Springer Geology

Sreepat Jain

Fundamentals
of Invertebrate
Palaeontology
Microfossils
Springer Geology

Series Editors
Yuri Litvin, Institute of Experimental Mineralogy, Moscow, Russia
Abigail Jiménez-Franco, Del. Magdalena Contreras, Mexico City, Estado de
México, Mexico
Soumyajit Mukherjee, Earth Sciences, IIT Bombay, Mumbai, Maharashtra,
India
Chalina Tatiana Olegovna, Inst. of Problems in Mechanics, Russian
Academy of Sciences, Moscow, Russia
The book series Springer Geology comprises a broad portfolio of scientific
books, aiming at researchers, students, and everyone interested in geology.
The series includes peer-reviewed monographs, edited volumes, textbooks,
and conference proceedings. It covers the entire research area of geology
including, but not limited to, economic geology, mineral resources, historical
geology, quantitative geology, structural geology, geomorphology, paleon-
tology, and sedimentology.

More information about this series at http://www.springer.com/series/10172

Sreepat Jain

Fundamentals
of Invertebrate
Palaeontology
Microfossils

123
Sreepat Jain
Department of Applied Geology
Adama Science and Technology
University
Adama, Ethiopia

ISSN 2197-9545 ISSN 2197-9553 (electronic)

Springer Geology
ISBN 978-81-322-3960-4 ISBN 978-81-322-3962-8 (eBook)
https://doi.org/10.1007/978-81-322-3962-8

© Springer Nature India Private Limited 2020

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or
part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way,
and transmission or information storage and retrieval, electronic adaptation, computer software,
or by similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are
exempt from the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in
this book are believed to be true and accurate at the date of publication. Neither the publisher nor
the authors or the editors give a warranty, expressed or implied, with respect to the material
contained herein or for any errors or omissions that may have been made. The publisher remains
neutral with regard to jurisdictional claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature

India Private Limited
The registered company address is: 7th Floor, Vijaya Building, 17 Barakhamba
Road, New Delhi 110 001, India
For my son Parth
And my wife Archna
Preface

The Fundamentals of Invertebrate Palaeontology (Microfossils) is a

follow-up of my earlier book, Fundamentals of Invertebrate Palaeontology
(Macrofossils), and like the latter, the present book does not deal with the
basic principles or classification (the latter is mentioned only in brief). This
book primarily deals in detailing up-to-date morphological characteristics
of the studied organism and hence is intended for undergraduate students or
for early researchers.
Through illustrations of over 3000 hand-drawn diagrams, this book is
divided into four sections with eleven chapters. The first section includes
organic-walled microfossils (Chitinozoans, Acritarchs, Pollen and Spores,
and Dinoflagellates), followed by the next section on phosphatic Conodonts.
The third section includes siliceous microfossils (Radiolarians and Diatoms),
and the last section is about calcareous microfossils (Ostracods, For-
aminifera, Calcareous nannofossils, and Calpionellids) (Fig. 1).
Each chapter starts with a very brief Introduction section emphasizing the
relevance of the studied group and includes a series of biostratigraphic tables
through time marking the stratigraphic position of index fossils. For this, a
computer program, TSCreator version 7.4 (www.timescalecreator.org), is
used that is based on the 2016 age model of Ogg et al. (2016).
The next section elaborates the general morphology of the studied group
through a series of detailed and well-labeled line diagrams (this is the main
emphasis of this book). Attempt is made to illustrate the morphology of the
group through line diagrams, but in one chapter (Foraminifera), actual
specimen photographs are used to better illustrate the finer nuances of the
group’s morphology. Additionally, for this section, throughout this book,
attempt is also made to follow a bulleted pattern to enumerate morphology.
However, due to the morphological complexity of the studied groups and to
maintain succinctness of the text, this bulleted approach could not be fol-
lowed in letter and spirit.
The general morphology section is followed by brief notes on classifi-
cation and geological history with up-to-date references at the end of the
chapter. At the end of this book, a list of all species illustrated is provided. In
this list, wherever possible, age and up-to-date species identification and
assignments have been taken due care (particularly so for the Calcareous
nannofossils species, where data from the Nannotaxa Web site proved
immensely useful).

vii
viii Preface

Fig. 1 The figure shows the layout the book - four sections with eleven chapters (the Pollen and Spores form a single
chapter). The first section includes organic-walled microfossils (Chitinozoans, Acritarchs, Pollen and Spores, and
Dinoflagellates), the next section deals with phosphatic Conodonts., third with siliceous microfossils (Radiolarians and
Diatoms), and the last section is about calcareous microfossils microfossils (Ostracods, Foraminifera, Calcareous
nannofossils, and Calpionellids)

This book would not have seen the light of the day without the support
and encouragement of my wife, Archna, and my son, Parth; this book is
rightfully dedicated to them.
I would also like to profusely thank Elsevier (Palaeoworld) for granting
permission to reproduce some of the figures in this book (specifics mentioned
in the respective figure captions), to Dr. Michael A. Kaminski (Saudi Arabia),
Editor-in-Chief, Micropaleontology, for not only granting permission but also
allowing photographs to be used for the Foraminifera chapter, to Dr. Jeremy
Young (UK) for granting permission to use figures for the Calcareous nan-
nofossils chapter, to Dr. Julio Rodriguez Lazaro for sending high-resolution
figures to use for the Ostracod chapter, and to Drs. Justyna Kowal-Kasprzyk
and Mohamed Benzaggagh for the Calpionellid chapter. I am also very
grateful to Artai A. Santos for providing insightful suggestions in improving
Preface ix

the Pollen and Spore chapter and to Dr. Mohamed Benzaggagh for the
Calpionellid chapter; all the above have also been duly acknowledged in the
figure captions. Last but not least, I thank Dr. Mikhail Rogov (Russia) for
literature help. I once again thank them all from the depth of my heart for
their time, help, and kindness.

Adama, Ethiopia Sreepat Jain

References

Ogg, J. G., Ogg, G., & Gradstein, F. M. (2016). A Concise Geologic Time Scale (pp. 240).
Amsterdam: Elsevier.
https://timescalecreator.org.
https://www.mikrotax.org/Nannotax3/.
Contents

1 Chitinozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2.1 Chamber Shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.2 Ornamentation and Surface Patterns . . . . . . . . . . . . . 9
2.3 Linear Linkage Arrangements. . . . . . . . . . . . . . . . . . 11
3 Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.1 Aboral Pole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.2 Aboral Scar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.3 Aboral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.4 Anti-apertural Pole . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.5 Apertural Plug . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.6 Apertural Pole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.7 Aperture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.8 Apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.9 Apical Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.10 Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.11 Axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.12 Basal Callus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.13 Basal Edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.14 Basal Margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.15 Basal Pore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.16 Basal Scar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.17 Base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.18 Callus (Basal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.19 Carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.20 Catenary Structure (Chain-Like Structure) . . . . . . . . 16
3.21 Central Cavity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.22 Chain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.23 Chamber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.24 Cluster . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.25 Coiled Chain (=Helicoidal Chain) . . . . . . . . . . . . . . 16
3.26 Collar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.27 Collarette (=Collar) . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.28 Copula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.29 Crests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.30 Diameter (=D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
xi
xii Contents

3.31 External Structures . . . . . . . . . . . . . . . . . . . . . . . . . . 16

3.32 Fenestrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
3.33 Flank . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.34 Flexure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.35 Glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.36 Inner Layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.37 Laciniated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.38 Linear Chain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.39 Linkage Structures . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.40 Lip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.41 Longitudinal Axis . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.42 Margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.43 Mesh-Like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.44 Mucron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.45 Neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3.46 Operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.47 Oral Scar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.48 Oral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.49 Ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.50 Outer Layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.51 Peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.52 Perforated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.53 Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.54 Prosome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.55 Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.56 Sculpture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.57 Septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3.58 Shoulder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
3.59 Sleeve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
3.60 Spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
3.61 Spiny Ornamentation . . . . . . . . . . . . . . . . . . . . . . . . 19
3.62 Vesicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
3.63 Wall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
4 Chitinozoan Affinity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
5 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
6 General History and Distribution . . . . . . . . . . . . . . . . . . . . . 24
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
2 Acritarchs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.1 Sphaeromorphs and Acanthomorphs . . . . . . . . . . . . . 29
2.2 Morphogroups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.3 Vesicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.4 Excystment Opening. . . . . . . . . . . . . . . . . . . . . . . . . 34
3 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
4 Geological History and Distribution . . . . . . . . . . . . . . . . . . . 34
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Contents xiii

3 Pollen and Spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
2 Spore Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
3 Pollen Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
3.1 Pollen Units . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
3.2 Polarity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3.3 Apertures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
3.4 Shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
3.5 Symmetry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
3.6 Size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
4 Sporoderm Stratification. . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
4.1 Exine Ornamentation . . . . . . . . . . . . . . . . . . . . . . . . 58
5 Earliest Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4 Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
2.1 Cyst Types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
2.2 Archeopyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
3 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
5 Conodonts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
2.1 Elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
2.2 Composite Elements . . . . . . . . . . . . . . . . . . . . . . . . . 96
2.3 Simple Elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
2.4 Element Notation and Apparatus . . . . . . . . . . . . . . . 107
3 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
4 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
6 Radiolarians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
2 Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
3 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
7 Marine Diatoms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
2 Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
3 Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
3.1 Gross Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . 134
3.2 Cell Wall Structure . . . . . . . . . . . . . . . . . . . . . . . . . . 134
xiv Contents

4 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

5 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
8 Ostracods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
2.1 The Carapace and Associated Structures . . . . . . . . . 143
2.2 Lamellae and Cardinals . . . . . . . . . . . . . . . . . . . . . . 145
2.3 Pore Canals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
2.4 Muscle Scars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
2.5 Carapace Orientation . . . . . . . . . . . . . . . . . . . . . . . . 148
2.6 Valve Closure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
2.7 Sculpture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
3 Sexual Dimorphism, Ontogeny, and Morphology
of the Carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
4 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
5 General History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
9 Benthic Foraminifera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
2 External Morphological Features . . . . . . . . . . . . . . . . . . . . . 171
2.1 Test Shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
2.2 Chamber Arrangement . . . . . . . . . . . . . . . . . . . . . . . 175
2.3 Chamber Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
2.4 Position of Primary Aperture . . . . . . . . . . . . . . . . . . 178
2.5 Aperture Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
2.6 Wall Structure and Test Composition . . . . . . . . . . . . 182
2.7 Sutures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
2.8 Ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
3 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
10 Calcareous Nannofossils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
2 Morphology of Coccoliths . . . . . . . . . . . . . . . . . . . . . . . . . . 194
2.1 Heterococcoliths . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
2.2 Holococcoliths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
3 Terminology for Heterococcoliths and Nannoliths . . . . . . . . 196
3.1 Coccolith Orientation . . . . . . . . . . . . . . . . . . . . . . . . 196
3.2 Central Area and Rim. . . . . . . . . . . . . . . . . . . . . . . . 202
3.3 Coccolith Outline and Axial Ratio . . . . . . . . . . . . . . 205
3.4 Ultrastructure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
3.5 Element Shapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
3.6 Element Modifications . . . . . . . . . . . . . . . . . . . . . . . 208
3.7 Types of Openings . . . . . . . . . . . . . . . . . . . . . . . . . . 210
3.8 Element Curvature . . . . . . . . . . . . . . . . . . . . . . . . . . 210
3.9 Parts of Rims . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Contents xv

