Moestrup 1989; Steidinger & Tangen 1996), Habitat and Locality: 4. tamarense is a widely distributed coastal and estuarine dinoflagellate species (Lebour 1925; Steidinger & Tangen 1996) mainly found in cold to cold-temperate waters in North America, Europe and Japan. However, this species has been reported from warmer waters around the world: Australia, Venezuela and the Gulf of Thailand (Balech 1995; Fukuyo et al. 1990; Hallegraeff 1991; Steidinger & Tangen 1996; Taylor et al. 1995). Alexandrium tamiyavanichi Balech, 1994 Plate 8, Figs. 1-6 Species Overview: Alexandrium tamiyavanichi is an armoured, marine, planktonic dinoflagellate. It is a producer of strong PSP toxins in the Gulf of Thailand. Taxonomical Description: A chain-forming species, A. tamiyavanichi typically occurs in chains of 8 cells or more. Single cells are small and round to slightly wider than long (Figs. 1,2). A small ventral pore (vp) is present on the first apical plate (1°)(Figs. 3-5). The thecal plates are thin and strongly porulated. Cells range in size between 31-41 um in length and 26-35 um in transdiameter width (Balech 1995; Fukuyo et al. 1989; Taylor et al. 1995), Nomenclatural Types: Holotype: Alexandrium tamivavanichi Balech, 1994; 217-219, figs. 1-6 Type Locality: Gulf of Thailand: Ang. Sila, Thailand Synonyms: Protogonyaulax cohorticula (Balech) Taylor, sec Kodama et al. (1988); non Gonyaulax cohorticula Balech, 1967 Thecal Plate Description: The plate formula for A, tamiyavanichi is: Po, 4', 6", 6c, 10s, 5", 2". The broad apical pore complex (APC) is triangular and narrows ventrally (Figs. 3,4). The apical pore plate (Po) is wide and oval with a large comma-shaped foramen (Figs. 3,4). Several small pores are present along the margin of the Po (Fig. 4). The anterior attachment pore (aap) is large, round and adjacent to the Po (Fig. 4). The 1’ plate is large and wide with straight sides, and is in direct contact with the Po (Figs. 3-5). A small ventral pore is present on the anterior right margin of this plate (Figs. 3- 5)(Balech 1967; 1995; Fukuyo et al. 1989; Taylor et al. 1995). The conical epitheca is wider than long with shoulders (Figs. 1,2). The hypotheca is slightly longer than the epitheca (Figs. 1,2). The deeply excavated cingulum is displaced in a descending fashion one time its width (Figs. 2,5). The sulcus is deep and widens posteriorly (Figs. 2,4,5). Two wing-like sulcal lists project anteriorly toward the antapex yielding two antapical spines (Figs. 1,5). The sulcus invades the epitheca via the distinctive anterior sulcal plate (s.a.); this plate is divided into two parts by a transverse rib (Fig. 4). It is the anterior extension of the s.a. plate which projects into a notch in the epitheca (Figs. 2,4,5).. The round posterior attachment pore, pap, is present in the center of the posterior sulcal plate (Fig. 6)(Balech 1967; 1995, Fukuyo et al. 1989; Taylor et al. 1995). Morphology and Structure: 4. tamiyvavanichi is a photosynthetic species. The transversely elongated nucleus is lunate shaped (Balech 1995). Reproduction: A. tamivavanichi reproduces asexually by binary fission. Ecology: 4. tamiyavanichi is a_ coastal planktonic species (Balech 1994). Toxicity: 4. tamivavanichi produces potent paralytic shellfish poison (PSP) toxins similar to those produced by 4. tamarense: gonyautoxins Harmful Marine Dinoflagellates 21 (GTX), and saxitoxin (SXT)(Fukuyo et al. 1989; Kodama et al. 1988). This species has been the main causative organism of PSP in Thailand waters (Kodama et al. 1988). Etymology: This species, ‘tamivavanichi’, was named in honor of Prof. Suthichai Tamiyavanich, researcher in red tides and toxic dinoflagellates in Thailand (Balech 1994; 1995). Species Comparison: 4. tamivavanichi is often and easily misidentified as A. cohorticula: cell size and outline is similar, both with an anterior extention of the s.a. plate, and both species are chain formers. However, there are number of substantial morphological differences between these two species: In 4. cohorticula, the epitheca is longer than wide; the Po is longer; the first apical plate, I', is thinner; the pap is larger and oval shaped; and the sulcal lists are larger and projected behind the hypotheca (Balech 1995). Chains of A. tamivavanichi can resemble A. catenella. The epitheca in A. tamiyavanichi, however, is conical in comparison to the rounded epitheca of A. catenella (Taylor et al. 1995). Habitat and Locality: 4. tamivavanichi is a coastal species that has only been reported from three warm-water localities: Gulf of Thailand (type locality), Manila Bay in the Philippines, and from the Andaman Sea, southwest of Thailand (Balech 1995). Cochlodinium polykrikoides Maregelef, 1961 Plate 9, Figs. 1-7 Species Overview: Cochlodinium polykrikoides is. an unarmoured, marine, — planktonic dinoflagellate species with a distinctive spiral- shaped cingulum. It is a common red tide former associated with fish kills in Japan and Korea. Taxonomic Description: Cochlodinium polykrikoides is an athecate species; i.e. without thecal plates. Cells are small, oval and slightly flattened dorso-ventrally (Figs. 1,2). Chains, rarely more than eight cells, are common (Figs. 1-4). An apical groove is present on the apex originating from the anterior end of the cingular and sulcal juncture and extending to the dorsal side of the epitheca. Cells range in size from 30- 40 um in length to 20-30 um in width (Silva 1967; Yuki & Yoshimatsu 1989; Fukuyo et al. 1990; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca is conical and rounded at the apex (Figs. 1,2,4). The hypotheca is bilobed (Fig. 1). The cingulum is deep and excavated (Figs. 1.2.4). It is displaced about 0.6 times the cell length, and descends in a distinct left-handed spiral of 1.8-1.9 turns around the cell. The narrow and shallow sulcus nearly runs parallel to the cingulum making 0.8-0.9 turns around the cell between the proximal and distal ends of the cingulum. The sulcus deepens and widens towards the antapex and divides the hypotheca into two asymmetrical lobes (Fig. 1). The right lobe is narrower and slightly longer than the left lobe (Silva 1967; Yuki & Yoshimatsu 1989; Fukuyo et al. 1990; Taylor et al. 1995; Steidinger & Tangen 1996). Trichocysts have been observed in this species, but the number per cell varies, and not all cells bear them. The presence and number of trichocysts increases with cell and culture age (Silva 1967). Nomenclatural Types: Holotype: Cochlodinium polvkrikoides Margelef, 1961: 76, fig. 27 Type Locality: Caribbean Sea: Puerto Rico Synonyms: Cochlodinium heterolobatum Silva, 1967 Morphology and Structure: C. polykrikoides is a photosynthetic species with numerous yellowish-green to brown chloroplasts, rod- shaped or ellipsoid in shape (Fig. 1). The nucleus is situated anteriorly in the epitheca (Figs. 2,4). A red stigma is present dorsally in the epitheca (Silva 1967; Yuki & Yoshimatsu 1989; Fukuyo et al. 1990; Taylor et al. 1995). Reproduction: CC. polykrikoides reproduces asexually by binary fission; plane of fission 1s oblique (Silva 1967). Ecology: C. polykrikoides is a_ planktonic species. It is a common ichthyotoxic 'red water’ bloom species in the northwestern Pacific. This species commonly forms cysts (Figs. 5-7) (Fukuyo et al. 1990; Taylor et al. 1995; Steidinger & Tangen 1996). 22 Harmful Marine Dinoflagellates Toxicity: Cochlodinium polykrikoides is a known red tide species associated with extensive fish kills and great economic loss in Japanese and Korean waters (Yuki & Yoshimatsu 1989; Fukuyo et al. 1990; Kim 1998). However, the actual toxin principles have yet to be ellucidated (Taylor et al. 1995). Ho and Zubkoff (1979) suggested that physical contact, not a released toxin, was the cause of oyster larvae (Crassostrea virginica) deformation — and mortality during a C. polykrikoides red tide in the York River (Virginia, USA). Species Comparison: C. polvkrikoides closely resembles two other Cochlodinium species: C. helix and C. helicoides. The degree of rotation of the cingulum and sulcus distinguish the former species from the latter two: a. the cingulum in C. polvkrikoides makes 1.8-1.9 turns around the cell, while in C. helix it is two turns and in C. helicoides it is 1.5 turns; and b. the sulcus turns 0.8 times between the proximal and distal ends of the cingulum in C. polvkrikoides, whereas it is | time in C. helix and 0.6 times in C. helicoides (Silva 1967). Habitat and Locality: C. polykrikoides is a cosmopolitan species found in warm temperate and tropical waters (Steidinger & Tangen 1996), This species was first reported from the Caribbean Sea along the southern coast of Puerto Rico (Margelef 1961). It has since been reported in northern Atlantic waters along the American east coast: Barnegat Bay, New Jersey (Silva 1967), and the York River, Virginia (Ho & Zubkoff 1979; Zubkoff et al. 1979). It is widely distributed in northwestern Pacific waters along the coasts of Japan and Korea (Fukuyo et al. 1990; Kim 1998). Coolia monotis Meunier, 1919 Plate 10, Figs. 1-8 Species Overview: Coolia monotis is an armoured, marine, benthic dinoflagellate species. It a toxic species with world-wide distribution. Taxonomic Description: Species in this genus are anterio-posteriorly compressed and = are observed in apical or antapical view. 30 0/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 fill 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. breve-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 al. 1973; Baden et al. 1982; Taylor et al. 1995). Harmful Marine Dinoflagellates 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. breve-like 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). Gymnodinium catenatum Graham, 1943 Plate 23, Figs. 1-7 Species Overview: Gymnodinium catenatum 1s 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 catenatum 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. 