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Kleine Senckenberg-Reihe | 54

Marine benthic dinoflagellates – unveiling their worldwide biodiversity | Hoppenrath | Murray | Chomérat | Horiguchi

This book is the fi rst summary of our knowledge of benthic dinofl agellate species.

Dinofl agellates are important primary producers and symbionts, but, at the same time, also consumers and para- sites. Species compositions of benthic habitats are quite distinct from those of planktonic habitats. Less than 10%

of the approximately 2000 described extant dinofl agellate species appear to be benthic. They occur in different types of habitats (chapter II) and their morphology, their behavior, and some of their life cycles (chapter VI) seem to be well adapted to the benthic life- style. Information on their geographic distribution is still very limited and is compiled herein (chapter V).

The study of harmful benthic dino- fl agellates started in the late 1970s when it was suspected that a benthic species, later named Gambierdiscus toxicus, was responsible for ciguatera fi sh poisoning, a type of human poi- soning linked to the consumption of certain species of tropical reef fi sh. As the number of ciguatera fi sh poison- ing incidents increases, and the dis- tribution of toxin producing benthic taxa seems to be expanding, detailed understanding of the species diversity and the ability to accurately identify them is becoming increasingly impor- tant (chapter VII).

Dinofl agellate classifi cation is current- ly undergoing changes and far from being settled, as new species and genera are discovered and systematic entities are rearranged. Many benthic dinofl agellate genera have unusual morphologies and appear to be only remotely related to known planktonic taxa, so that molecular phylogenetic analyses frequently show little statistic - al support for any relationship (chap- ter IV). Benthic species display unique thecal plate arrangements compared to planktonic species, e.g. Adenoides, Amphidiniella, Cabra, Planodinium, Sabulodinium, Rhinodinium (chap- ter III). Therefore, no classifi cation on higher rank levels (e.g. family, order) was used throughout this book. Gen- era (and species within a genus) are presented in alphabetical order.

This book presents the fi rst compre- hensive identifi cation help for benthic dinofl agellates. At the same time it

Kleine Senckenberg-Reihe | 54

Mona Hoppenrath | Shauna A. Murray | Nicolas Chomérat | Takeo Horiguchi Marine benthic dinoflagellates – unveiling their worldwide biodiversity

2014 | 276 pp. | 93 fi gs | 8 tabs | 14.8 x 21 cm | paperback | 19.90 € | ISBN 978-3-510-61402-8 www.schweizerbart.de/9783510614028 | www.senckenberg.de

22 III. Taxonomy – Adenoides III. Taxonomy

Adenoides [Aden: gland; eidos: sight – neutral]

Adenoides Balech Publication: Balech, 1956, Revue Algologique

2, pp. 30–31, Figs 1–8.

Type species: A. eludens (Herdman) Balech.

Plate formula: APC 4’ 6c 4s 5’’’ 5p 1’’’’

or APC 4’ 6c 5s 5’’’ 3p 2’’’’.

Description: Thecate genus with laterally flattened cells with a minute, depressed and scarcely visible epitheca. Shallow cingulum without displacement almost at the anterior cell end. No precingular plate series.

Remarks: A taxonomic problem with the original description of the type species has been discussed in detail in Hoppen- rath et al. (2003, pp. 385, 389) who rein- vestigated and revised the description of A. eludens. Whether a second Adenoides species, described by Herdman (1922) as Amphidinium species and transferred to Adenoides by Dodge (1982; without own observations), really exists, is not clear.

Because of this uncertainty it has not been included herein.

Adenoides eludens (Herdman) Balech Publication: Balech, 1956, Revue Algologique

2, p. 30.

Basionym: Amphidinium eludens E.C. Herd- man; Herdman 1922, Proceedings and Transactions of the Liverpool Biological Society 36, pp. 22–23 (26), Figs 1, (2).

Illustrations: Figs 5, 6.

Size: 25–40 µm long, 22–28 µm deep.

Plate formula: APC 4’ 6c 4s 5’’’ 5p 1’’’’

or APC 4’ 6c 5s 5’’’ 3p 2’’’’.

Chloroplasts: Two lobed brown peridinin- chloroplasts.

Description: Round to oval, asymmetrical, laterally flattened cells with minute depressed and scarcely visible epitheca.

The hypotheca is longer dorsally than ventrally. Smooth thecal plates with pores. Shallow cingulum without dis- placement almost at the anterior cell end. Short and slightly depressed sulcus with one flagellar pore located in the anterior third of the cell. No precingular plate series. Two conspicuous large pores at the dorsal posterior end. Two striking pyrenoids visible as rings because of the starch sheaths. Nucleus in the lower dor- sal hyposome half.

Distribution: Sandy sediments. Port Erin, Isle of Man, UK (Herdman 1922); North Suther- land, Scotland, UK (Dodge 1989); North German Wadden Sea, Germany (Hoppen- rath 2000b, Hoppenrath et al. 2003); Nor- mandy, France (Paulmier 1992); Roscoff, Brittany, France (Balech 1956, Dodge and Lewis 1986); Elba, Italy (Hoppenrath un- publ. obs.); Arabian Gulf, Kuwait (Saburova et al. 2009, Al-Yamani and Saburova 2010);

Sea of Japan, Russia (Konovalova and Se-

III. Taxonomy – Adenoides 23 Fig. 5: Adenoides eludens. A–C: Different focal planes, note the ring-like starch sheath around the pyrenoid (arrow); p = pusule, n = nucleus. Scale bars: 10 µm.

