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.
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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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