1)(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 pm in width to 34-65 um in length. Chain-forming cells are slightly smaller with sizes ranging from 27-43 um in width to 23-60 ym 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 catenatum Graham, 1943: 259-262, figs. 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 reproduces asexually by binary fission. 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 deficiency 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 | 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 um 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 Dinoflagellates Cysts are not only a reseeding tool, but also a disbursement agent: G. catenatum — was introduced to Australian waters via ships! ballast water (Hallegraeff & 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 dinoflagellate 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 misidentified. 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; Hallegraeff 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 (Hallegraeff 1991), Gymnodinium mikimotoi 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 um in length to 14-35 um in width (Takayama & Adachi 1984; Fukuyo et al. 1990; Hallegraeff 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 times its width (Figs. 1,5). The sulcus slightly invades the epitheca extending from above the cingulum to the antapex (Figs. 1,5)(Takayama & Adachi 1984; Fukuyo et al. 1990; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Gymnodinium mikimotoi Oda, 1935: 35-48, figs. 1-3 Type Locality: NW Pacific Ocean: Gokasho Bay, Japan Synonyms: Gymnodinium Adachi, 1984 Gyrodinium aureolum Hulburt, sensu Braarud and Heimdal, 1970 nagasakiense Takayama and 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; Hallegraeff 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Reproduction: G. = mikimotoi — reproduces asexually by binary fission; cells divide obliquely Harmful Marine Dinoflagellates 4] 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 warmer 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 shellfish in Southland, NZ (dubbed ‘Southland toxin') after a Gymnodinium ct. 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 fish 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 flatter (dorso-ventral 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 aureolum 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. awreolum 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 pulchellum 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 um in length to 11-16 um 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 I-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). Nomenclatural Types: Holotype: Gymnodinium pulchellum 1994: 32, fig. 58 Type Locality: Tasman Sea: Melbourne, Australia Synonyms: Gymnodinium type '84-K Onoue et al., 1985 Larsen, Hobsons Bay, Morphology 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 asexually by binary fission. reproduces 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 fish 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 Gymnodinium 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 fish 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 ‘pu/chellum’ 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 sanguineum Hirasaka, 1922 Plate 26, Figs. 1-4 Species Overview: Gymnodinium sanguineum is an unarmoured, marine, planktonic dinoflagellate species. This cosmopolitan species is a red tide former that has been associated with fish and shellfish mortality events. Taxonomic Description: Gymnodinium sanguineum 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 um 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 cingulum is left-handed and displaced I- 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; Fukuyo 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 (Hallegraetf 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 0/00). 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 & Coats 1993; Voltolina 1993). Gymnodinium veneficum Ballantine, 1956 Plate 27, Figs. 1-3 Species Overview: Gymnodinium veneficum is an unarmoured, marine, planktonic dinoflagellate 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 um in length to 7-14 um in width (Ballantine 1956; Dodge 1982; Taylor et al. 1995). The epitheca and hypotheca 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, figs. 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 et al. 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 et al. 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 galatheanum (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. 1). Cells range in size from 9-17 um in length to 8-14 um in width (Braarud 1957; Taylor et al. 1995; Steidinger & Tangen 1996). The epitheca and hypotheca are both round (Figs. |-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 flagellum (Figs. 1-3). The short and narrow sulcus slightly invades the epitheca adjacent to the apical groove (Figs. 1,2,4)(Braarud 1957; Taylor et al. 1995; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: 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, II] Woloszynskia micra Leadbeater and Dodge, 1966 Harmful Marine Dinoflagellates 45 Basionym: Gymnodinium galatheanum Braarud, 1957 Morphology and Structure: G. ga/atheanum is a photosynthetic species with several round chloroplasts. 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 aureolum, 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 et al. 1995). Lingulodinium polyedrum (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 of Lingulodinium polyedrum are angular, roughly pentagonal and polyhedral-shaped (Fig. 1). Cells range in size from 40-54 um in length and 37-53 um 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: Gonyaulax polyedra Stein, 1883: p. 13, pl. 4, figs. 7-9 Type Locality: unknown Synonyms: Gonyaulax polyedra Stein, 1883 Lingulodinium machaerophorum (Deflandre and Cookson) Wall, 1967 (cyst) Hystrichosphaeridium machaerophorum Deflandre and Cookson, 1955 (cyst) Thecal Plate Description: The plate formula for L. polvedrum 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, comes in direct contact with the Po, and bears a ventral pore on its right side (Figs. 1,2,4). The deeply excavated cingulum is nearly equatorial, and displaced one to two times its width. It 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. 1,2,4)(Kofoid 1911; Dodge 1985: Dodge 1989; Lewis & Burton 1988; Fukuyo et al. 1990; Steidinger & Tangen 1996). Morphology and Structure: L. polvedrum 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 1971; Jeffrey et al. 1975). Reproduction: = Ll. — polyedrum reproduces asexually 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 1911). 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 et al. 1990). This species forms colorless spherical spiny cysts (35-50 um in diameter). The numerous tapering spines can reach up to 17 um in length, all 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 dinoflagellate cyst) Lingulodinium machaerophorum (Deflandre and Cookson) Wall, 1967 was discovered to be the resting spore of L. polvedrum (Wall 1967; Fensome et al. 1993). Marasovic (1989) reported production of temporary resting cysts in a waning red tide dominated by L. polvedrum 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. pol/yedrum is a widely distributed species found in warm temperate and subtropical waters of coastal areas (Kofoid 1911; Dodge 1985; 1989; Steidinger & Tangen 1996). Noctiluca scintillans (Macartney) Kofoid et Swezy, 1921 Plate 30, Figs. 1-4 Species Overview: Noctiluca scintillans is an unarmoured, marine planktonic dinoflagellate species. This large and distinctive bloom forming species has been associated with fish and marine invertebrate mortality events. Taxonomic Description: Noctiluca 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 flagellum 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 um 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 scintillins Macartney, 1810: 264-265, pl. 15, figs. 9-12 Type Locality: North Sea: Herne Bay, Kent, England Synonyms: Medusa scintillins Macartney, 1810 Noctiluca miliaris 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: Nocéiluca 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. N. 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 & 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; Hallegraetf 1991; Taylor et al. 1995; Steidinger & Tangen 1996). Remarks: This species is frequently referred to as N. miliaris 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 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. 1)(Faust et al. 