Fig. 6: Adenoides eludens. A: Left lateral view. B, C: Right lateral view; note the thecal pores in C. D–H: Drawings of the plate pattern. D: Left lateral. E: Right lateral. F: Dorsal.

G: Ventral. H: Epitheca, cingulum and sulcus. Scale bars: 10 µm.

88 III. Taxonomy – Gambierdiscus

Fig. 38: Gambierdiscus spp. drawings of the epithecal plate patterns. Relative sizes to scale.

III. Taxonomy – Gambierdiscus 89 Fig. 39: Gambierdiscus spp. drawings of the hypothecal plate patterns. Relative sizes to scale.

68 III. Taxonomy – Bysmatrum

Fig. 26: Bysmatrum granulosum, SEM (photos courtesy of A. Couté and L. Ten-Hage).

A: Ventral view. B: Apical view, epitheca. C: Left lateral view. D. Oblique view of the hy- potheca. Scale bars: 10 µm.

IV. Phylogeny and systematics 203

genus type species habitatnote

Hypnodinium Klebs 1912 H. sphaericum

Klebs F

Manchudinium Skvortzov 1972M. sinicum

(Skvortzov) Skvortzov

F

Phytodinedria Pascher 1944 P. aeruginea

Pascher F taxonomic junior synonym

of Cystodinedria sensu Popovský and Pfiester 1990 Phytodinium

Klebs 1912 P. simplex Klebs F

Pyramidodinium Horiguchi et Sukigara 2005

P. atrofuscum Horiguchi et Sukigara

M

Rhizodinium Baumeister in Bourrelly 1955

R. gessneri (Baumeister) Loeblich et Loeblich

F

Spiniferodinium Horiguchi et Chihara 1987

S. galeiforme Horiguchi et ChiharaM

Stylodinium Klebs 1912

S. globosum Loeblich et LoeblichF, M Tetradinium Klebs 1912

T. javanicum Klebs

F

F = freshwater, M = marine, B = brackish Table 1: Dinoflagellates classified as ‘phytodinialean’ taxa in the past.

aims to lend support in order to im- prove monitoring efforts worldwide.

About 190 species in 45 genera are presented in detail, illustrated with more than 200 color images, approxi- mately 150 scanning electron micro- graphs, and more than 250 drawings.

www.schweizerbart.de/9783510614028 | www.senckenberg.de

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Kleine Senckenberg-Reihe | 54

Marine benthic dinoflagellates – unveiling their worldwide biodiversity | Hoppenrath | Murray | Chomérat | Horiguchi

Table of Contents

Greetings 4

Foreword 5

Contents 7

Acknowledgements 10

I. Introduction 12

II. ‘Materials & Methods’ 16

Habitats 16

Sampling 16

Extraction = separation from

the substrate 19

Fixation and Electron

Microscopy (EM) 19

Culturing 21

Quantification 21

III. Taxonomy 22

Adenoides 22

Alexandrium 24

Amphidiniella 25

Amphidiniopsis 27

Amphidinium 41

Ankistrodinium 56

Apicoporus 58

Biecheleria 61

Bispinodinium 62

Bysmatrum 64

Cabra 70

Coolia 74

Dinothrix 80

Durinskia 82

Galeidinium 85

Gambierdiscus 86

Glenodinium 95

Gymnodinium 96

Gyrodinium 103

Halostylodinium 107

Herdmania 109

Heterocapsa 111

Katodinium 112

Moestrupia 115

Ostreopsis 116

‘Peridinium’ partim = new genus 126

Pileidinium 128

Plagiodinium 129

Planodinium 130

Polykrikos 132

Prorocentrum 134

Pseudothecadinium 152

Pyramidodinium 154

Rhinodinium 155

Roscoffia 156

Sabulodinium 160

Scrippsiella 163

Sinophysis 165

Spiniferodinium 173

Stylodinium 175

Symbiodinium spp. 177

Testudodinium 178

Thecadinium 180

Togula 188

Vulcanodinium 191

IV. Phylogeny and systematics 193 Phylogeny of the morpho logical

adaptations 194

Amphidinium 195

Amphidiniopsis, Archaeperidinium,

Herdmania – Peridiniales 195 Cabra, Rhinodinium, Roscoffia –

Podolampadaceae 196

Coolia, Gambierdiscus, Ostreopsis –

Gonyaulacales 196

Prorocentrum & Adenoides 197 Sinophysis & Sabulodinium 197

‘Dinotoms’ – Dinothrix, Durinskia, Galeidinium, ‘Gymnodinium’ quadrilo- batum, ‘Peridinium’ quinquecorne 198 Dinoflagellate taxa with cryptophyte-

(klepto)chloroplasts 198

The phytodinialean dinoflagellates

(‘Phytodiniales’) 198

V. Biogeography 209

VI. Ecology 212

Attachment 213

Life cycles 213

Tide pools 213

Vertical migration 214

Blooms 214

Spatial distribution 216

Temporal distribution 216

Quantitative Data 217

VII. Toxins of benthic dinoflagellates and benthic harmful algal blooms 218

Introduction 218

Gambierdiscus 219

Ostreopsis 223

Coolia 223

Prorocentrum 226

Amphidinium 227

Alexandrium 227

Vulcanodinium 227

References 234

Taxonomic index 266

Useful web pages 272

Picture credits 273

Authors’ Addresses 274

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