3.10 Directions on the Rim . . . . . . . . . . . . . . . . . . . . . . . 211

3.11 Element Arrangement as Seen in Plan View . . . . . . 212
3.12 Coccolith Arrangement . . . . . . . . . . . . . . . . . . . . . . . 212
4 Terms Used in Describing the Parts of a Holococcolith. . . . 213
4.1 Holococcolith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.2 Crystallite Arrangement . . . . . . . . . . . . . . . . . . . . . . 213
4.3 Plug . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.4 Rim . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.5 Block . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.6 Cavity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.7 Pore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.8 Proximal Flange . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.9 Proximal Ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.10 Wall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.11 Distal Cover . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
4.12 Proximal Plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
4.13 Septum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
4.14 Depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
5 Terms Used in Describing the Entire Holococcolith . . . . . . 215
5.1 Cavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
5.2 Septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
5.3 Solid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
5.4 Morphological Types . . . . . . . . . . . . . . . . . . . . . . . . 215
5.5 Central Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
5.6 Structural Types . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
5.7 Orientation in Profile . . . . . . . . . . . . . . . . . . . . . . . . 216
5.8 Orientation in Plan View . . . . . . . . . . . . . . . . . . . . . 216
5.9 Structures Spanning Central Area . . . . . . . . . . . . . . . 216
5.10 Structures Closing Central Area . . . . . . . . . . . . . . . . 218
6 Crystallography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
6.1 Crystallographic Orientation . . . . . . . . . . . . . . . . . . . 219
7 Nannolith Shapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
7.1 Dibrachiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
7.2 Compact . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
7.3 Radiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
7.4 Rod-Shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
7.5 Central Body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
7.6 Free Rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
7.7 Convex Outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
7.8 Stellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
8 Additional Taxon-Based Terminology . . . . . . . . . . . . . . . . . 222
8.1 Braarudosphaeraceae . . . . . . . . . . . . . . . . . . . . . . . . 222
8.2 Ceratolithaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
8.3 Discoasters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
8.4 Fasciculiths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
8.5 Helicosphaera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
8.6 Nannoconids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
8.7 Sphenoliths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
xvi Contents

9 Brief Note on Classification and Illustration of Major

Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
10 Geological History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
11 Calpionellids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
2 General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
3 Geological History and Distribution . . . . . . . . . . . . . . . . . . . 265
4 Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275

Species List . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317
About the Author

Dr. Sreepat Jain is a Professor at the Department of Applied Geology,

Adama Science and Technology University, Adama, Ethiopia. With over two
decades of teaching and research experience, Prof. Jain holds two doctorates,
one from India and the other from the United States. Prof. Jain has been
awarded the “TA Excellence in Teaching” for meritorious teaching in the
United States and has also received the prestigious “Prof. S. K. Singh
Memorial Gold Medal” for best research paper from the Palaeontological
Society of India (India). He has published several research articles in national
and international peer-reviewed journals and has authored four books, two
of which, Fundamental of Physical Geology (Springer) and Fundamental
of Invertebrate Palaeontology: Macrofossils (Springer) are recommended
for undergraduate Earth Science courses at several American universities.
Both these Springer books have also received excellent international reviews.
Prof. Jain also serves a reviewer for several international geoscience journals.

xvii
 Chitinozoa
1

also proved useful in delineating the Ordovician–

1 Introduction
Silurian GSSP boundaries (Vandenbroucke 2004;
Verniers and Vandenbroucke 2006), auxiliary
The Chitinozoa (Greek words “chitin” = organic;
GSSP boundaries (Hennissen et al. 2010), and
“zoa” = animal) are flask-, urn-, or bottle-shaped,
stage boundaries (Bergström et al. 2009; Webby
hollow organic vesicles of uncertain biological
et al. 2010a). The latter includes the GSSP of the
affinity, first described by Eisenack (1930) from
Sandbian (Vandenbroucke 2004), Pridoli (Kriz
the erratic boulders strewn near the town of
et al. 1986), Lochkovian (=Silurian–Devonian
Königsberg in Eastern Prussia (present-day
boundary) (Paris et al. 1981), Pragian (Chlupác
Kaliningrad, Russia). The term “Chitinozoa”
et al. 1985), and Famennian (Paris et al.1996).
was introduced because it was thought that their
The GSSP is the Global Boundary Stratotype
walls were of chitin (a naturally occurring
Section and Point, a reference point on a strati-
biopolymer; a polysaccharide) and that the vesi-
graphic section that defines the lower boundary of
cles were of a zoological origin (Eisenack 1931).
a stage on the geologic time scale..
The chitinozoans first appeared in the Early
Ordovician, evolved rapidly in the Paleozoic, and
thereafter went extinct by the end of the Devonian
(Fig. 1(1)). Although few chitinozoans have been
2 General Morphology
reported from the Carboniferous and Permian, but
The chitinozoan vesicle (Fig. 2(1)) ranges from 30
these are considered either suspect or reworked.
to 1500 lm, but most are between 150 and
Recently, the earliest chitinozoans have been
300 lm long (Gabbott et al. 1998). The vesicle
reported from the Cambrian (*510 Ma), from
possesses a two-layered *5-lm thick wall com-
southern China (Shen et al. 2013) (see Fig. 1).
posed of a dark brown or black chitin-like sub-
These earliest forms (Eisenackitina? sp.) are
stance called pseudochitin (Fig. 2(1)). Although,
well-preserved phosphatized flask-shaped vesi-
so far, even in well-preserved specimens, no
cles (see Fig. 1(2–10).
chitin-related organic compounds have been
The fact that the chitinozoans occur in almost
reported (Jacob et al. 2007). The chitinozoan
all types of marine deposits, and that their walls
vesicle shows three major morphological types
are resistant to oxidation, thermal alteration, and
(Fig. 2(1, 3, and 4)); the vesicle also shows vari-
the recrystallization of rock matrix, makes them
ations in its shape (Fig. 2(5 and 6)). However, the
an indispensable tool for regional and global
basic element of a vesicle is the bulging chamber
biostratigraphic correlation. They are among the
called the Central cavity that once housed the
most useful index taxa for the Ordovician–Sil-
organism (Figs. 2 and 3).
urian duration (Tables 1, 2, 3 and 4) and have

© Springer Nature India Private Limited 2020 1

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_1
2 1 Chitinozoa

Table 1 Chitinozoan biozonation (465–459 Ma) (continued…)

2 General Morphology 3

Fig. 1 Cambrian Stage 5 chitinozoan Eisenackitina? sp. from Gaotai Formation in Duyun, Guizhou, China (after Shen
et al. 2013). 1: Stratigraphic position of Eisenackitina? sp., showing that Chitinozoa have their roots in the Cambrian
explosion, and illustrating possibility of benthic origin for the group (after Shen et al. 2013). Genus-level diversify
curve of marine metazoans is after Alroy et al. (2008), and chronological distribution and biodiversification (10–70:
Mean number of species per million years) of chitinozoans are based largely on Grahn and Paris (2011). GOBE—Great
Ordovician Biodiversification Event. 2–10 after Shen et al. (2013). 2–3: Vase-shaped vesicle; 3: Detail of aperture
with flaring collarette; 4: Detail of copula with perforations (arrowed) around base. 5–7: Vase-shaped vesicle with
broken aperture: 5: Lateral view; 6: Posterior view showing hollow copula blocked by residue near boundary between
copula and vesicle bottom; 7: Reverse view of 5 showing perforations (arrowed) around base of copula. 8–10: Internal
mold showing broken aperture and copula with laterally expanding edge: 8: Lateral view; 9: Posterior view showing
blocked vesicle; 10: Detail of perforations? (arrows) around edge of the copula
4 1 Chitinozoa

Table 2 Chitinozoan biozonation (458–444 Ma) (continued…)

2.1 Chamber Shape 2.1.2 Spherical

The shape of the chamber also varies greatly
from being lenticular to cylindrical (Fig. 4(1–7)).
A note on the naming of chamber shapes is
briefly enumerated below (for terminology see
Fig. 4(8 and 9)). However, it must be kept in
mind that the collarette (see Fig. 2(1); detailed
later in the chapter) is not considered when
defining chamber shapes. Major chamber shapes
are as follows:
2.1.1 Lenticular
The chamber width is significantly larger than
the length (D > L), the margin is round, and the
base is convex (Fig. 4(1)). Examples: Calpi-
chitina and Fungochitina.
The chamber width is equal to or close to its length
(D  L), margin is inconspicuous, and the base is
rounded with a shoulder (or exceptionally evagi-
nated as in Margachitina elegans) (Fig. 4(2)).
Examples: Hoegisphaera and Sphaerochitina.
2.1.3 Hemispherical
The chamber length is close to half the chamber
diameter (L = D/2), base is flat, margin is
sharp or blunt, and the shoulder is conspicuous
(Fig. 4(3)). Examples: Bulbochitina and
Cyathochitina.
2.1.4 Ovoid
The length of the chamber is greater than the
chamber diameter, but less than three times the
2 General Morphology 5

Table 3 Chitinozoan biozonation (443–420 Ma) (continued…)

6 1 Chitinozoa

Table 4 Chitinozoan biozonation (410–359 Ma)

2 General Morphology 7

Fig. 2 Chitinozoan morphological features, types of vesicle wall surfaces and vesicle shapes. 1–4: Main
morphological features of the three principal types of chitinozoan vesicles (modified after Paris 1981); 5–6: Variations
in vesicle shapes

diameter (3D > L > D), base is convex to roun- Angochitina, though, more rarely truncated as in
ded, and the margin and flexure are inconspicu- Lagenochitina dalbyensis or ogival as in
ous (Fig. 4(4)). Examples: Desmochitina and Lagenochitina conifundus.
8 1 Chitinozoa