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 tm) that can only be observed at the SEM level (Figs. 1,2). Cells have a dorsoventral diameter of 80-108 um, and a transdiameter of 46-59 um (Faust et al. 1996). Nomenclatural Types: Iconotype: Ostreopsis heptagona Norris, Bomber and Balech, 1985: fig. 1 Type Locality: Gulf of Mexico: Knight Key, Florida, USA Thecal Plate Description: The plate formula of Ostreopsis heptagona is: Po, 3', 7", 6c, 68?, Vp, Rp, 5", Ip, 2""(Fig. 5). The epitheca contains 11 plates. The apical pore plate (Po) is 15 um long, narrow and curved (Figs. 1,3), situated between apical plates 1', 2' and 3', with a long, slit-like apical pore. The I' 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 (lp) 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 et al. 1985). The cingulum is equatorial and narrow (Figs. 1-3). Within the cingulum the Vo ts 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. hepfagona reproduce asexually 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) |' plate that occupies the left center of the epitheca (this plate is hexagonal, six-sided, in all other species of this genus) (Faust et al. 1996; Steidinger & Tangen 1996); and b)_ the pentagonal and dorso-ventrally elongate Ip plate in the hypotheca (Faust et al. 1996). In O. heptagona plate 5" is pentagonal as it contacts plates 1', 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 |', while 6" is hexagonal and contacts two apical plates, |' 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: |p 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 lenticularis 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 Gambier 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. 1)(Faust et al. 1996). Cells of Ostreopsis lenticularis are lenticulate to broadly oval (Figs. 1,2). The cell surface is smooth and covered with randomly spaced pores (0.4 um 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 um and a transdiameter of 57-63 tum (Faust et al. 1996; Fukuyo 1981), Nomenclatural Types: Holotype: Ostreopsis lenticularis Fukuyo, 1981: figs. 30-34 Type Locality: South Pacific Ocean: Gambier and Society Islands, and New Caledonia Thecal Plate Description: The plate formula of Ostreopsis lenticularis is: Po, 3', 7", 6c, 68?, Vp, Rp, 5", lp, 2'""(Fig. 6). The epitheca contains 11 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 I|' plate is large, irregularly pentagonal-shaped, and situated in the center (Figs. 1,5)(Faust et al. 1996). The hypotheca ts 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 lenticularis 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 lenticularis reproduces asexually by binary fission. Ecology: O. Jenticularis 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 (Ballantine et al. 1988). Species Comparisons: Ostreopsis /enticularis 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. Jenticularis has a_ slightly pointed ventral area while G. foxicus has a round and indented one (Fukuyo 1981). O. lenticularis is also similar to O. siamensis in shape and thecal plate configuration (Fukuyo 1981). Habitat and Locality: Populations of O. lenticularis were originally found in the Gambier 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 & Tindall 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 first apical plate (1') on the epitheca (Fig. 1)(Faust et al. 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 um and a transdiameter of 118-134 um. 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 um) with smooth edges (Fig. 3)(Quod 1994; Faust et al. 1996). Nomenclatural Types: Holotype: Ostreopsis 1994; fig. | Type Locality: West Indian Ocean: Saint Leu, Reunion Island, Mascareignes Archipelago Quod, mascarenensis 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 um) with an array of minute openings (Fig. 4). The 1' plate is large, long and hexagonal, 102 um long and 40 um wide (Fig. 1). 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 et al. 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: QO. 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 1',2',3',1" and Ip (Quod 1994), Habitat and Locality: Populations of O. mascarenensis can be commonly found in Harmful Marine Dinoflagellates S| shallow (2-Sm) barrier reef environments and coral reefs in the SW Indian Ocean. This species has been observed as an epiphyte on Turbinaria sp., Galaxaura 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 dinoflagellate 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 (1') on the epitheca (Fig. 1)(Faust et al. 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 um diameter)(Figs. 1-4). Cells have a dorsoventral diameter of 47-55 um = and transdiameter of 27-35 um (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, 68?, Vp, Rp, 5", lp, 2"". In the epitheca, the |' 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 um) (Fig. 2) (Faust et al. 1996). 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 (Nakajima et 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 1s 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 dinoflagellate species. It was first identified 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 first apical plate (1') on the epitheca (Fig. 1)(Faust et al. 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. |-3). Large (0.5 um diameter) and small (0.1 um diameter) pores are present (Fig. 4). Cells have a dorsoventral diameter of 108-123 um and a transdiameter of 76-86 um (Faust et al. 1996). Nomenclatural Types: Holotype: Ostreopsis siamensis Schmidt, 1902: figs. 5-7 Type Locality: Gulf of Thailand: Thailand Thecal Plate Description: The plate formula for Ostreopsis siamensis is: Po, 3', 7", 6c, 68?, 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 um long (Fig. 3). The |' 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 um), 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 I["" 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 asexually by binary fission. reproduces 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 (Nakajima et al. 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; Ballantine et al. 1988; Faust 1995; Faust & Morton 1995). They have also Harmful Marine Dinoflagellates 53 been associated with sand in the Caribbean (Faust et al. 1996). Pfiesteria piscicida Steidinger et Burkholder, 1996 Plate 36, Figs. 1-9 Species Overview: Pfiesteria piscicida is a putatively toxic dinoflagellate 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 et al. 1996). Taxonomic Description: Pfiesteria piscicida is a polymorphic and multiphasic dinoflagellate species with a number of unicellular stages throughout its life cycle: bi- and triflagellated 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 biflagellated stages, zoospores, have two size groups: 5-8 um (gametes) and 10-18 um (Fig. 3). The larger triflagellated stage, 25-60 um, is a planozygote with the features of a vegetative cell along with one transverse and two longitudinal flagella (Fig. 4). Cyst stages, with highly resistant cell walls, range in size from 25-33 um (Fig. 5). The flagellated forms are typically planktonic and ephemeral, whereas the cyst stages are benthic (Steidinger et al. 1996). Nomenclatural 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 piscimortuis Burkholder et al., 1995 "ohantom dinoflagellate" 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 of P. 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 (1') is rhomboid in shape (Fig. 6). The broad and shallow cingulum is without lists, and descends almost | 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 et al. 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 algal 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 et al. 1996; Lewitus et al. 1999). Reproduction: Biflagellated zoospores reproduce asexually via temporary cysts. Sexual reproduction has also been documented for this species: biflagellated zoospores —_ produce anisogamous gametes (Fig. 3), which fuse to produce triflagellated planozygotes (two longitudinal flagella and one transverse) (Fig. 4). Sexual and asexual reproduction can occur on 54 Harmful Marine Dinoflagellates either a fish or algal diet (Tester, P., petsoliel communication). Species Comparisons: P. piscicida is a distinct free-living estuarine dinoflagellate (Fensome 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, microfauna, finfish 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, biflagellated 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 dinoflagellate species, a large food vacuole allows P. piscicida to phagocytize large prey items (Gaines & Elbrachter 1987; Schnepf & Elbrachter 1992; Burkholder et al. 1998). Pfiesteria piscicida is a strong ichthyotoxic dinoflagellate 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; Noga et al. 1996; Steidinger et al. 1996). P. piscicida and possibly other Pfiesteria-like 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 et al. 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 Pfiesteria and Pfiesteria-like species in several states, including Maryland and North Carolina (P. Tester, personal communication). Habitat and Locality: Pfiesteria piscicida was first identified from the Pamlico Sound in North Carolina. Since its emergence; however, P. piscicida and Pfiesteria-like 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 et al. 