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 | 54Mona Hoppenrath | Shauna A. Murray | Nicolas Chomérat | Takeo Horiguchi Marine benthic dinoflagellates – unveiling their worldwide biodiversity2014 | 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) BalechPublication: 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: F igs 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 habitat note

Hypnodinium Klebs 1912

H. sphaericumKlebs

F

Manchudinium Skvortzov 1972

M. sinicum(Skvortzov) Skvortzov

F

Phytodinedria Pascher 1944

P. aerugineaPascher

F taxonomic junior synonymof Cystodinedria sensuPopovský and Pfiester 1990

Phytodinium Klebs 1912

P. simplex Klebs F

Pyramidodinium Horiguchi et Sukigara 2005

P. atrofuscum Horiguchi et Sukigara

M

RhizodiniumBaumeister in Bourrelly 1955

R. gessneri (Baumeister)Loeblich et Loeblich

F

SpiniferodiniumHoriguchi et Chihara 1987

S. galeiformeHoriguchi et Chihara

M

StylodiniumKlebs 1912

S. globosumLoeblich et Loeblich

F, M

TetradiniumKlebs 1912

T. javanicumKlebs

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.

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Kleine Senckenberg-Reihe | 54

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

Table of Contents

Greetings 4Foreword 5Contents 7Acknowledgements 10

I. Introduction 12

II. ‘Materials & Methods’ 16Habitats 16Sampling 16Extraction = separation from the substrate 19Fixation and Electron Microscopy (EM) 19Culturing 21Quantification 21

III. Taxonomy 22Adenoides 22Alexandrium 24Amphidiniella 25Amphidiniopsis 27Amphidinium 41Ankistrodinium 56Apicoporus 58Biecheleria 61Bispinodinium 62Bysmatrum 64Cabra 70Coolia 74Dinothrix 80Durinskia 82Galeidinium 85Gambierdiscus 86Glenodinium 95Gymnodinium 96Gyrodinium 103Halostylodinium 107

Herdmania 109Heterocapsa 111Katodinium 112Moestrupia 115Ostreopsis 116‘Peridinium’ partim = new genus 126Pileidinium 128Plagiodinium 129Planodinium 130Polykrikos 132Prorocentrum 134Pseudothecadinium 152Pyramidodinium 154Rhinodinium 155Roscoffia 156Sabulodinium 160Scrippsiella 163Sinophysis 165Spiniferodinium 173Stylodinium 175Symbiodinium spp. 177Testudodinium 178Thecadinium 180Togula 188Vulcanodinium 191

IV. Phylogeny and systematics 193Phylogeny of the morpho logical adaptations 194Amphidinium 195Amphidiniopsis, Archaeperidinium, Herdmania – Peridiniales 195Cabra, Rhinodinium, Roscoffia – Podolampadaceae 196Coolia, Gambierdiscus, Ostreopsis – Gonyaulacales 196Prorocentrum & Adenoides 197Sinophysis & Sabulodinium 197

‘Dinotoms’ – Dinothrix, Durinskia, Galeidinium, ‘Gymnodinium’ quadrilo- batum, ‘Peridinium’ quinquecorne 198Dinoflagellate taxa with cryptophyte-(klepto)chloroplasts 198The phytodinialean dinoflagellates(‘Phytodiniales’) 198

V. Biogeography 209

VI. Ecology 212Attachment 213Life cycles 213Tide pools 213Vertical migration 214Blooms 214Spatial distribution 216Temporal distribution 216Quantitative Data 217

VII. Toxins of benthic dinoflagellates and benthic harmful algal blooms 218Introduction 218Gambierdiscus 219Ostreopsis 223Coolia 223Prorocentrum 226Amphidinium 227Alexandrium 227Vulcanodinium 227

References 234Taxonomic index 266Useful web pages 272Picture credits 273Authors’ Addresses 274

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