Fig. 3 Additional morphological features of a chitinozoan showing chamber details including sleeve, operculum, and
attachment (modified after Paris 1981; Paris et al. 1999)

2.1.5 Conical 2.1.7 Cylindrical

With straight, tapering flanks, flat base, and sharp
or blunt margin (Fig. 4(5)). Examples: Bursa-
chitina, Euconochitina, and Cyathochitina.
2.1.6 Claviform
The chamber length exceeds three times the
maximum diameter (L > 3D), base is rounded,
margin is inconspicuous, and the shoulder is
absent (Fig. 4(6)). Examples: Clavachitina and
Laufeldochitina.
The chamber length is several times that of the
diameter, flanks are straight and parallel, the base
is flat, and the margin is sharp or blunt (Fig. 4
(7)). Example: Rhabdochitina.
The vesicle’s oral end (apertural pole) bears
the aperture which is usually produced into a
neck; the aboral end (toward Apex) is broader
and closed (Figs. 2(1–4), and 3). The aperture is
occluded by a separate operculum (Figs. 2(3),
and 3(1–3)), whose form and position are of
2 General Morphology
Fig. 4 Chamber shapes (1–
7) and criteria for chamber
measurement (8–9)
great taxonomic importance. The aperture is
either situated directly on the chamber (Fig. 2(3))
or at the distal end of a tube-like neck (Fig. 3(3)).
The apertural pole (Fig. 2(1–4)) represents the
top of the vesicle such that in a chain-like
structure, the upper (or last) vesicle possess a free
aperture (Fig. 3(6 and 7)). In certain genera, the
operculum is deeply recessed within the neck
(i.e., the apertural tube) so that adhesion of the
adjacent vesicle is achieved by a basal, tubular
appendage called the copula (Fig. 3(2–5)).
9
2.2 Ornamentation and Surface
Patterns
The junction between the flanks and the bottom
of the chamber (the anti-apertural end) is called
the margin (=Basal margin; see Fig. 5(1)). Both
the shape (Fig. 5(2–5)) and the ornamentation
(processes, Fig. 5(1); carina, Fig. 5(6–15); and
spines, Fig. 5(16–22)) and surface patterns of the
wall (=Sculpture: Fig. 5(1 and 23–27)) are of
taxonomic importance for suprageneric and
10 1 Chitinozoa

Fig. 5 Margin, Carina, ornamentation, and sculpture. 1–5: Variability in vesicle margin; 6–15: Variability in Carina;
16–22: Ornamentation of the vesicle wall; 23–27: Surface patterns of the vesicle wall (Sculpture) (modified after Paris
1981; Paris et al. 1999)

generic assignment (see also Paris et al. 1999).
The carina (Fig. 5(6–15)) is an important exter-
nal structure and corresponds to an extension
(annular evagination) of the outer layer, perpen-
dicular to the axis of the vesicle and may be
located either below, on, or above the margin. It
can be thick (as in Laufeldochitina embranous,
Cyathochitina kuckersiana, and Pterochitina
perivelata), may extend horizontally (as in
Cyathochitina vaurealensis), or flare anti-
aperturalwards (as in Laufeldochitina). It may
also be complete (as in Cyathochitina), perfo-
rated, reticulated (as in Sagenachitina and Bal-
tochitina) or laciniated (as in Pogonochitina
2 General Morphology
spinifera). The carina may be associated either
with a glabrous or spiny wall and/or apical
structures. But the carina does not coexist with
processes (=homologous elements; see Fig. 5
(1)). The chitinozoans also exhibits varied orna-
mentation that includes all kinds of spiny
extensions longer than 2 µm, occurring on the
vesicle (Fig. 5(1 and 16–22)). They could be
simple (Fig. 5(16)), bifurcated or branching
spines (Fig. 5(17)), bi- or multi-rooted spines
(Fig. 5(18)), anastomosed spines or those con-
nected by a mesh-like structure (Fig. 5(19)),
spines in rows (Fig. 5(20)), crowns or crests or
randomly distributed on part or all of the vesicle
(Fig. 5(21 and 22)). The spiny ornament, which
is usually hollow, may coexist with any of the
other elements (i.e., carina, and apical structures,
except for a glabrous surface). It is of prime
importance to distinguish eroded spines (rounded
scars) from a truly glabrous surface. Spines,
when located on the margin, and forming a
crown are then called, processes (Fig. 5(1)).
They display the same range of complexity as the
spines occurring on the flanks (Fig. 5(1)), but in
addition, they may have a spongy or a cell-like
texture (as in Plectochitina). The surface patterns
(Fig. 5(23–27)) on the chitinozoan wall is also
very varied and can be smooth or scabrate, ver-
miculate (Fig. 5(23)), foveolate (Fig. 5(24)), felt-
like (Fig. 5(25)), spongy (Fig. 5(26)), or micro-
granulous surfaces, including tubercles and cones
(Fig. 5(27)). When the granules are <2 µ high,
the wall surface is regarded as glabrous.
A glabrous surface may or may not be associated
with a carina or with apical structures (as in
Armoricochitina reticulifera), ridges (as in
Laufeldochitina stentorin) or ribs (as in Marga-
chitina catenaria).
The other two external structures, beside car-
ina, are the Apical structures (such as the scar,
callus, mucron, copula, and peduncle; see Figs. 2
and 3) and Sleeve (Fig. 3(2)). The apical struc-
tures usually serve as a linkage medium with the
operculum of the preceding vesicle. The Sleeve
(Fig. 3(2)) corresponds to a partially or totally
unstuck outer layer that may extend beyond the
margin and even far beyond the base of the
vesicle (as in Pellichitina, Cutichitina, and
11
Velatachitina). This term is not extended to
various types of vertical folding (as in Calpi-
chitina velata) or to irregular detachment of the
outer layer (as in Desmochitina juglandiformis).
The collarette, a fragile membranous contin-
uation of the vesicle wall, may be present around
the aperture (Fig. 6(1)). The plug, sealing this
aperture, is either called an operculum (Fig. 2(3))
or a prosome (Fig. 2(2)). Additionally, collarette
(Fig. 6(1–5)), Neck (Fig. 6(9–11)), and the base
of the chitinozoan organism (Fig. 6(1 and 12–
18)) can also be varied. The prosome closes the
apertural tube or the opening of the chamber, but
when neck is lacking, it is called an operculum
(Fig. 7(3)). Both operculum and prosome are
extended aborally by Rica (Fig. 7(2 and 3)), a
membranous expansion lining the upper part of
the chamber (Fig. 7(1–3)). The operculum has a
disk-like shape with a flat, slightly concave or
convex distal surface (Fig. 7(3)) and is extended
toward the chamber with a flaring membranous
expansion, the Rica (Fig. 7(3)).
Many chitinozoans are found united in long
chains or clusters, where the vesicles are welded
together at the operculum (i.e., the oral pole) and
at the base (i.e., the aboral pole) (see Fig. 7).
2.3 Linear Linkage Arrangements
Most vesicles are found isolated, but in rare
instances, they occur in clusters and chains.
Although, it is assumed that these isolated vesicles
were part of a larger chain-like structure, called
Linear Catenary Structures. The latter represent
the linkage between two successive vesicles,
reflecting the connection of the aperture of
one individual with the bottom of the next one
(Fig. 7). In this regard, three types of linear
arrangements are noted (Fig. 7). The linkage by
simple juxtaposition (Fig. 7(1)) is the simplest but
rare due to its fragility in preservation. In this, the
lips (i.e., the apertural borders) of one individual is
in contact with the convex anteapertural part of
the next one (as in Angochitina, Ramochitina,
Ancyrochitina, and Cyathochitina). In the linkage
with simple adherence, another rare type, the
flaring lips envelope the convex anti-apertural part
12
Fig. 6 Collarette, Neck, and Base (1). 2–8: Variability in collarette; 9–11: Variability in neck; 12–18: Variability in
base
of the following specimen with no contact
between the apex and the operculum of two suc-
ceeding vesicles. It is noted in taxa that have a
globulous chamber and (or) those without neck
(as in Eisenackitina and Desmochitina). The most
common type is the linkage with double adhesion
(or adherence) (Fig. 7(2)). The contact is large
between the lips and the rounded bottom of the
following vesicle which is strengthened by the
fixation of its mucron (or copula) to the center of
the operculum of the preceding vesicle, best noted
in Cingulochitina, Linochitina, Desmochitina,
Urochitina, and Pterochitina (Fig. 7(2)). The third
type is the reinforced linkage, a more advance
type, where modifications of the vesicle occur
1 Chitinozoa
(Fig. 7(4)). The apex of the chamber extends
toward the anteapertural pole and covers the op-
erculum of the preceding vesicle so that it
becomes totally dependent of the other (as in
Margachitina). For all the above, it should be
noted that no internal communication between
vesicles has so far been noted, and hence, clusters
are not considered to be a “colony.”
A brief description of the terms used in the
chapter (stratigraphic occurrence of major terms in
Fig. 8) and elaborated below to help better under-
stand the complex chitinozoan morphology (terms
are after various authors: Combaz et al. 1967;
Cramer 1967; Jansonius 1970; Jenkins 1970; Eise-
nack 1968, 1972; Laufeld 1974 and Paris 2006).
3 Terminology 13

Fig. 7 Types of linear linkage arrangements (modified after Paris 1981; Paris et al. 1999)

3.3 Aboral
3 Terminology
It is the part of the vesicle where the base and
3.1 Aboral Pole basal scar occur; aperture is at the opposite end
of it (Fig. 3(1)).
It is synonymous with Base and Basal pole
(Fig. 2(1)).
3.4 Anti-apertural Pole
3.2 Aboral Scar
It is opposite the aperture and, at the lower part of
It is synonymous to the basal scar (Fig. 3(1)). the vesicle (Figs. 2(1) and 3(1)).
14 1 Chitinozoa