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; Burkholder & Glasgow 1997). Prorocentrum arenarium Faust, 1994 Plate 37, Figs. 1-6 Species Overview: Prorocentrum arenarium is an armoured, marine, sand-dwelling, benthic dinoflagellate species. This toxic species is associated with coral rubble and colored sand in tropical embayments 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 um 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 Dinoflagellates 55 vary from kidney-shaped to oblong (Figs. 1-5), with an average size of 0.62 um long and 0.36 um wide. Spacing between poroids is 2-3 um. 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 periflagellar 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 um) (Fig. 1)(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: Prorocentrum arenarium is a photosynthetic species with a prominent central pyrenoid and a_ posterior nucleus (Fig. 6). A small (2-3 um), narrow, tubular, peduncle-like structure in the periflagellar 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 jm)(Faust 1990b), but larger than P. ruetzlerianum (cell diameter 28-35 wm) (Faust 1990b) and P. compressum (cell diameter 36 m)(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 um (Faust 1991): P. maculosum has about 85-90 valve poroids and 65-75 marginal poroids with a diameter of 0.6 um (Faust 1993b). The architecture of the periflagellar area of P. arenarium, With no ornamentation (Faust 1994), is similar to that of P. concavum, P. ruetzierianum (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 PP. 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 balticum (Lohmann) Loeblich HI, 1970 Plate 38, Figs. 1-4 Species Overview: Prorocentrum balticum is an armoured, marine, planktonic, bloom-forming dinoflagellate 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 um 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. 1,3,4). 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 periflagellar pores are different sized (Fig. 3)(Dodge 1975; Toriumi 1980; Steidinger & Tangen 1996; Faust et al. 1999), Nomenclatural Types: Holotype: Exuviaella baltica Lohmann, 1908: 265, plate 17, fig. la,b Type Locality: unknown Synonyms: Prorocentrum pomoideum Bursa, 1959 Exuviaella aequatorialis Hasle, 1960 Morphology and Structure: = Prorocentrum balticum is a photosynthetic species with a round nucleus situated posteriorly (Dodge 1975; Dodge 1982; Toriumi 1980). Reproduction: P. ba/ticum 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's highest growth rates were at low salinities (10-15 0/00). Toxicity: Although toxicity in P. balticum 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 et al. 1999). Because of its small size, records of P. balticum 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 belizeanum Faust, 1993 Plate 39, Figs. 1-9 Species Overview: Prorocentrum belizeanum is an armoured, marine, benthic dinoflagellate 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 um in length and 50-55 um in width. Valves are concave in the center (Figs. 2,4) (Faust 1993a). 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 um 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 periflagellar 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 periflagellar 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 1993: figs. 1,2 Type Locality: Caribbean Sea: Twin Cays, Belize, Central America belizeanum — Faust, 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 binary fission. Ecology: P. belizeanum ts 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 (DTX1)(Morton et al. 1998). Species Comparison: Only a few round or near- round Prorocentrum species are known: P. belizeanum is larger then P. hoffmannianum (45- 55 um long and 40-45 um wide)(Faust 1990b) and larger than P. compressum (36 wm 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. hoffmannianum _ has approximately 670 areola per valve (diam.=1.0- 1.15 um), and P. ruetzlerianum has about 550 pentagonal-shaped areola per valve (diam.=1.0 um)(Faust 1990b). The architecture of the periflagellar area of P. belizeanum is similar to P. ima (Taylor 1980) and the planktonic species P. playfairi (Croome & Tyler 1987). P. hoffmannianum (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. Jima (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. hoffmannianum (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. hoffmannianum, 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 dinoflagellate. 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 um in length and 38-45 um in width. The valve surface is covered with 1000- 1100 prominent shallow areolae. The areolae are round to oval with smooth edges (Figs. 1,3) and often observed with a central pore (0.8 ym 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. 1,2)(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. 1)(Fukuyo 1981; Faust 1990b), Nomenclatural Types: Holotype: Prorocentrum 1981: figs. 13-19,49 Type Locality: Pacific Ocean: French Polynesia, New Caledonia and the Ryukyu Islands concavum Fukuyo, Morphology and Structure: Prorocentrum concavum 1S 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 (Tindall et al. 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. Jima (Fukuyo 1981; Faust 1990b), but P. dima 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). P. concavum 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). Prorocentrum faustiae Morton, 1998 Plate 41, Figs. 1-4 Species Overview: Prorocentrum faustiae is an armoured, marine, benthic dinoflagellate 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 um long and 38-42 um wide. Small pores (0.1 tm 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 periflagellar 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, 1998: 567, figs. 1-4 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 faustiae reproduces asexually by binary fission. Harmful Marine Dinoflagellates 59 Ecology: Prorocentrum faustiae is a benthic species epiphytic on macroalgae (Morton 1998). Toxicity: P. faustiae is a diarrhetic shellfish poison (DSP) toxin-producing species producing okadaic acid (OA) and_ Dinophysistoxin-1 (DTX1)(Morton 1998). Species Comparison: Prorocentrum faustiae is similar in shape and size to P. hoffmannianum (45-55 tum long and 40-45 um 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 of non-planktonic dinoflagellates (Morton 1998). Habitat 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, benthic dinoflagellate 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 um long and 40- 45 um wide. Both valves are slightly concave in the center. The intercalary band is smooth and appears as a flared 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 um 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. 1,2)(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. /ima, 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. hoffmannianum 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. reticulatum (Faust 1997), P. sabulosum (Faust 1994), P. belizeanum (Faust 1993a) and P. hoffmannianum (Faust 1990b) share a distinct feature in the periflagellar area: three small accessory pores adjacent to a periflagellar pore (Faust 1997). Both P. hoffmannianum and P. belizeanum have a prominent flared curved apical collar on the left valve bordering the periflagellar 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. hoffmannianum 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). Prorocentrum lima (Ehrenberg) Dodge, 1975 Plate 43, Figs. 1-7 Species Overview: Prorocentrum lima is an armoured, marine, benthic dinoflagellate species with world-wide distribution. Taxonomic Description: P. Jima 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 um in length and 20-28 um 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 um in diameter)(Figs. 1,3), and about 60-100 larger round to oblong unevenly distributed valve pores with trichocysts (0.48 tm 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. Jima 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. 1,2,6)(Faust 1991; Steidinger & Tangen 1996). Nomenclatural Types: Holotype: Prorocentrum lima (Ehrenberg) Dodge, 1975: 109, figs. 1E,F, plate 1B,C Type Locality: unknown Synonyms: Exuviaella marina Cienkowski, 1881 Exuviaella lima (Ehrenberg) Bitschli, 1885 Exuviaella marina var. lima (Ehrenberg) Schiller 1933 Basionym: Cryptomonas lima Ehrenberg, 1860 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. /ima 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 Dinoflagellates 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. /ima 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 (70- 75 um 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-| (DTX1)(Marr et al. 1992); Dinophysistoxin-2 (DTX2)(Hu et al. 1993); and Dinophysistoxin-4 (DTX4)(Hu et al. 1995). Species Comparison: P. Jima 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. hofjfmannianum). 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. hoffmannianum are also similar in shape to P. /ima, 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). Prorocentrum maculosum 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 um long and 30- 40 um 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 um), 2-4 «um apart (Fig. 3). Valve center is devoid of poroids (Figs. 1,2,6)(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. 1,2)(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 periflagellar collar (Fig. 4)(Faust 1993b), Nomenclatural 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. /ima (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). In P. lima the thecal surface is smooth with round pores; only the larger flagellar pore is surrounded by a curved periflagellar collar (Faust 1991). 