Marine Benthic Dinoflagellates

Dinoflagellates are important primary producers, symbionts and, at the same time, heterotrophic consumers and parasites. The species composition in benthic habitats is quite distinct from planktonic habitats. Our understanding of benthic dinoflagellate biodiversity, biogeography, toxicology and ecology has improved but is still rudimentary. Benthic harmful algal blooms have attracted increasing interest because of the impact of ciguatera poisoning, the most important food-borne disease of non-bacterial origin worldwide, which is caused by benthic dinoflagellate species. Ciguatera ­poisoning appears to have increased worldwide in recent years.

This publication is an updated summary of the taxonomy of currently described taxa and includes morphological and molecular genetic information for species identification. It contains the most comprehensive review of benthic dinoflagellate toxins published so far. The book also describes methods of study, discusses phylogenetics and evolution, and highlights their scientific relevance as well as the health and economic impacts of benthic dinoflagellates for society.

This book is a fundamental contribution to improving the monitoring of benthic dinoflagellates worldwide: 242 species in 63 genera are presented, illustrated with more than 240 colour images, about 250 electron micrographs and more than 330 drawings.

Greetings   4
Foreword   5
Contents   6

I. Introduction   9

II. Materials & Methods   14
Habitats   14
Sampling   14
Extraction = separation from the substrate   16
Light Microscopy (LM) and epifluorescence   17
Fixation and Electron Microscopy (EM)   18
Culturing   19
Quantification   19
Molelcular Genetic Methods of Detection and Quantification   20
Toxicity tests and toxin analyses   22

III. Taxonomy   23
Adenoides   23
Aduncodinium   26
Ailadinium   28
Alexandrium   30
Amphidiniella   32
Amphidiniopsis   34
Amphidinium   55
Ankistrodinium   70
Ansanella   72
Apicoporus   74
Bindiferia   76
Bispinodinium   78
Blixaea   80
Bysmatrum   83
Cabra   91
Caladoa   94
Carinadinium   96
Ceratocorys   100
Chiharadinium   101
Chrysodinium   104
Coolia   105
Coutea   116
Dinothrix   116
Durinskia   120
Fukuyoa   124
Gambierdiscus   127
Glenodinium   140
Gymnodinium   141
Gyrodinium   146
Halostylodinium   151
Herdmania   152
Heterocapsa   154
Katodinium   155
Laciniporus   157
Madanidinium   160
Moestrupia   160
Nusuttodinium   162
Ostreopsis   165
Pachena   165
Paragymnodinium   181
Pellucidodinium   183
Pentaplacodinium   184
Pileidinium   185
Plagiodinium   187
Planodinium   188
Polykrikos   190
Prorocentrum   192
Psammodinium   216
Pseudadenoides   218
Pseudothecadinium   220
Pyramidodinium   222
Rhinodinium   224
Roscoffia   228
Sabulodinium   229
Sinophysis   231
Speroidium   240
Spiniferodinium   241
Stylodinium   244
Symbiodiniales   246
Testudodinium   246
Thecadinium   250
Togula   255
Vulcanodinium   258

IV. Phylogeny and systematics   260
Phylogeny of the morphological adaptations   260
Amphidinium   261
Amphidiniopsis, Archaeperidinium, Herdmania - Peridiniales   262
Cabra, Rhinodinium, Roscoffia – Podolampadaceae   263
Gonyaulacales   263
Prorocentrum & Pseudadenoides/Adenoides   266
Sinophysis & Sabulodinium   267
‘Dinotoms’ – Blixaea, Dinothrix, Durinskia   267
Dinoflagellate taxa with cryptophyte-(klepto)chloroplasts - Nusuttodinium   268
The phytodinialean dinoflagellates (‘Phytodiniales’)   268

V. Toxins of benthic dinoflagellates and benthic harmful algal blooms   269
Introduction   269
Gambierdiscus & Fukuyoa   270
Ostreopsis   279
Coolia   282
Prorocentrum   286
Amphidinium   291
Alexandrium   292
Durinskia   292
Vulcanodinium   293

VI. Relevance for science and society   295
Important areas for basic research   295
1. General dinoflagellate evolution   295
2. Choloroplast evolution in (benthic) dinoflagellates   296
3. Nutritional supplements and natural products from benthic dinoflagellates   299
4. The genetics and evolution of dinoflagellate toxicity   300
Health, social and economic impacts   301
Ostreopsis and palytoxins   301
Gambierdiscus and ciguatoxins   304
Vulcanodinium and pinnatoxins   308
Prorocentrum and okadaic acid   309
Impacts of BHABs on marine biota   310
Conclusions   311

References   312
Taxonomic index   363
Acknowledgements   369
Useful web pages   371
Picture credits   372
Authors’ Addresses   373
Imprint   376