Fig. 8 Chitinozoan morphological characters through time (modified after Paris et al. 1999)
3 Terminology
3.5 Apertural Plug
Simple or complex plug seals the aperture. It is
represented either by the Prosome or by the
Operculum (Figs. 2(1 and 2) and 7(2),
respectively).
3.6 Apertural Pole
It is the upper part of the vesicle (Fig. 2(1)).
3.7 Aperture
It is main opening of the vesicle and occurs
adorally, located at a collar, neck, lip, or chamber
(Fig. 2).
3.8 Apex
It is the point of emergence of the axis of sym-
metry on the base of the chamber (Fig. 2(1)).
3.9 Apical Structure
This includes (basal) scar, (basal) callus, mucron,
copula, and peduncle (Figs. 2(1) and 3(1)).
3.10 Appendices
These are singular processes at the basal edge;
they may be simple or branched (=synonymous
with processes; Fig. 5(1)).
3.11 Axis
It is an imaginary line that joins the center of the
aperture to the apex (Fig. 2(1)). It is the axis of
symmetry of the vesicle.
15
3.12 Basal Callus
It is the thickened portion of the vesicle wall at
the center of the base (Fig. 3(1)).
3.13 Basal Edge
It is the bent portion of the vesicle wall that
forms an edge separating the base and the
chamber flank; it often bears appendices, carina,
or other elements of ornamentation (Fig. 5(1)).
3.14 Basal Margin
It is synonymous with the basal edge (Fig. 3(1)).
3.15 Basal Pore
It is the opening at the center of the basal scar
(Fig. 3(1)).
3.16 Basal Scar
It is the hollow (usually circular) surface at the
center of the base, or at the basal callus (Fig. 3
(1)) and reflects the connection between the
interiors of subadjacent chambers.
3.17 Base
It is the portion of the vesicle wall situated
aborally to the basal edge. It is part of the
chamber opposite to the aperture (=anti-apertural
end or chamber bottom) (Fig. 2(1)).
3.18 Callus (Basal)
It is the short stub-like thickening on the apex
(Fig. 3(1)).
16
3.19 Carina
It is the circular membrane that extends distally
at the basal edge (like the brim of a hat). It is
located below, on, or above the margin (Fig. 5(1
and 6–15)).
3.20 Catenary Structure (Chain-Like
Structure)
Illustrates vesicles connected “aperture to base,”
along their axis (straight, curved or coiled chain)
or vesicles connected “neck to margin,” with a
coiled pattern (Fig. 7).
3.21 Central Cavity
It is the inner cavity corresponding to the
Chamber (Fig. 2(1)).
3.22 Chain
Several vesicles are linked longitudinally with
one another (Fig. 5).
3.23 Chamber
It is the part of the vesicle (generally bulging in
nature) situated below the neck or below the
collarette when present (Fig. 2).
3.24 Cluster
It is the group of vesicles of the same species and
connected by their flanks (with free aperture)
(Fig. 5).
3.25 Coiled Chain (=Helicoidal Chain)
The vesicles are connected either “neck to mar-
gin,” i.e., with a free aperture, or “aperture to
1 Chitinozoa
base,” i.e., aperture linked to the succeeding
vesicle (Fig. 7).
3.26 Collar
It is the end of the vesicle that expands orally;
part of the oral tube that extends from the neck or
chamber (in forms with no neck) to the aperture
(Fig. 3(1, 3 and 4)).
3.27 Collarette (=Collar)
It is the thinned cylindrical or flaring part of the
neck, or of the vesicle wall when the neck is
absent; it surrounds the aperture (Fig. 2(1)).
3.28 Copula
It is the tubular, membranous expansion sur-
rounding the apex (Fig. 3(3–5)).
3.29 Crests
It is the vertical rows of free or connected spiny
ornamentation, or of web-like to continuous
membranes (Fig. 5(20)).
3.30 Diameter (=D)
The term refers to the maximum diameter of the
chamber (Fig. 4(8 and 9)).
3.31 External Structures
It includes carina, apical structures, and the
sleeve (Fig. 6).
3.32 Fenestrate
It is a naturally perforate membrane, carina,
collar, etc. (Fig. 6(10 and 11)).
3 Terminology
3.33 Flank
It is the portion of the vesicle wall that extends
between the basal edge and, either the shoulder
flexure, or collar (Fig. 2(1)).
3.34 Flexure
It is the concave zone separating the flank from
the neck (Fig. 2(1)).
3.35 Glabrous
It is the state of the vesicle surface that lacks the
spiny ornamentation (Fig. 5(23)).
3.36 Inner Layer
It is the internal membrane of the wall usually
acting as a framework for the vesicle (Fig. 2(1)).
3.37 Laciniated
It is the deeply indented borders of a carina
(Fig. 5(8 and 9)).
3.38 Linear Chain
It is the straight, curved, or coiled catenary
structure where the apertural pole of one vesicle
is fixed to the base of the following one; the
aperture is not free (Fig. 7).
3.39 Linkage Structures
These intervening elements in the connection of
two successive vesicles in a catenary structure
(operculum, apical structures lips, base) (Fig. 7).
17
3.40 Lip
It is the swelling around the aperture of a neck-
less and collarless vesicles. It is also the distal
part of the collarette (or of the neck), surrounding
the aperture (Fig. 2(1)).
3.41 Longitudinal Axis
It is the geometrical axis linking the centers of
the aperture and base (Fig. 2(1)).
3.42 Margin
It is the transition zone between the base and the
flanks (Fig. 2(1)). It may be inconspicuous,
rounded, blunt, or sharp. The margin bears major
morphologic elements such as carina, processes,
and sleeve.
3.43 Mesh-Like
Spiny ornamentation is making up a network or a
trelliswork on the surface of the wall (Fig. 5
(19)).
3.44 Mucron
It is the thickened, raised rim surrounding the
apical pit (Fig. 3(3)).
3.45 Neck
It is the tubular structure expanding apertural-
ward from the chamber; it is frequently termi-
nated by a collarette (Fig. 2(1)). In collarless
forms, it is the portion of the vesicle that extends
between the collar or aperture and the flexure.
18
3.46 Operculum
It is the disk-like plug sealing the aperture of the
vesicles and lacking a neck (Fig. 7(2 and 3)). It is
a membranous expansion flaring
anti-aperturalward (=Rica), and its thickness may
equal or slightly exceed the thickness of the
vesicle wall.
3.47 Oral Scar
It is the circular boss with a pit or pore at its
center, situated at the external surface of the
operculum (Fig. 3(1)).
3.48 Oral
The end of the vesicle where the operculum and
aperture occur (Fig. 2(2)).
3.49 Ornamentation
They are the larger external–morphological ele-
ments of the vesicle, such as appendices, pro-
cesses, auricles, and spines (Figs. 2(1) and 6(16–
22)). The ornamentation does not include the
surface sculpture (Fig. 6(23–27)).
3.50 Outer Layer
It is the external membrane forming the vesicle
wall together with the inner layer (Fig. 2(1)); the
vesicle ornaments (carina, processes, spiny or-
namentation) arise from evaginations of this
outer layer, which may also be glabrous (Figs. 2
(1), 3(3) and 6).
3.51 Peduncle
It is a solid, short or elongated cylindrical
structure extending from the apex (Fig. 2(3)).
1 Chitinozoa
3.52 Perforated
It refers to closely distributed holes occurring on
the carina (Fig. 5(12 and 13)) or on the collarette
(Fig. 6(8)).
3.53 Processes
These can be simple or complex, short or very
long spines (Fig. 6(1)). They are usually hollow
but may have a cell-like structure.
3.54 Prosome
This is the internal plug situated at the base of the
neck (Fig. 2(2)). It may be simple (disk-like;
Fig. 7(3)) or complex (tubular structure with
several or numerous horizontal septa, i.e., “ac-
cordion pleated prosome;” Fig. 2(2)). Prosome is
not involved in vesicle linkage.
3.55 Rica
It is a membranous flanging part of the prosome
or operculum, lining the upper part of the
chamber (Figs. 2(1–3) and 6(1–3)).
3.56 Sculpture
It includes the morphology of the vesicle surface
(Fig. 5(1)) and seen on the external and internal
surfaces, operculum, and large ornamentation
elements. Sculpture types includes laevigate,
granulate, verrucate, reticulate, rugate, or spongy
(Fig. 5(23–27)).
3.57 Septa
It is the horizontal membranous partition (from a
few to more than 20) within the prosome (Fig. 2
(2)).
3 Terminology
3.58 Shoulder
It is the convex area at the top of the flanks, just
below the flexure (Fig. 2(1)).
3.59 Sleeve
It is the partially or totally attached membrane
(usually the outer layer) covering the vesicle and
extending beyond the base (Fig. 3(2)).
3.60 Spines
These include all kinds of simple to complex
elongate expansions of the outer layer (Fig. 5
(16)); spine length must be at least twice their
width and exceeds 2 µm; and they are most
frequently hollow.
3.61 Spiny Ornamentation
These include vesicles bearing spines, crests,
and/or processes randomly distributed or arran-
ged in rows or crowns (Fig. 5(16–22)).
3.62 Vesicle
It is the basic chitinozoan individual, including
the wall of the chamber and the neck (and col-
larette when present) as well as the apertural plug
(Fig. 2(1)).
3.63 Wall
It is the organic envelope of a chitinozoan and
includes both the outer and inner layer (Fig. 3(2)).
4 Chitinozoan Affinity
The chitinozoans have been linked to diverse life
forms across different Kingdoms—Protozoa
(Testacea, Foraminifera, Dinoflagellata),
19
Metazoa (Annelida, Gastropoda, Graptolites),
and Fungi (Riding 1980). They have been sug-
gested as being protected eggs or cysts produced
by a marine metazoan due to their arrangement in
elongate organic-walled sheets (“cocoons”)
(Kozlowski 1963). This “egg case hypothesis”
has now been accepted as the most plausible
interpretation (see Miller 1996; Gabbott et al.
1998; Paris and Nõlvak 1999; Armstrong and
Brasier 2005).
5 Classification
The classification by Paris (1981) is the followed
by most workers where the highest subdivision is
the order (Table 5). The order Operculatifera
groups all chitinozoans with an operculum,
whereas order Prosomatifera belongs to those
with a prosome (Table 5). The lower ranks are
families (such as Desmochitinidae), followed by
subfamilies (such as Desmochitininae), genera
and species; infraspecific ranks such as sub-
species and varieties (formas) are avoided
(Table 5). Later, Paris et al. (1999) proposed a
suprageneric classification based on diagnostic
features (Fig. 8) whose hierarchy was established
on statistical and evolutionary grounds.
However, some workers still prefer to use an
alphabetical listing of taxa (as proposed by
Eisenack 1972); they consider the suprageneric
classification of organisms of unknown biologi-
cal affinities (as chitinozoans) to be meaningless.
In this suprageneric classification, the orders are
based on the nature of the vesicle seal, either an
operculum for the order Operculatifera or a
prosome for the order Prosomatifera (see
Table 5; see also Paris et al. 1999 for details).
The Operculatifera are characterized by an op-
erculum, reduced oral tubes (usually with a col-
larette, but lacking a neck). Desmochitinidae is
the sole family with six subfamilies; Desmo-
chitina (Early Ordovician–Late Silurian; Fig. 9)
had a relatively small subspherical vesicle with
short lips but no neck and was commonly united
in chains. The Prosomatifera (Table 5) have
prosome and well-developed necks. Prosomat-
ifera has two families, the Conochitinidae
20 1 Chitinozoa