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. hoffmannianum (Faust 1990b), P. compressum (Abe 1967; Dodge 1975), P. playfairi and P. foveolata (Croome & Tyler 1987). Etymology: The name ‘macu/osum’ 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 1s 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 um long and 20-25 um 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 Dinoflagellates 63 houses a prominent curved periflagellar collar adjacent to the auxiliary pore (Figs. 1,2,5). Opposite is a smaller periflagellar plate adjacent to the flagellar pore (Fig. 5). The large periflagellar collar (2 X 6 um) may appear as an apical spine, and has been reported as such (Fukuyo 1981; Carlson 1984). Both valves are excavated (Figs. 1,2)(Faust 1990b). Nomenclatural Types: Holotype: Prorocentrum mexicanum Tafall, 1942: plate 34, 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 periflagellar 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 periflagellar 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 dinoflagellate. This is a cosmopolitan species in cold temperate to tropical waters. Taxonomic Description: P. micans is a bivalvate species often observed in valve view. Cells of this species are highly variable in shape and size (Figs. 1-5)(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 um long) on the left valve (Figs. 1,3). Cells are medium-sized (35-70 um long, 20-50 wm 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. 1,4-6)(Wood 1954; Toriumi 1980; Dodge 1975; 1982; 1985; Fukuyo et al. 1990; Steidinger & Tangen 1996; Faust et al. 1999), The periflagellar area is a relatively small, shallow, broad triangular depression situated 64 Harmful Marine Dinoflagellates apically on the right valve off-center (Fig. 3). Two periflagellar pores are present: one large flagellar pore and one smaller auxiliary pore (Fig. 3). Adjacent to the flagellar pore is a small, slightly curved periflagellar plate (Fig. 3). The large pointed apical spine lies adjacent to the periflagellar area, directly opposite the periflagellar plate (Fig. 3)(Taylor 1980; Toriumi 1980). Nomenclatural Types: Holotype: Prorocentrum 1834: 307 Type Locality: North Sea: near Kiel, Berlin, Germany Synonyms: Cercaria sp. Michaelis, 1830 Prorocentrum schilleri Bohm in Schiller, 1933 Prorocentrum levantinoides Bursa, 1959 Prorocentrum pacificum Wood, 1963 micans Ehrenberg, Morphology 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 Prorocentrum. 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 0/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 & Berthome 1988). Species Comparison: This species varies considerably in shape and size and may be confused with closely related species; e.g. P. gracile, P. scutellum and P. caribbaeum. — 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 periflagellar 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 Prorocentrum 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 um long to 10-15 um 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 periflagellar 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 periflagellar 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 triangulatum 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 mininum is a photosynthetic species with golden-brown chloroplasts, one large pyrenoid and two pusules. The nucleus is broadly ellipsoidal and posteriorly situated (Parke & Ballantine 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). Cells 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 periflagellar 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 Harmful Marine Dinoflagellates Prorocentrum ruetzlerianum Faust, 1990 Plate 48, Figs. 1-6 Species Overview: Prorocentrum ruetzlerianum is an armoured, marine, benthic dinoflagellate species. This species is associated with floating detritus and sediment in tropical embayments of the Caribbean Sea. Taxonomic Description: P. ruetz/erianum 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 um. 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 um 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 periflagellar 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). Nomenclatural Types: Holotype: Prorocentrum ruetzlerianum Faust, 1990: figs. 21-23 Type Locality: Caribbean Sea: Belize, Central America Twin Cays, Morphology and = Structure: = Prorocentrum ruetzlerianum is a photosynthetic species with golden chloroplasts, a centrally located pyrenoid (Figs. 4,5), and a posterior nucleus (Faust 1990b). 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. hoffmannianum has approximately 670 round to oval areolae per valve (1.1 um = diameter)(Faust 1990b); P. belizeanum has about 853-1024 round to oval areolae per valve (0.73 um = diameter)(Faust 1993a); and P. sabulosum has about 391 round to oval areolae per valve (1.3) um 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 tamarense. 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. hoffmannianum)., 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 Gambierdiscus 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 (CQ, and carbonates) are utilized for growth, metabolism and reproduction. auxotrophy - Heterotrophic nutritional mode in which specific external organic compounds (usually the vitamins By, 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 Dinoflagellates 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). cingular 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. Prorocentrum). 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 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. DAPI (4',6-Diamidino-2-Phenylindole) - A highly specific and sensitive fluoresceing DNA stain used in epifluorescent microscopy to observe structures containing DNA. DAPI 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 cell 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 ventrally; one flagellum 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). inward Harmful Marine Dinoflagellates 69 dinophysoid - Group of dinokont type dinoflagellates. Members of this group are the only thecate dinoflagellates 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 et al. 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. epifluorescence microscopy - A method of microscopy used to view light-excited regions of an organism stained with a fluorochrome dye. epiphyte - 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 flagella emerge located in the periflagellar 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 unarmoured 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 (diploid-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 monilatum. 70 Harmful Marine Dinoflagellates lacerate - With a deeply and irregularly incised margin. lanceolate - Tapering at both ends. lateral - In desmokonts 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. megacytiec 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 (Steidinger & Tangen 1996). mesokaryotic - Dinoflagellate nucleus which possesses characteristics of both prokaryotes and eukaryotes. mixotrophy - Heterotrophic nutritional mode of some photosynthetic dinoflagellates in which ingestion of food particles is required for existence. mucocyst - An ejectile organelle; a minute structure that emerges through pores in the theca of armoured dinoflagellates 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 Shellfish Poisoning (NSP) - A human neurological disease caused by the ingestion of toxic marine shellfish (filter- 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 dinoflagellates 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 dinoflagellate 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, finger-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 (desmokonts) 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 periflagellar pores, the auxilliary pore (A) and_ the flagellar pore (F). Accessory pores, apical spine(s), periflagellar = collars and/or periflagellar 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 maculosum). A_ periflagellar collar can be flared and/or protuberant, or appear as a winged spine (e.g. Prorocentrum mexicanum). 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 epitheca (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 Dinoflagellates 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). transdiameter - 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 dinoflagellates that releases filamentous or fibrillar threads when discharged. tycoplanktonie - 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 (1') 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 1996). ventral ridge - In dinokonts, an identifiable ridge on the right side of the sulcal intrusion onto the epitheca. vermiculate - 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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Sexual reproduction in the dinoflagellate Noctiluca miliaris Suriray. J. Phycol. 6: 122-126. Zingone, A., M. Montresor & D. Marino 1998. Morphological variability of the potentially toxic dinoflagellate Dinophysis sacculus (Dinophyceae) and its taxonomic relationships with D. pavillardii and D. acuminata. Eur. J. Phycol. 