Table 5 Chitinozoan classification (see text for explanation) (after Paris et al. 1999)
5 Classification 21

Fig. 9 Representative examples of Desmochitinidae (see text for explanation and Table 5)
22 1 Chitinozoa

Fig. 10 Representative examples of Conochitinidae (see text for explanation and Table 5)
5 Classification 23

Fig. 11 Representative
examples of Lagenochitinidae
(see text for explanation and
Table 5)
24
(Fig. 10) and the Lagenochitinidae (Fig. 11),
distinguished by the relationship between the
chamber and the neck in Lagenochitina (Early
Ordovician–Early Silurian), the operculum is
recessed within the neck and possesses a rela-
tively large vesicle with a cylindrical neck.
Representative genera belonging to Desmochi-
tinidae, Conochitinidae, and Lagenochitinidae
are illustrated in Figs. 9, 10, and 11, respectively.
6 General History and Distribution
The flask-shaped Eisenackitina? is the oldest chiti-
nozoan (*510 Ma; Shen et al. 2013) (Fig. 1(1)).
This occurrence is followed by a gap, and then,
during the Ordovician (485.4 ± 1.9–
443.8 ± 1.5 Ma), morphologically smooth vesicles
of desmochitinid and conochitinid types are noted;
the conochitinid stock gradually dwindled through
the Silurian and died out at its end. The more
complex tanuchitinids appeared in the Arenig
(Ordovician), while the sphaerochitinids appeared a
little later in the Caradoc (Ordovician) that charac-
terize Silurian and Devonian assemblages. Chiti-
nozoan with stout basal horns and appendix-bearing
forms are typical of many Silurian and Early
Devonian assemblages, although the Late Devonian
forms are more often covered with short spines.
Chains of chitinozoan are prominent in Silurian and
Devonian assemblages. In the Carboniferous,
chitinozoans are rare. Although, few specimens
have been reported from the Permian (Tasch 1973),
but these are considered as fungal spores.
References
Alroy, J., Aberhan, M., Bottjer, D. J., Foote, M., Fürsich,
F. T., Harries, P. J., et al. (2008). Phanerozoic trends
in the global diversity of marine invertebrates.
Science, 321(5885), 97–100.
Armstrong, H. A., & Brasier, M. D. (2005). Microfossils
(p. 304). Oxford: Blackwell.
Bergström, S. M., Chen, X. U., Gutiérrez-Marco, J. C., &
Dronov, A. (2009). The new chronostratigraphic
classification of the Ordovician System and its rela-
tions to major regional series and stages and to d13C
chemostratigraphy. Lethaia, 42, 97–107.
1 Chitinozoa
Chlupáč, I., Lukeš, P., Paris, F., & Schönlaub, H.
P. (1985). The Lochkovian-Pragian boundary in the
Lower Devonian of the Barrandian area (Czechoslo-
vakia). Jahrbuch der Geologischen Bundesanstalt,
128, 9–41.
Combaz, A., Calandra, F., Jansonius, J., Millepied, P.,
Poumot, C., & Van Oyen, F. H. (1967). Microfossiles
organiques du Paléozoïque. Les Chitinozoaires (2):
Morphographie (pp. 1–58). Éditions du Centre
National de la Recherche Scientifique, Paris.
Cramer, F. H. (1967). Chitinozoans of a composite
section of Upper Llandoverian to basal Lower Gedin-
nian sediments in northern Léon, Spain. A preliminary
report. Bulletin de la Société belge de Géologie, de
Paléontologie et d’Hydrologie 75, 69–129.
Eisenack, A. (1930). Neue Mikrofossilien des baltischen
Silurs – Vorlaufige Mitteilung. Naturwissenschaften,
18, 180–181.
Eisenack, A. (1931). Neue Mikrofossilien des baltischen
Silurs. I. Palaeontologische Zeitschrift, 13, 74–118.
Eisenack, A. (1968). Über Chitinozoen des baltischen
Gebietes. Palaeontographica A, 131, 137–198.
Eisenack, A. (1972). Chitinozoen und andere Mikrofos-
silien aus der Bohrung Leba, Pommern. Palaeonto-
graphica Abteilung A, 140, 64–87.
Gabbott, S. E., Aldridge, R. J., & Theron, J. N. (1998).
Chitinozoan chains and cocoons from the Upper
Ordovician Soom Shale lagerstätte, South Africa:
Implications for affinity. Journal of the Geological
Society, 155, 447–452.
Grahn, Y., & Paris, F. (2011). Emergence, biodiversifi-
cation and extinction of the chitinozoan group. Geo-
logical Magazine, 148(2), 226–236.
Hennissen, J., Vandenbroucke, T. R. A., Chen, X., Tang,
P., & Verniers, J. (2010). The Dawangou auxiliary
GSSP (Xinjiang autonomous region, China) of the
base of the Upper Ordovician Series: Putting global
chitinozoan biostratigraphy to the test. Journal of
Micropalaeontology, 29, 93–113.
Jacob, J., Paris, F., Monod, O., Miller, M. A., Tang, P.,
George, S. C., et al. (2007). New insights into the
chemical composition of chitinozoans. Organic Geo-
chemistry, 38, 1782–1788.
Jansonius, J. (1970). Classification and stratigraphic
application of Chitinozoa. In Proceedings of North
American Paleontological Convention 1969, Part G,
pp. 789–808.
Jenkins, W. A. M. (1970). Chitinozoa. In B. F. Perkins
(Ed.), Geoscience and Man 1. Louisiana: Baton
Rouge.
Kozlowski, R. (1963). Sûr la nature des chitinozoaires.
Acta Palaeontologica Polonica, VIII, 424–445.
Kriz, J., Jaeger, H., Paris, F., & Schönlaub, H. P. (1986).
Pridoli—The fourth subdivision of the Silurian sys-
tem. Jahrbuch der Geologischen Bundesanstalt, 129,
291–360.
Laufeld, S. (1974). Silurian Chitinozoa from Gotland.
Fossiis and Strata, 5, 1–130.
Miller, M. F. (1996). Chitinozoa. In J. Jansonius & D.
C. McGregor (Eds.), Palynology: Principles and
References
applications (pp. 307–336). American Association of
Stratigraphic Palynologists Foundation.
Paris, F. (1981). Les Chitinozoaires dans le Paleozoique
du sud-ouest de 1’Europe (cadregéologique - tude
systematique – biostratigraphie) (Vol. 26, 496 p).
Memoire de la Sociéte géologique et minéralogique de
Bretagne, Rennes.
Paris, F. (1996). Chitinozoan biostratigraphy and palaeoe-
cology. In J. Jansonius & D. C. McGregor (Eds.),
Palynology: Principles and applications (Vol. 2,
pp. 531–552). American Association of Stratigraphic
Palynologists Foundation, Dallas.
Paris, F. (2006). Chitinozoans: A fascinating and myste-
rious microfossil-group. Technical-card and direction
for use. https://doi.org/10.13140/RG.2.1.2462.7682.
Paris, F., Grahn, Y., Nestor, V., & Lakova, I. (1999).
A revised chitinozoan classification. Journal of Pale-
ontology, 73, 549–570.
Paris, F., & Nõlvak, J. (1999). Biological interpretation
and paleobiodiversity of a cryptic fossil group: The
“chitinozoan animal”. Geobios, 32, 315–324.
Paris, F., Laufeld, S., & Chlupáč, I. (1981). Chitinozoa of
the Silurian-Devonian boundary stratotypes in Bohe-
mia. Sveriges Geologiska Undersökning Serie Ca, 51,
1–29.
25
Riding, J. B. (1980). The nature and affinity of the
Chitinozoa—A review. Journal of the University of
Sheffield Geological Society, 7, 262–268.
Shen, C., Aldridge, R. J., Williams, M., Vandenbroucke,
T. R. A., & Zhang, X.-G. (2013). Earliest chitinozoans
discovered in the Cambrian Duyun fauna of China.
Geology, 41, 191–194.
Tasch, P. (1973). Chitinozoa, Part III. In Paleobiology of
the Invertebrate (pp. 821–833). New York, Toronto:
John Wiley & Sons, Inc.
Vandenbroucke, T. (2004). Chitinozoan biostratigraphy
of the Upper Ordovician Fågelsång GSSP, Scania,
southern Sweden. Review of Palaeobotany and Paly-
nology, 130, 217–239.
Verniers, J., & Vandenbroucke, T. R. A. (2006). Chiti-
nozoan biostratigraphy in the Dob’s Linn
Ordovician-Silurian GSSP, Southern Uplands, Scot-
land. GFF, 128, 195–202.
Webby, B. D., Paris, F., Droser, M. L., & Percival, I. G.
(2004). The great Ordovician biodiversification event
(p. 496). New York: Columbia University Press.
 Acritarchs
2

offer the earliest glimpse in assessing the history

1 Introduction
of protistan biodiversity (Knoll 1994, 2006;
Vidal and Moczydlowska-Vidal 1997) (Fig. 1).
The Acritarcha (Greek words: achritos: “uncer-
The protists include organisms with organic
tain”; arché: “origin”) was coined by Evitt (1963)
(acritarchs, dinoflagellates, chitinozoans), cal-
to include marine microplanktons (incorporating
careous (coccolithophores, foraminifers), or
all organic-walled microfossils of unknown ori-
siliceous (diatoms, radiolarians) skeletons.
gin) recorded from the early Paleozoic rocks (541–
The oldest acritarchs (leiosphaerids) have
252 Ma). The Acritarcha (an informal incertae
been recorded in 1.8-billion-year-old (Ga = giga-
sedis group) are small, eukaryotic, unicellular,
annum = billion year ago) rocks, while some
organic-walled, resistant, morphologically varied,
other organic-walled fossils like bacterial sheaths
and long-ranging forms of unknown biological
and coccoids date as far back as 3.4 Ga (Javaux
affinity; a polyphyletic group with multiple clades
et al. 2010; Buick 2010) (Fig. 1). Theses smooth
(Butterfield 2004, 2005).
walls (sphaeromorphs) are considered as the
Most Palaeozoic forms are possible cyst
oldest acritarchs (Javaux 2014), whereas the
stages of diverse algal groups such as dino-
oldest ornamented acritarchs (with concentric
phyceans, prasinophyceans, chlorophyceans, or
striations) are 1.8 Ga and interpreted as probable
zygnemaphyceans (Colbath and Grenfell 1995;
eukaryotes. The oldest spiny acritarchs (acan-
Mendelson 1987; Butterfield 2004). Of them,
thomorphs) dated as 1.5 Ga are decidedly
only a few are of non-algal affinity and may
eukaryotic. Hence, these single-celled acritarchs
represent crustacean eggs or exoskeletal remains
are an important window for our understanding
of higher organisms, plant spores, fungal paly-
of the early life on Earth as they are the first
nomorphs, remains of cyanobacteria, or other
record of complex and larger cells of eukaryotes
groups (Servais et al. 1997; Butterfield 2005).
(animals, plants, fungi, and protists). Acritarchs
Hence, “acritarcha” is a traditional group that
reigned for more than one billion years as the
includes single-celled, organically preserved
most complex organisms on the planet. Apart
vesicles. Most of these vesicles are assumed to be
from bacterial microfossils and Stromatolites, the
eukaryotic, i.e., their cell contains a nucleus and
acritarchs have one of the longest records than
other organelles which are enclosed in mem-
any other fossil group (Fig. 1). More impor-
branes. The filamentous organic-walled micro-
tantly, acritarchs played a role in increasing the
fossils of unknown biological affinities are not
amount of oxygen in the oceans during the
included among acritarchs. The acritarchs are
Neoproterozoic that eventually paved the way for
among the oldest eukaryotes in the fossil record
(Zhang 1986; Yan 1991; Butterfield 2015) and

© Springer Nature India Private Limited 2020 27

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_2
28 2 Acritarchs

Fig. 1 Earliest acritarchs and major acritarch events. 1–3: Mesoproterozoic (1.7–1.4 billion years old) fossils from the
Ruyang Group, China (after Agić 2016). 1: Dictyosphaera macroreticulata is a sphaeromorph with a distinct surface
pattern; 2: Spine-bearing taxon Shuiyousphaeridium macroreticulatum; 3: Double-walled acritarch Pterospermopsi-
morpha insolita, consisting of a central body and an outer envelope; 4: Chronology of major geological, atmospheric, and
biological events (modified after Beraldi-Campesi 2013). The oldest eukaryotic-like fossils (acritarchs; black bar) found
so far (and that possibly need oxygen for metabolic capabilities) are the *3200 Ma old forms recorded from estuarine
environments suggesting that life achieved a relatively rapid global presence and had diversified enough (see Javaux et al.
2010; Buick 2010). By 2500 Ma there is a marked change into an oxygenated atmosphere, although oxygen is likely to
have been accumulating beforehand (see also Ohmoto 2004). These oxygenation events occurred in pulses and are
correlated with a putative origin of oxygenic photosynthesis by cyanobacteria and the tentative emergence of life at
3800 Ma (Lazcano and Miller 1994)

the rise of animals and other large and complex Proterozoic oceans, are widely studied, largely
organisms (Fig. 1). due to their widespread distribution, the variety
The acritarchs, probably the earliest eukary- of types of rock they are recovered from, and the
otic phytoplankton—primary producers in the similarity of their body plan to present-day
1 Introduction 29

non-skeletal plankton. Some Paleozoic acritarchs 2 General Morphology

have proved to be very useful in dating tec-
tonized and metamorphically affected lithological The acritarchs range in size between 30 and
units where no other fossils are preserved 200 µm. However, Buick (2010) suggests that
(Molyneux 1999; Montenari et al. 2000). They organic-walled microfossils >50 µm are termed
have also proved very useful in biostratigraphy as acritarchs. But, recently Javaux et al. (2010)
and paleoenvironmental studies (Huntley et al. recorded well-preserved organic-walled micro-
2006; Munnecke et al. 2012; Wang et al. 2014). fossils that were as big as 300 µm from
A case in point is the distinctive marker Tappa- 3.2 Ga-old shallow-marine siliciclastic rocks in
nia–Dictyosphaera–Valeria assemblage zone the Mesoarchaean (2.8–3.2 Ga) shales and silt-
representing a time interval from *1650 to stones of the Moodies Group, South Africa.
*1400 Ma (Yin et al. 2005; Agić et al. 2015;
Javaux and Knoll 2017; Adam et al. 2017). This
2.1 Sphaeromorphs
assemblage zone has a widespread distribution of
and Acanthomorphs
an ecologically coherent biota recorded from
China (Ruyang Group; Yin 1997; Yin et al.
Acritarchs are either smooth (sphaeromorphs;
2005), Australia (Roper Group; Javaux et al.
Fig. 2(1)) or covered in spines (acanthomorphs;
2001; Javaux and Knoll 2016), Siberia (Kamo
Fig. 2(2)). The sphaeromorphs are the oldest
Group; Nagovitsin 2009), and the USA (Greyson
acritarch group (1.8 Ga; Lamb et al. 2009) (see
Formation, Belt Supergroup, Montana; Adam
Fig. 2(3)). The unornamented (smooth) and
et al. 2017). The assemblage is typified by the
spherical forms are called leiosphaerids. The
process-bearing Tappania, reticulate-walled
sphaeromorphs may, at times, bear a variety of
Dictyosphaera/Shuiyoushaeridium, and concen-
surface ornamentation/sculpture such as mesh-
trically striated Valeria, offering key insights into
work, pores, or corrugation. They also display
the early evolution of the clade, and enabling
several shapes such as spheroidal, ovoidal (el-
practical applications to paleobiogeography and
lipsoidal), and fusiform (spindle-shaped) and
biostratigraphic correlation. Tappania is consid-
range in size from 62 to 216 µm (Lamb et al.
ered within or immediately adjacent to the
2009). The sphaeromorph walls may be psilate,
crown-group Eukarya (but see also Javaux and
scabrate, verrucate, rugulate, fossulate, but not
Knoll 2016), as also Dictyosphaera and Shuiy-
truly spinous.
ousphaeridium (Pang et al. 2013; Agić et al.
The acanthomorphs (Figs. 2(4–12) and 3), on
2015). Hence, these oldest eukaryotic organisms
the other hand, are ornamented with spiny pro-
also carry valuable phylogenetic and paleobio-
trusions called “Processes” (Fig. 3(1)) that vary
logical implications for constraining the early
in size, shape and distribution along the vesicle
evolution of eukaryotes (Adam et al. 2017).
surface (Fig. 3). These, at the Cambrian–
As a note, it must be mentioned that a broader
Ordovician boundary, are remarkably complex
group of organically preserved, acid-extracted
and diverse in shape (Fig. 4). Here, it must be
microfossils are called palynomorphs or palyno-
mentioned that some palynologists use “sculp-
flora. These include acritarchs, dinoflagellates,
ture” for elements that are <5 µm long, and
pollen, spores, chitinozoans, and fungi. Palynol-
“processes” for larger element sizes.
ogy (Greek word paluno meaning “strew” or
“dust”; the study of dust) is the scientific disci-
pline that studies these fossils, and their modern 2.2 Morphogroups
counterparts. Palynology is an interdisciplinary
science drawing from and applied to geology, The acritarchs, based on their complex processes
botany, climate studies, and forensics. This and diverse shapes are broadly divided into the
chapter deals with acritarchs. following morphogroups (see Fig. 2):
30 2 Acritarchs

Fig. 2 Common acritarch morphotypes and their stratigraphic ranges. 1–2: Those without ornamentation (spines) are
called sphaeromorphs, and spine-bearing ones are called acanthomorphs (see text for explanation). 3: Stratigraphic
ranges of common acritarch morphotypes. Arrows indicates those subgroups that have a post-Permian record (this
record is poorly known and thus, not shown). The width of lines roughly corresponds to the number of species recorded
for the particular interval. 4–13: Common acritarch morphotypes. All acritarchs are vesicles made by an organic wall
that varies in thickness and ultrastructure. Some acritarchs may also bear an opening (excystment) (see text for
explanation)
2 General Morphology 31

Fig. 3 Acanthomorph morphology (1), types of surface sculpture (2–12), process terminations (13–17) and excystment
openings (18–23). See text for explanation
32 2 Acritarchs

Fig. 4 Ornamentation of selected Cambrian–Devonian acritarchs (not to scale) (modified after Martin 1993). 1:
Retisphaeridium howellii; 2: Cristallinum cambriense; 3: Timofeeva lancarae; 4: Vulcanisphaera turbata; 5:
Cristallinum randomense; 6: Vulcanisphaera africana; 7: Veryhachium minutum; 8: Cymatiogalea velifera; 9:
Cristallinum dentatum; 10: Striatotheca principalis; 11: Dilatisphaera wimanii; 12: Tunisphaeridium eisenackii; 13: D.
williereae; 14: Psenotopus condrocheus; 15: Neoveryhachium carminae; 16: Cymbosphaeridium pilar; 17: Daillydium
pentaster; 18: Umbellasphaeridium saharicum; 19: Ammonidium exoticum; 20: Winwaloeusia ranulaeforma
2 General Morphology 33