33: 259-273. Zotter, J. 1979. Exuviaella baltica: a bloom organism of the Galveston Bay system. /n: D.L. Taylor & H.H. Seliger (eds.), Toxic Dinoflagellate Blooms, — Elsevier/North- Holland, New York: 195-198. Zubkoff, P.L., J.C. Munday, Jr., R.G. Rhodes & J.E. Warinner, [Il 1979. Mesoscale features of summer (1975-1977) dinoflagellate blooms in the York River, Virginia (Chesapeake Bay Estuary). /n: D.L. Taylor & H.H. Seliger (eds.), Toxic Dinoflagellate Blooms, Elsevier/North-Holland, New York: 279-286. Harmful Marine Dinoflagellates 97 (Prakash & Taylor 1966) (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 PLATE 1 | (Prakash & Taylor 1966) (redrawn from Balech 1995) 4 drawings. Fig. 3. Ventral view: 1° plate bears ventral pore (vp). Hypotheca 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 Ly Hajlegraeff et al. 1991) Alexandrium catenella, Fig. 1. SEM: ventral view. Two cell chain. Cells round; wider than long. Rounded apex and slightly concave antapex. Cingulum deep and lipped; sulcus deeply impressed and widens posteriorly. Hypotheca with prominent sulcal lists (arrows). Fig. 2. LM: four cell chain. Cells anterior- posteriorly compressed. Fig. 3. LM: apical view. First (redrawn from Balech 1995) apical plate (1') comes in direct contact with apical pore plate (Po). Ventral pore absent from 1” plate. Fig. 4. SEM: apical pore complex (APC). Foramen fishhook shaped; anterior attachment pore (aap) adjacent. Fig. 5. Line drawing: thecal plates depicted. Fig. 6. LM: resting cyst elliptical with rounded ends. Harmful Marine Dinoflagellates 79 PLATE 3 (Hallegraeff 1991) is = 1 (Hallegraeff 1991) APC P foramen F /\ ¢ ; yf) =| bY wy g 3 al A . 4 B 5 a & Fukuyo) (redrawn from Balech 1995) 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 Alexandrium minutum. 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 1X its width. Sulcus shallow (sa=anterior sulcal plate). Apical pore plate (Po) in direct contact with 1° plate. Fig. 2. LM: ventral 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 Dinoflagellates PLATE 4 (Walker & Steidinger 1979) (redrawn from Balech 1995) 4 Alexandrium monilatum. 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 I' plate. Anterior (redrawn from Balech 1995) (Walker & Steidinger 1979) 5 aap foramen (redrawn from Balech 1995) 3 (Walker & Steidinger 1979)" 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 Dinoflagellates 10] (Fukuyo etal.) A Fe Ka Vp > 4 (redrawn from Balech 1995) Alexandrium ostenfeldii. Figs. 1-3. LM. Fig. 1. 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 (1') forms a 90 degree angle at the point where vp and 4' plate come in contact. Apical pore complex (APC) with comma-shaped foramen. Figs. 3-4. Line drawings. (Larsen & Moestrup 1989) (Fukuyo et al.) PLATE 5 (redrawn from Balech 1995) 3 plate wider than high. Fig. 3. Ventral view: 6” Cingulum (C) slightly excavated. Fig. 4. APC and I’ plate: a. Po in direct contact with 1°; b. Po in indirect contact with 1° 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 (Fukuyo et al.) oy ge 1 (Montresor 1995) 2 (Fukuyo et al.) (Fukuyo et al.) . tay , oe (redrawn from Balech 1995) ¢ ' oq ' A (redrawn from Balech 1995) 5 6 (Montresor 1995) Alexandrium pseudogonyaulax. Figs. 1-4. LM. Fig. 1. Ventral view. Cell broadly pentagonal; wider than plate margin. Fig. 4. Epitheca: apical view. 1’ plate does not come in contact with Po. Po oval and long. Epitheca short and dome-shaped. Hypotheca longer than epitheca. Cingulum shallow and barely displaced. Fig. 2. Dorsal view. Antapex obliquely concave. Fig. 3. Epitheca: ventral view. Apical pore plate (Po) with comma-shaped foramen. I’ plate pentagonal with large wide ventral pore (vp) on 4' longitudinal on apex. Figs. 5-6. Line 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 (Fukuyo et al.) (L664 ‘Ie 18 yeesbayeH) Alexandrium tamarense. Fig. 1. LM. Two cell chain: cells small to medium; slightly longer than wide, nearly spherical. Cingulum (C) deeply escavated and lipped. Left hypothcal lobe slightly larger than right. Nucleus (n) visible. Figs. 2-4. SEM. Fig. 2. Two cell chain: cingulum displaced 1X its width. Deep sulcus (s) widens posteriorly. Fig. 3. Epitheca: apical view. Apical pore plate (Po) rectangular; narrows (redrawn from Balech 1995) 5 PLATE 7 i(Hallegraeff et al, 1991) ventrally. Po and first apical plate (1°) 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 (Hallegraeff 1991). Fig. 5. Line drawing. Fig. 6. LM. Oblong resting cyst with rounded ends, reddish lipid bodies; covered in mucilage. 104 Harmful Marine Dinoflagellates PLATE 8 (je 8 OANyN 4) \ (redrawn from Balech 1995) s) Alexandrium tamivavanichi. Figs. 1-3. LM. Fig. 1. Two cell chain: cells medium-sized; round to slightly wider than long. Epitheca with shoulders. Fig. 2. Cells stained with calcofluor white: cingulum displaced 1X its width; sulcus widens posteriorly. Fig. 3. Apical view: apical pore plate (Po) houses comma-shaped foramen. First apical plate (1') with ventral pore (vp). Figs. 4-5. Line drawings. Fig. 4. 1' a A (redrawn from Balech 1995) 4 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 of s.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 (Fukuyo et al.) (Fukuyo et al.) (‘12 3 OANyN4) (‘je 3 OANyNY) 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 PLATE 9 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 flexible lists (arrowheads). Ventral pore present (arrow). Fig. 2. Dorsal view: apical pore plate (arrow), Po, located off-center on epitheca. Fig. 3. Antapical view: hypothecal plates. Fig. 4. Smooth edged thecal pores unevenly distributed. Fig. 5. Po about 12 um long, slightly S) (after Fukuyo 1981) EPITHECA 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 flagella (arrows). Fig. 8. Line drawing: thecal plate arrangement. Harmful Marine Dinoflagellates 107 PLATE 11 . 1998) 3) ‘Zingone et al. 1998 CINGULAR 6 ingone et al’ 1998) (Redrawn from Steidinger & Tangen 1996) Dinophysis acuminata. Figs. 1-5. SEM: lateral view. Fig. 1. Cell oval and rotund; thecal surface with shallow depressions and scattered pores. Left sulcal list (LSL) extends beyond midpoint of cell. Well- developed cingular lists: anterior cingular list (ACL); posterior cingular list (PCL). C=cingulum. 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. Megacytic 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. ld). Fig. 5. Large dorsal nucleus (N). Small, blunt projections on tapered antapex (arrow). Fig. 6. Line drawing. 108 Harmful Marine Dinoflagellates PLATE 12 (Dodge 1985) LEFT SULCAL LIST (Redrawn from Steidinger & Tangen 1996) 4 Dinophysis acuta, Fig. 1. SEM: lateral view. Cell oblong and robust: theca heavily areolated. Well developed cingular lists (CL) and left sulcal list (LSL). Pointed antapex. Figs. 2-3. LM: lateral view (from Larsen & Moestrup 1992: figs. 2a,d: scale bars=20 um). Fig. 2. Large areolae, each with a pore (arrows). Fig. 3. Widest point below mid-section (dashed line) aligned with third sulcal rib (arrow). Fig. 4. Line drawing. Harmful Marine Dinoflagellates 109 Dinophysis caudata. Figs. 1-2. 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. PLATE 13 POSTERIOR PROJECTION 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. 110 Harmful Marine Dinoflagellates PLATE 14 LSL (Larsen & Moestrup 1992) 2 The (Larsen & Moestrup 1992) 3 (redrawn from Steidinger & Tangen 1996) Dinophysis fortii. Fig. 1. SEM: lateral view. Left Figs. 2-3. LM: lateral view. Fig. 2. Cell subovate with sulcul list (LSL) long and well-developed. Right sulcal a wide round posterior bottom (dorsal bulge)(arrows). list (RSL) present. Cingulum (C) obscures low and Fig. 3. LSL supported by three strong ribs (arrows). small epitheca. Thecal surface covered with areolae. Smoothly convex dorsal margin. Fig. 4. Line drawing. Harmful Marine Dinoflagellates 11] (redrawn from Steidinger & Tangen 1996) Dinophysis mitra. Figs. 1-4. SEM. Fig. 1. Lateral view: cell broad and wedge-shaped; epitheca visible. Left sulcal list (LSL) short (arrow). Right sulcal list (RSL) small (arrowhead). Theca heavily areolated. Fig. 2. Epitheca cap-like: greatly reduced. LSL supported by three short ribs (arrows). Ventral hypothecal margin concave below LSL (arrowheads). PLATE 15 ) (Fukuyo et al. Fig. 3. Dorsal view: hypothecal margin smoothly convex. Short anterior cingular list (ACL) and posterior cingular list (PCL) supported by 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 & “al 4 (Larsen & Moestrup 1992) Dinophysis norvegica. Fig. 1. SEM: lateral view. Cell heavily areolated with pointed antapex and posterior protrusions (arrowheads). 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. 1; pointed antapex. Fig. 3. Robust cell with rounded antapex. Moestrup 1992) 2 smn eats - aa — (Larsen & Moestrup 1992) S (Larsen & Moestrup 1992) 3° (redrawn from Steidinger & Tangen 1996) 6 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. Fig. 5. Large posterior nucleus (n). Pointed antapex with posterior projections (arrows). Fig. 6. Line drawing. Right sulcal list depicted (RSL). Harmful Marine Dinoflagellates 113 rr Nae eau less 77744 $,5)5) Sle a ae (een, 5 (redrawn from Steidinger & Tangen 1996) Dinophysis rotundata. Figs. 1-2. SEM: lateral view. Fig. 1. Cell broadly rounded. Small cap-like epitheca (e) not obscured by cingular lists. Right suleal list (arrow). Fig. 2. Left sulcal list (LSL) (large arrow), over 1/2 the cell length, widens posteriorly. Surface pores present (small arrows). PLATE 17 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 Dinoflagellates PLATE 18 Dinophysis sacculus. Figs. 1-3. SEM: lateral view. Fig. 1. Cell oblong with rounded _ posterior. Hypotheca long, margins undulate. Thecal surface coarsely areolated. 