2.2.1 Sphaeronorphs (Spharia = Ball) features of their body plan (Fig. 3). These are (see
These are simple spherical-shaped (Fig. 2(4)). Fig. 3(1)): (a): the resilient carbonaceous vesicle
(the hollow body of an acritarch; (b): the round or
2.2.2 Acanthomorphs split openings in the wall, called excystment
(Acantha = Thorn) structure or Pylome, and (c): processes (spiny
These have spherical bodies with spines that protrusions). Hence, an acritarch can be described
usually open into the body (Fig. 2(5)). Examples: based on vesicle morphology, type of excystment
Baltisphaeridium, Micrhystridium, and opening and process morphology (including the
Multiplicisphaeridium. process tip morphology) (Fig. 3(1)). The vesicle
and excystment opening are briefly discussed
2.2.3 Prismatomorphs (Prisma = Prism) below (the processes have been dealt above).
These possess prismatic to polygonal forms and
the edges form a flange or crest that may be
2.3 Vesicle
serrated (Fig. 2(6)).
An acritarch is composed of a hollow central
2.2.4 Netromorphs (Netro = Spindle)
body called a Vesicle (Fig. 3(1)). The vesicle
These have a fusiform spindled-shape body with
wall thickness is generally a fraction of a µm, but
one or more spines (Fig. 2(7 and 8)). Example:
may reach *4 µm (Eiserhardt 1989). In Tas-
Leiofusa.
manites newtonii it is 2–7 µm thick, in
Baltisphaeridium it is 2–3 µm and in Orthos-
2.2.5 Herkomorphs (Herkos = Wall)
phaeridium insculptum, the wall is *2 µm thick.
These have a roughly spherical form and divided
The wall of an acritarch is made of
into polygonal fields (Fig. 2(9)). Examples: Ric-
sporopollenin-like compound called Polyter-
ulasphaera and Herkomorphs.
penes. The sporopollenin is an oxidative polymer
of carotenoids and carotenid esters (Brooks and
2.2.6 Oomorphs (Oon = Egg)
Shaw 1968; see also Bernard et al. 2015). The
There have an egg-shaped form with ornamen-
acritarch vesicle wall is made up of one or two
tation confined to the poles (Fig. 2(10)).
layers that may be joined or separate. If separate,
the terms endo-, meso-, and ectophragm (pe-
2.2.7 Diacromorphs (Akron = Summit)
riphragm) are used (Fig. 3(1)). The periphragm
These are spherical to ellipsoidal forms with or-
(= ectophragm), with or without endophragm,
namentation confined to the poles (Fig. 2(11)).
forms projections (processes/spines) that may be
Example: Acanthodiacrodium.
regularly or irregularly distributed, solid or hol-
low (Fig. 3). The latter two are important diag-
2.2.8 Polygonomorphs (Gonia = Angle)
nostic characters for species identification as only
These have a body defined by the number and
one is present, solid or hollow, in a species.
position of the spines, that are often triangular or
The patterns on the surface (surface sculpture)
square in outline (Fig. 2(12)). Example:
of the vesicle wall are varied (Fig. 3(2–12)). The
Veryhachium.
processes vary greatly and may be simple or
divided, distally free, coalescent or enclosed in a
2.2.9 Pteromorphs (Pteros = Wing)
peripheral membrane (Fig. 3(1 and 13–17)). The
These have a roughly spherical central zone often
acritarch ornamentation is also variable (Fig. 4)
compressed and surrounded by wing-like lamel-
and in size, too, ranging from a fraction of a µm to
las. Under light microscope, they resemble a
several tens or even a hundred. If the processes on
fried egg (Fig. 2(13)).
an individual vesicle are of one type, they are
However, having said that, the acritarchs show
called homomorphic, but if there are more than one
tremendous variation in size and shape and are
type, the processes are termed as heteromorphic.
generally categorized based on three shared
34
2.4 Excystment Opening
The excystment opening is preserved as a lateral
or transverse split (Figs. 3(1 and 18–23) and 5).
The opening is generally straight or curved with
variably gaping (Fig. 3(1 and 18)) but sometimes
divides the vesicle into two parts (Fig. 3(19)).
Median split (Fig. 3(19)) divides the vesicle into
two more or less equal halves (as in Riculas-
phaera, Hemisphaeridium, and Orthosphaerid-
ium). A split that does not completely encircle
the vesicle is called partial splitting (Loeblich
and Tappan 1969), and is synonymous with
“Cryptosuture” of Lister (1970), “Ornamented
circumferential break” of Reaugh (1978), and
“Ornamented lateral rupture” of Le Hérissé
(1984) (see Fig. 3(18 and 21)). However, Tappan
(1980) treated lateral ruptures as a specific type
of simple rupture, generally observed in elongate
or fusiform acritarchs (Fig. 3(21 and 22)), but
not on those with spherical central bodies. An
Epityche (a specialized variant of the partial
rupture; as in Veryhachium) is a simple, curved
rupture, invisible before partial opening (Fig. 3
(23)). It is always in the form of a flap but
sometimes with a median projection corre-
sponding to an indentation on the opposite edge.
A Pylome (of Sarjeant 1967; = “cyclopyle” of
Eisenack 1969) is an excystment structure that
has a circular or subcircular opening with a
rounded or subangular outline and closed by a
plug-like operculum (or lid), prior to excystment
(Miller 1987) (Fig. 3(20)). As it is preformed, its
position can be detected before excystment, on
the vesicle surface. A pylome may be bordered
by a thickened rim or collar, or by a tubular neck.
Generally, a single pylome is present but very
rarely a second one is present at the opposite pole
(Fig. 3(20)). A cryptopylome is a circular to
subcircular (rarely subangular) opening caused
by the removal of a part of the vesicle wall; it is
invisible before excystment (Tappan 1980).
Pseudopylome, a structure rarely present, is
caused by the thickening of the vesicle wall,
resembling the pylome rim or collar, but typi-
cally occurs opposite to the true pylome (as in
Polyancisrrodoms; Fig. 3(20)); rarely, a true
second pylome is noted as in Asketopalla
2 Acritarchs
formosula and Dicommopalla macadamii.
Endopylome is a circular, operculate opening of
the endophragm, topped by a simple split in the
ectophragm (Le Hérissé 1989).
Unlike sculpture/ornamentation (Fig. 4), the
presence of an excystment (Fig. 5) does not help
in the identification of a species, as there are
members of the same species, with and without
an opening. Varied morphological characters
enumerated above are illustrated through time in
Fig. 6.
3 Classification
The acritarch classification is morphology-based
and depends on vesicle symmetry, shape and
ornamentation as also on the number of wall
layers and the type of opening (Lister 1970;
Tappan 1980; Fensome et al. 1990). Natural
classifications of acritarchs has rarely been
attempted or accepted. The artificial scheme
introduced by Downie et al. (1963) has been
widely accepted or the one proposed by Cramer
(1970) who classified acritarchs in alphabetical
order, excluding those forms with algal affinities
(see Servais 1996 for a detailed discussion).
4 Geological History
and Distribution
The oldest sphaeromorphs are
*1.8-billion-year-old recorded from North
China (Peng et al. 2009; Lamb et al. 2009); the
first acanthomorphs (from Australia, China,
Siberia and USA), appeared shortly afterward in
the Mesoproterozoic. This latter interval is
marked by relatively low diversity, until about
one billion years ago (Agić 2016). Nevertheless,
this period encompassed the initial diversification
of acritarchs as seen in the appearance of
microfossils with various sculpture patterns
(Dictyosphaera and Valeria), protrusions from
the vesicle (as in Germinosphaera) and outer
envelopes (as in Pterospermopsimorpha and
Simia) (see also Fig. 6). Well-preserved micro-
fossils of Tappania and other acritarchs have
4 Geological History and Distribution 35

Fig. 5 Successive appearance of major types of excystment opening (not to scale) (modified after Martin 1993). 1:
Sphaeromorph with pylome and operculum-like structure; 2: Sphaeromorph with equivocal split, either accidental or at
line of cryptosuture; 3: Volkovia with munium; 4: Revinotesta with munitium; 5: Cymatiogalea with pylome and
operculum of definite contour; 6: Corollasphaeridium with gaping opening; 7: Asketopalla with two pylomes; 8:
Polyancistrodorus with pylome and pseudopylome; 9: Veryhachium with epityche; 10: Diexallophasis with preformed
split; 11: Visbysphaera with endopylome; 12: Onondagaella with epibystra

been recorded from the coastal facies of the early
Mesoproterozoic (1.5 Ga) Roper Formation of
north central Australia (Javaux et al. 2001, 2003,
2010). Zhang (1997) described 1.85 Ga-old
sphaeromorphic acritarchs from the Changzhou-
gou Formation in northern China and from the
Meso-Neoproterozoic Ruyang Group (1150–
950 Ma).
The early Neoproterozoic (ca. 900–770 Ma)
witnessed an increase in morphological diversity
(Fig. 7), well-documented from various localities
in Russia and the Arctic Canada. But, because of
36 2 Acritarchs
 4 Geological History and Distribution 37

b Fig. 6 Stratigraphic occurrences of morphological characters (modified after Huntley et al. 2006). Huntley et al. (2006)
noted that the Paleoproterozoic acritarchs typically had spherical vesicles with the occasional medial split (Schizofusa
sinica), enveloping membrane (Pterospermopsimorpha pileiformis), internal bodies (Nucellosphaeridium magnum), or
concentric surface ornamentation (Thecatovalvia annulata and Valvimorpha annulata). The Mesoproterozoic saw the
first appearance of elliptical vesicles (Fabiformis baffinensis), ten process-related characters (Shuiyousphaeridium
macroreticulatum and Tappania plana), vesicle plates (S. macroreticulatum and Dictyosphaera delicata), pores in
vesicle walls (Tasmanites volkovae), and multicelled and colonial appearance (Majasphaeridium sp. and Satka
squamifera). The Neoproterozoic saw new body plan such as polyhedral vesicles (Octoedryxium truncatum),
bulb-shaped vesicles (Sinianella uniplicata), medusoid vesicles (Multifronsphaeridium pelorium), barrel-shaped
vesicles (Artacellularia kellerii), triangular and hair-like processes (Cymatiosphaera wanlongensis and Dasysphaerid-
ium trichotum), funnel-tipped processes (Briareus borealis), processes that fuse at the tips (ectophragm acanthomorph
from Butterfield and Rainbird 1998), and flange ornamentation about the vesicle equator (Simia simica). In the
Cambrian two new morphological characters appeared, a costae meshwork that surrounds the vesicle (Retisphaeridium
brayense) and crest–ornamentation–equatorial ornamentation like flange but that did not circumvent the vesicle,
resulting in wing-like structures (Pterospermella solida)

their low initial diversity and lengthy period of
evolutionary stasis, early eukaryotes were proba-
bly not ecologically significant until the middle or
end of the Neoproterozoic or the Paleozoic, when
the fossil records show increased species diversity
(Fig. 7). The first of these diversifications took
place in the late Tonian period (800–700 Ma),
before the global Snowball Earth glaciation
events, when the envelope-bearing and
ornamented forms became more common.
Thereafter, the Ediacaran witnessed the diversifi-
cation of large, spine-bearing forms such as
Tanarium.
Large acritarchs perished after the Protero-
zoic–Cambrian transition (541 Ma), presumably
because of increased predation pressure from the
newly evolved zooplankton. The acritarchs
diversified parallel to the Cambrian and

Fig. 7 Acritarch assemblages, global acritarch diversity, and stratigraphic ranges of potential index fossils (width of
bars represents relative abundance and/or diversity). Biotic and glacial events are also indicated (modified after Gaucher
and Sprechmann 2009)
38 2 Acritarchs

Ordovician radiations of marine invertebrates dominant eukaryotic phytoplankton, being