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. " , * +5 4 oO ; Bro P50 O00 0°2900 Ooo o' o (after Stein 1883) Smooth thecal surface with pores. Metacytic zone (M) devoid of pores. Figs. 4-5. LM: lateral view. Fig. 4. Hypotheca 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 Dinoflagellates 115 i ® o © = Qo = Oo 1] 2] —_ s Cc i?) —_— oO Iongitudinal ridges SEM. Fig. 1. Ventral view: cell large, elongate and quadrilateral. Gonyaulax polygramma. Figs. 1-3. 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 PLATE 21 TI |} x tc} we] MD) or) | ak, <@) fe), OO} (O66L 1 38 OAnyN4) VS (redrawn from Steidinger & Tangen 1996) 6 ol (LF) flagella 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 Gymnodinium breve. Fig. 1. SEM: ventral view. Cell small, wider than long, dorso-ventrally flattened. Cell nearly square in outline; prominent apical process (AP) 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. (redrawn from Steidinger & Joyce 1973) 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 YUKI & YOSHIMATSU 1987) (YUKI & YOSHIMATSU 1987):/e% (YUKI & YOSHIMATSU 1987) Gymnodinium catenatum. Figs. 1-3. SEM: ventral 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 (LF) and transverse (TF) flagella visible. Fig. 3. Chain cells (redrawn from Stesdinger & Tangen 1996) PLATE 23 €, AG om (Blackburn et al. 1989) 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 Dinoflagellates PLATE 24 Gymnodinium mikimotoi. Figs. 1-4. SEM. Fig. 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 by 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 (arrow)(after Fukuyo et al.). Fig. 4. Cell compressed dorso-ventrally (after Fukuyo et al.). Figs. 5-7. LM. Fig. 5. Cingulum displaced 2 times its width (arrows)(from Larsen & Moestrup 1989: fig. 16g). Fig. 6. Large nucleus (N) in left lobe of hypotheca. Fig. 7. Vegetative division. Division plane oblique. Harmful Marine Dinoflagellates 121 Gymnodinium pulchellum. Figs. 1-2. SEM: ventral view. Fig. 1. Cell — aall 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- developed apical groove: reverse S-shape. Transverse flagellum (TF) housed in cingulum. Sulcus slightly PLATE 25 (Larsen 1994) Mee), AG (after Larsen 1994) 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 122 Harmful Marine Dinoflagellates PLATE 26 (0661 ‘Ie 38 OAN\N4) (after Dodge 1982) 4 Gymnodinium sanguineum. Figs. 1-3. LM. Cell large, view. Deep cingulum median, displaced 1-2 times its pentagonal, and slightly dorso-ventrally flattened. width. Sulcus deeply notches hypotheca. Apical Cells vary in shape and size. Fig. 1. Ventral view. groove present (arrow). Fig. 3. Cell deeply pigmented; Epitheca and hypotheca nearly equal in size: epitheca central nucleus (n). Fig. 4. Line drawing. Spindle- conical, hypotheca bilobed (arrows). Fig. 2. Ventral shaped chloroplasts radially arranged. Harmful Marine Dinoflagellates 123 (9g6| aujueljeg Jaye) Gymnodinium 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 PLATE 27 (after Dodge 1982) (961 eunueyeg Jaye) 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 (Steidinger) | (Steidinger) (redrawn from Steidinger & Tangen 1996) AG (after Taylor et al. 1995) 3 4 Gyrodinium galatheanum. Figs. 1-2. SEM: ventral Cingulum wide, houses transverse flagellum (single view. Fig. 1. Cell small, oval to round, with distinct arrow). Longitudinal flagella present (double arrow). apical groove (AG). Cingulum (C) displaced 3 times Fig. 3. LM: ventral view. Cingulum deeply excavated its width. Short and narrow sulcus (S) slightly invades (arrows). Nucleus (N) large and central. Fig. 4. Line epitheca. Fig. 2. Epitheca and hypotheca round. drawing. Harmful Marine Dinoflagellates 125 Dodge 1985) "™ (Lewis & Burton 1988) Lingulodinium polyedrum. 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 (1') long and narrow. Apical pore plate PLATE 29 Dodge 1985) (redrawn from Steidinger & Tangen 1996) 4 (Lewis & Burton 1988) (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 Dinoflagellates PLATE 30 a (Bg61 dnyssoy 9 uesie)) (O661 1e 38 OANyN4) Noctiluca scintillans. Figs. 1-3. LM. Fig. 1. Cells large, balloon-shaped, nearly spherical, and colorless. A single flagellum housed in the ventral groove (arrow). Fig. 2. Cytoplasmic strands extend from nucleus (near the groove) to cell perifery. Posterior TE (redrawn from Steidinger & Tangen 1996) 4 Engulfed cell (arrowheads). Fig. 3. Asexually dividing cell. Fig. 4. Line drawing. Deep and wide ventral groove (VG) houses the tooth (TO), an extension of the cell wall. Striated tentacle (TE). Harmful Marine Dinoflagellates 127 Ostreopsis heptagona. Figs. 1-4. SEM. Fig. 1. Epithecal view: cells broadly oval, oblong and pointed. Long curved apical pore plate, Po, off-center (arrow). Plate 1' heptagonal and distinctive. Fig. 2. Hypothecal view: plate Ip pentagonal and dorso- ventrally elongate. Fig. 3. Po long, narrow and curved. (after Norris et al. 1985) PLATE 31 EPITHECA a" X Narrow mucilage strands cover cell surface. Fig. 4. Ventral view: location of ventral opening (arrow), ventral plate (asterisk), and rigid plate (asterisk) within cingulum. Fig. 5. LM. Two cells. Fig. 6. Line drawing: thecal plate arrangement. 128 Harmful Marine Dinoflagellates PLATE 32 8 (after Fukuyo 1981) Ostreopsis lenticularis, Figs. 1-5. SEM. Fig. 1. Shallow cingulum with smooth edge. Small sulcus Epithecal view: cell lenticulate to broadly oval. hidden (arrow). Fig. 5. Location of ventral opening Curved off-center apical pore plate with a slit-like (arrow), ventral plate (asterisk), and rigid plate apical pore (arrow). Plate I’ irregularly pentagonal. (arrowheads) within cingulum. Fig. 6. Line drawing: Fig. 2. Hypothecal view: plate Ip central and thecal plate arrangement. Figs. 7,8. LM. Fig. 7. pentagonal. Fig. 3. Smooth thecal surface. Round Cytoplasma granulated; posterior nucleus (n). Fig. 8. pores with smooth raised edges. Fig. 4. Hypothecal Distinct cingular list. ventral view: cell anterio-posteriorly compressed. Harmful Marine Dinoflagellates 129 Ostreopsis mascarenensis. Figs. 1-5. SEM. Fig. 1. Epitheca: inner thecal surface. Cell very large, broadly ovate, large plates. Plate 1' elongate and hexagonal. Apical pore plate (Po) nearly straight. Fig. 2. Hypotheca: plate Ip long and wide. Fig. 3. Smooth cell surface with round pores: pores with two small openings (arrows). Fig. 4. Po with long narrow apical pore; small pores line the opening (arrowheads). Figs. PLATE 33 5-6. Ventral view of epitheca. Fig. 5. Cell compressed anterio-posteriorly; cingulum narrow with smooth edge. Small sulcus hidden (arrow). Fig. 6. Location of ventral opening (large arrow), ventral plate (asterisk), and rigid plate (arrowheads) within cingulum, Pores with ejected trichocysts (small arrows). Fig. 7. LM. Epitheca: Po (arrow) and cingulum in focus. Fig. 8. Line drawing: hypotheca plate arrangement. 130 Harmful Marine Dinoflagellates PLATE 34 Ostreopsis ovata. Figs. 1-5. SEM. Fig. 1. Epithecal view: cell slender and tear-shaped. Apical pore plate (Po) off-center (arrow). Plate 1” large and hexagonal. Cingulum wide with narrow lists. Fig. 2. Hypothecal view: plates delicate. Plate Ip long and narrow. Fig. 3. Po: short and straight, adjacent to plate 2'. Fig. 4. G\ HYPOTHECA on (after Fukuyo 1981) Thecal surface smooth with scattered small pores. Suture line uneven and bumpy (arrows). Fig. 5. Hypothecal view: ventral opening (arrow), ventral plate (asterisk), and rigid plate (arrowhead) on cingulum. Fig. 6. LM. Large posterior nucleus. Fig. 7. Line drawing: thecal plate arrangement. Harmful Marine Dinoflagellates 131 v Ostreopsis siamensis. Figs. 1-6, SEM. Fig. 1. Epithecal view: cell broad and tear-shaped. Thecal surface smooth with scattered pores. Apical pore plate (Po) off-center (arrow). Narrow cingulum with smooth edge. Plate 1’ narrow and pentagonal. Fig. 2. Hypothecal view: plate 1p long and pentagonal. Fig. 3. Po: long, curved and narrow. Fig. 4. Large and small PLATE 35 (after Fukuyo 1981) 8 pores on thecal surface. Fig. 5. Ventral view: location of ventral opening (arrow), ventral plate (asterisk), and rigid plate (arrowhead) on cingulum. Fig. 6. Hypothecal view: Vo (arrow) and Rp (arrowhead). Fig. 7. LM. Hypotheca. Fig. 8. Line drawing: thecal plate arrangement. 132 Harmful Marine Dinoflagellates PLATE 36 (Steidinger et al/1996) eX (Steidinger et al. 1996) Steidinger et al, 1996) —— (Steidinger et al. 1996) Pfiesteria piscicida: Figs. 1-4. SEM. Figs, 1-2. Biflagellated stage resembles a gymnodinioid cell. Cells small, oblong and thecated. Fig. 1. Plate sutures apparent (arrows). Both flagella present (arrowheads). Fig. 2. Peduncle deployed (arrow). Fig. 3. Biflagellated stage with 2 size groups: large vegetative (V) cell; small gamete (G) cell. Fig. 4. Flagellated stage. Planozygote: larger triflagellated stage; similar to vegetative cell with 2 longitudinal flagella (arrows) adjacent to peduncle (P). Fig. 5. LM. Triple layer cyst (after Steidinger et al. 1996) F ot al. 3986) (after Steidinger et al. 1996) (arrows): benthic stage. Nucleus (N) stained with DAPI (courtesy of P. Tester). Figs. 6-7. SEM: biflagellate stage. Fig. 6. Epithecal plate morphology: thecal nodules apparent. Small la plate triangular (arrowhead). Plate 1” rhomboid (after Steidinger et al. 1996). Fig. 7. Apical view of APC: Po (arrowhead), cp. X plate. Figs. 8-9. Line drawings: plate tabulation. Fig. 8. Apical view: epitheca. Fig. 9. Ventral view: thecal nodules depicted. Harmful Marine Dinoflagellates 133 PLATE 37 Prorocentrum arenarium. Figs. 1-5. SEM. Fig. 1. Right valve: cells round to ovoid. Periflagellar area is a broad, V-shaped depression. Short longitudinal flagellum visible (arrowhead). Marginal poroids present (arrows). Fig. 2. Left valve: surface smooth, with scattered valve and marginal poroids (arrows). Fig. 3. Lateral view: intercalary band smooth; marginal poroids evenly spaced (arrowheads). Fig.4. Marginal poroids oblong to kidney-shaped. Fig. 5. Periflagellar area: triangular and unornamented with large flagellar pore (f) and smaller auxiliary pore (a). Fig. 6. LM. Right valve: posterior nucleus (n) and prominent central pyrenoid (arrow). 134 Harmful Marine Dinoflagellates PLATE 38 (Faust et al. 1999) Toriumi 1980) (redrawn from Steidinger & Tangen 1996) 4 Prorocentrum balticum. Figs. 1-3. SEM. Fig. 1. Valve scattered rimmed pores (arrows). Fig. 3. Periflagellar view: cell round to spherical, covered with many tiny region: two different sized pores and two small apical spines. Apical spine apparent. Intercalary band broad, projections (arrows). Fig. 4. Line drawing. transversely striated (arrows). Fig. 2. Surface with Harmful Marine Dinoflagellates 135 Prorocentrum belizeanum. Figs. 1-6, SEM. Fig. 1. Right valve: cell round to oval; surface heavily areolated. Fig. 2. Left valve: anterior margin with flared curved apical collar. Marginal areolae visible. Fig. 3. Lateral view: valve center concave; intercalary band smooth and wide. Fig. 4. Apical view: apical area with rounded lip; both valves excavated. Fig. 5. Areolae round to ovoid with smooth margins; some PLATE 39 with pores. Fig. 6. Periflagellar area: auxiliary pore (a) surrounded by curved periflagellar collar (arrows); adjacent to flagellar pore (f). Left valve with flared apical collar (arrowheads). Fig. 7. Left valve: central pyrenoid (arrow) and posterior nucleus (n). Fig. 8. LM: right valve: flagella present. Fig. 9. Line drawing: areolae arrangement (after Faust 1993a). 136 Harmful Marine Dinoflagellates PLATE 40 Prorocentrum concavum. Figs. 1-4. SEM. Fig. 1. Right valve. Cell ovate and heavily areolate. Valve center devoid of areolae. Left valve with anterior apical ridge (arrowhead). Fig. 2. Lateral view. Valve center concave and flattened. Intercalary band granulated and horizontally striated. Fig. 3. Valve areolae round to oval with smooth edges: some with eee *. e * e,%¢ ° fgeete sete “es oe? small central pores. Fig. 4. Periflagellar area a V- shaped depression. Two pores: small auxiliary pore (a); large flagellar pore (f). Figs. 5-6. LM (M.A. Faust). Fig. 5. Right valve. Fig. 6. Left valve. Fig. 7. Line drawing: areolae arrangement. (Figs. 1-4,7 atter Faust 1990b) Harmful Marine Dinoflagellates 137 PLATE 41 Prorocentrum faustiae. Figs. 1-4. SEM. Fig. 1. Right valve. Cells broadly ovate to rotundate with slightly concave center. Valve surface rugose. Periflagellar area situated apically. Fig. 2. Left valve: apical region slightly excavated. Fig. 3. Intercalary band wide and transversely striated. Small marginal pores evenly spaced along cell perifery (arrows). Fig. 4. Periflagellar area: apical view. Broad V-shaped depression; larger flagellar pore (f) adjacent to smaller auxiliary pore (a). (All figures donated by S.L. Morton) 138 Harmful Marine Dinoflagellates PLATE 42 iS) Prorocentrum hoffmannianum. Figs. 1-4. SEM. Fig. 1. Right valve: cell ovoid, tapering slightly apically. Valve surface areolated, slightly concave. Curved apical collar (arrow). Fig. 2. Left valve: distinct flared apical collar bordering periflagellar area (arrowheads). Marginal areolae large. Intercalary band smooth. Fig. 3. Areolae round to ovoid with smooth margins. Some with small pores (arrows). Fig. 4. Periflagellar area: ee | r e Oe fe fie hee **, e Fee Stan ea 8 a eee ieee eobos 4%, 4° Stoee e., Lr ) te "eevee ats Se wee sett, ee had ESP tot e Py re . vefec rs : spiae° He * oO flagellar pore (f) surrounded by flared periflagellar collar (arrowheads), adjacent to auxiliary pore (a); pores equal in size. Fig. 5. LM. Left valve: central pyrenoid (arrow); posterior nucleus (n). Intercalary band appears striated (M.A. Faust). Fig. 6. Line drawing: areolae arrangement. (Figs. 1-4.6 after Faust 1990b) Harmful Marine Dinoflagellates 139 P.LIMA 1¥ Prorocentrum lima. Figs. 1-3. SEM. Fig. 1. Right valve. Cells oblong to ovate with narrowed anterior. Marginal pores and scattered surface pores present: valve center devoid of pores. Intercalary band smooth and wide. Fig. 2. Left valve; bacteria attached (arrows). Fig. 3. Periflagellar area: shallow. broad, V- shaped depression on right valve. Flared periflagellar PLATE 43 ta sé 9 % 5 ue Mey > é ‘ ‘ collar encircles auxiliary (a) pore (arrow): larger flagellar pore (f) adjacent (after Faust 1991). Figs. 4-7. LM. Fig. 4. Thecal pore arrangement. Fig. 5. Right valve with central pyrenoid (arrow). Fig. 6. Left valve and posterior nucleus (n). Fig. 7. Triple-layered resting cyst. (Figs. 1,2,4-7 after Faust 1993c) 140 Harmful Marine Dinoflagellates PLATE 44 Prorocentrum maculosum, Figs. 1-4. SEM. Fig. 1. Right valve: cell broadly ovate, narrowing apically. Valve surface rugose with scattered poroids; valve center devoid of poroids. Marginal pores evenly spaced (arrows). Fig. 2. Left valve: anterior end flat to slightly concave with raised apical ridge (arrows). Valve margins appear as a flange around cell. Fig. 3. Valve poroids: unevenly distributed on valve surface; circular to oblong or kidney-shaped. Fig. 4. « ° « bad e ° e o% . e a e ee bd e e s - « Sf «6 * as * e a ? e ° e 7 a *.* . ° e « *~« « e e e e¢ * e? . . e a a} eo. . e . e « « . . ° 3 « s ee * bd "“,* ce? *“ 26% «© * * a e ° s « e e e« @e Ped . e e "ee Pad «oye e ty Periflagellar area: broad V-shaped depression on right valve. Apical ridge (raised margin) on left valve. Flagellar (f) and auxiliary (a) pores surrounded by protuberant periflagellar collar (arrowheads); equal in size. Fig. 5. LM. Right valve: central pyrenoid (P) and large posterior nucleus (N) (M.A. Faust). Fig. 6. Line drawing: valve poroid and marginal pore arrangement (Figs. 1-4,6 after Faust 1993b) Harmful Marine Dinoflagellates 141 Prorocentrum mexicanum. Figs. 1-5. SEM. Fig. 1. Right valve: cell oval. Periflagellar collar curved and prominent (arrow). Trichocyst pores radially arranged (arrowheads). Fig. 2. Left valve. Apical area excavated (M.A. Faust). Fig. 3. Lateral view: cell ovate to convex; intercalary band broad and transversely striated. Cell surface rugose. Fig. 4. Trichocyst pores round with smooth edge, within deep PLATE 45 furrowed depressions. Fig. 5. Periflagellar area: small, V-shaped shallow depression. Prominent curved periflagellar collar (double arrows) adjacent to auxiliary pore: protuberant periflagellar plate (single arrow) opposite and adjacent to flagellar pore. Fig. 6. LM. Right valve: radial pore arrangement visible (M.A. Faust). Fig. 7. Line drawing: trichocyst pore arrangement. (Figs. 1,3-5,7 after Faust 1990b) 142 Harmful Marine Dinoflagellates PLATE 46 ( Toriumi 1980) t Prorocentrum micans. Figs. 1-3. SEM. Fig. 1. Right valve: cell tear-drop shaped; rounded anteriorly, pointed posteriorly, broadest in the middle. Apical spine (AS) winged, Rugose thecal surface. Intercalary band smooth and wide. Fig. 2. Heart-shaped cell. Apical spine missing. Fig. 3. Periflagellar area: small, (after Dodge 1982) shallow triangular depression on right valve. Flagellar (f) and auxiliary (a) pores present; curved periflagellar plate adjacent to f. Large winged AS directly opposite. Figs. 4-5. LM: Left valve. Winged AS visible. Fig. 5. Empty theca with visible trichocyst pores (arrows). Fig. 6 Line drawing: trichocyst pore arrangement. Harmful Marine Dinoflagellates 143 PLATE 47 "ne thee 4 4 ji re : SS : “a ' Sa i 5 ee: | eT broad-based “ spines “‘} i (redrawn from Steidinger & Tangen 1996) Prorocentrum minimum. Figs. 1-4. SEM. Fig. 1. Right (arrow) adjacent to a. Intercalary band wide: valve. Cell oval; broad truncate apical region. Thecal surface with numerous short broad spines. Small scattered pores (arrows). Fig. 2. Lateral apical view. Periflagellar area with 2 pores: large flagellar (f) and small auxiliary (a). Small apical spine (arrowhead) adjacent to f; small curved forked periflagellar collar transversely striated. Fig. 3. Cells oval: ventrally flattened. Fig. 4. Apical view. Short thecal spines and small scattered pores (arrows). Figs. 5-6. LM. Surface features and intercalary band visible. Fig. 7. Line drawing. (Figs. 1-6 after Faust 1974) 144 Harmful Marine Dinoflagellates PLATE 48 Prorocentrum ruetzlerianum. Figs. 1-3. SEM. Fig. I. Right valve: cell round to ovoid, covered with pentagonal areolae. Cell surface rugose. Fig. 2. Anterio-lateral view. Each areola with small circular pore at its base (arrows). Intercalary band broad, transversely rugose, Fig. 3. Periflagellar area: small, shallow, unornamented depression on right valve; large flagellar (f) pore and smaller auxiliary (a) pore. Figs. 4-5. LM (M.A. Faust). Right valve: striated valve margins (small arrows); large central pyrenoid (large arrow). Fig. 6. Line drawing: areolae arrangement. (Figs. 1-3,6 after Faust 1990b)