(Fig. 7). But by the end of the Devonian
(*360 Ma), the group declined in diversity, with
only basic body plans such as those of leio-
sphaerids still present, eventually leading to a
rapid reduction after the end-Permian mass
extinction, by about 252 Ma (Fig. 7). By the start
of the Mesozoic era (marked by smaller forms,
<20 µm), the acritarchs were no more the
replaced by dinoflagellates and biomineralizing
coccolitophores and diatoms (the “red algal lin-
eages”). The acritarchs were still present
throughout the Mesozoic and the Cenozoic
(Fig. 7), but were not as abundant as before, and
generally include organically preserved unicellu-
lar organisms that have not been identified; these
are known as Problematica. Figures 8 and 9

Fig. 8 Representative acritarchs from Neoproterozoic and Precambrian–Cambrian boundary (not to scale) (modified
after Martin 1993). 1: Pterospermopsimorpha; 2: Vandalosphaeridium; 3: Octoedryxium; 4: Podolina; 6:
Micrhystridium; 5: Comasphaeridium, 7: Rhopalophora; 8: Skiagia. 9: Asteridium tornatum; 10: Annulum
squamaceum; 11: Comasphaeridium strigosum; 12: Comasphaeridium velvetum; 13: Skiagia orbicularis; 14: Skiagia
ornata; and 15: Skiagia scottica
4 Geological History and Distribution 39

Fig. 9 Selected acritarchs in relation to the Cambrian–Ordovician boundary (not to scale) (modified after Martin
1993). 1: Arbusculidium ornatum (= A. ramusculosum); 2: Arbusculidium destombesii (= A. frondiferum); 3: Ladogella
sp.; 4: Actinotodissus achrasi; 5: Actinotodissus sp.; 6: Actinotodissus achrasi; 7: Acanthodiacrodium angustum; 8:
Vulcanisphaera africana; 9: Vulcanisphaera cf. V. africana; 10: Vulcanisphaera imparilis; 11: Buedingiisphaeridium
tremadocum; 12: Buedingiisphaeridium tremadocum; 13: Visbysphaera? occultata; 14: Visbysphaera? fecunda; 15:
Baltisphaeridium crinitum; 16: Dasydiacrodium glabrum; 17: Dasydiacrodium cf. glabrum; 18: Ephelopalla media; 19:
Villosacapsula ceratioides; 20: Villosacapsula globosa; 21: Corollasphaeridium wilcoxianum; 22: Cymatiogalea sp.;
23: Cymatiogalea sp.; 24: Cymatiogalea cuvillieri; 25: Daillydium pentaster; and 26: Multiplicisphaeridium abnorme
40 2 Acritarchs

Fig. 10 Ranges of commonly recorded Cambrian–Devonian acritarch taxa

4 Geological History and Distribution
illustrate representative acritarchs through time.
Figure 10 gives ranges of major commonly
recorded Cambrian–Devonian taxa.
References
Adam, Z. R., Skidmore, M. L., Mogk, D. W., &
Butterfield, N. J. (2017). A Laurentian record of the
earliest fossil eukaryotes. Geology, 45, 387–390.
Agić, H. (2016). Fossil Focus: Acritarchs. Palaeontology
Online, 6, Article 11, 1–13.
Agić, H., Moczydłowska, M., & Yin, L.-M. (2015).
Affinity, life cycle, and intracellular complexity of
organic-walled microfossils from the Mesoproterozoic
of Shanxi, China. Journal of Paleontology, 89, 28–50.
Beraldi-Campesi, H. (2013). Early life on land and the
first terrestrial ecosystems. Ecological Processes, 2,
1–17.
Bernard, S., Benzerara, K., Beyssac, O., Balan, E., &
Brown, G. E., Jr. (2015). Evolution of the macro-
molecular structure of sporopollenin during thermal
degradation. Heliyon, 1(2), e00034.
Brooks, J., & Shaw, G. (1968). Chemical structure of the
exine of pollen walls and a new function for
carotenoids in nature. Nature, 219, 532–533.
Buick, R. (2010). Early life: Ancient acritarchs. Nature,
463(7283), 885–886.
Butterfield, N. J. (2004). A vaucheriacean alga from the
middle Neoproterozoic of Spitsbergen: Implications
for the evolution of Proterozoic eukaryotes and the
Cambrian explosion. Paleobiology, 30, 231–252.
Butterfield, N. J. (2005). Probable proterozoic fungi.
Paleobiology, 31, 165–182.
Butterfield, N. J. (2015). Early evolution of the Eukaryota.
Palaeontology, 58, 5–17.
Butterfield, N. J., & Rainbird, R. H. (1998). Diverse
organic-walled fossils, including “possible dinoflagel-
lates,” from the early Neoproterozoic of arctic Canada.
Geology, 26(11), 963–966.
Colbath, G. K. & Grenfell, H. R. (1995). Review of
biological affinities of Paleozoic acid–resistant,
organic–rich eukaryotic algal microfossils (including
“acritarchs”). Review of Palaeobotany and Palynol-
ogy, 86, 287–314.
Cramer, F. H. (1970). Distribution of selected Silurian
acritarchs. Revista Española de Micropaleontologia.
Numero Extraordinario, 1, 1–203.
Downie, C., Evitt, W. R., & Sarjeant, W. A. S. (1963).
Dinofiagellates, hystrichosphères, and the classifica-
tion of acritarchs. Geological Sciences, 7(3),
6 pp. (Stanford University Publications, California).
Eisenack, A. (1969). Zur Systematik einiger paläozoischer
Hystrichosphären (Acritarcha) des baltischen Gebi-
etes. Neues Jahrbuch für Geologie und Paläontologie
Abhandlungen, 133, 245–266.
41
Eiserhardt, K.-H. (1989). Baltisphären aus Gotländer
Ojlemyrflint (Acritarcha, Oberordoviz, Geschiebe,
Schweden). Mitteilungen aus dem
Geologisch-Paläontologischen Institut der Universität
Hamburg H. 58, 79–129.
Evitt, W. R. (1963). A discussion and proposals concern-
ing fossil dinoflagellates, hystrichospheres and acri-
tarchs. Proceedings of the National Academy of
Sciences, 49, 158–164.
Fensome, R. A., Williams, G. L., Barss, M. S., Freeman,
J. M., & Hill, J. M. (1990). Acritarchs and fossil
prasinophytes: An index to genera, species and
infraspecific taxa. In American Association of Strati-
graphic Palynologists Contributions Series, 25, 771 p.
Gaucher, C., & Sprechmann, P. (2009). Neoproterozoic
acritarch evolution. In C. Gaucher, A. N. Sial, G.
P. Halverson, & H. E. Frimmel (Eds.),
Neoproterozoic-Cambrian Tectonics, Global Change
and Evolution: A focus on southwestern Gondwana.
Developments in Precambrian Geology (Vol. 16,
pp. 319–326). Elsevier.
Huntley, J. W., Xiao, S., & Kowalewski, M. (2006). 1.3
billion years of acritarch history: An empirical mor-
phospace approach. Precambrian Research, 144, 52–68.
Javaux, E. J. (2014). Acritarchs. In Encyclopedia of
astrobiology. https://doi.org/10.1007/978-3-642-
27833-4_23-3.
Javaux, E. J., & Knoll, A. H. (2016). Micropaleontology
of the lower mesoproterozoic roper group, Australia,
and implications for early eukaryotic evolution. Jour-
nal of Palaeontology, 91(2), 199–229.
Javaux, E. J., & Knoll, A. H. (2017). Micropaleontology
of the lower Mesoproterozoic Roper Group, Australia,
and implications for early eukaryotic evolution. Jour-
nal of Paleontology, 91(2), 1–31.
Javaux, E. J., Knoll, A. H., & Walter, M. R. (2001).
Morphological and ecological complexity in early
eukaryotic ecosystems. Nature, 412, 66–69.
Javaux, E. J., Knoll, A. H., & Walter, M. R. (2003).
Recognizing and interpreting the fossils of early
eukaryotes. Origins of Life and Evolution of the
Biosphere, 33(1), 75–94.
Javaux, E. J., Marshall, C. P., & Bekker, A. (2010).
Organic-walled microfossils in 3.2-billion-year-old
shallow-marine siliciclastic deposits. Nature, 463
(7283), 934–938.
Knoll, A. H. (1994). Proterozoic and Early Cambrian
protists: Evidence for accelerating evolutionary tempo.
Proceedings of the National Academy of Sciences of
the United States of America, 91, 6743–6750.
Knoll, A. H. (2006). Life on a young planet: The first
three billion years of evolution on earth (p. 296).
Princeton University Press.
Lamb, D. M., Awramik, S. M., Chapman, D. J., & Zhu, S.
(2009). Evidence for eukaryotic diversification in the
similar to 1800 million-year-old Changzhougou For-
mation, North China. Precambrian Research, 173,
93–104.
42
Lazcano, A., & Miller, S. L. (1994). How long did it take
for life to begin and evolve to cyanobacteria? Journal
of Molecular Evolution, 39, 549–554.
Le Hérissé, A. (1984). Upper Ordovician lower Silurian
organic walled microphytoplankton from the Nar
Borehole, Gotland, Sweden. In J. Utting (Ed.),
Abstract of the 6th International Palynology Confer-
ence (p. 87). Calgary.
Le Hérissé, A. (1989). Acritarches et kystes d’algues
Prasinophycées du Silurien du Gotland, Suède.
Palaeontographica Italica, 71, 298 pp.
Lister, T. R. (1970). The acritarchs and chitinozoa from
the Wenlock and Ludlow Series of the Ludlow and
Millichope areas, Shropshire. Part 1. Palaeontograph-
ical Society Monographs, 124(1), 100.
Martin, F. (1993). Acritarchs: A review. Biological
Reviews, 68, 475–538.
Mendelson, C. V. (1987). Acritarchs. In J. Lipps (Ed.),
Fossil prokaryotes and protists (pp. 62–86). Knox-
ville: University of Tennessee.
Miller, M. A. (1987). A diagnostic excystment suture in
the Silurian Acritarch Circinatisphaera aenigma gen.
et sp. nov. Palynology, 11, 97–105.
Molyneux, S. G. (1999). A reassessment of Manx Group
acritarchs, Isle of Man. 23–32. In N. H. Woodcock, D.
G. Quirk, W. R. Fitches, & R. P. Barnes (Eds.), In sight
of the suture: The Palaeozoic geology of the Isle of
Man in its Iapetus Ocean context (Vol. 160, 376 pp).
London: Geological Society, Special Publications.
Montenari, M., Servais, T., & Paris, F. (2000). Palyno-
logical dating (acritarchs and chitinozoans) of Lower
Palaeozoic phyllites from the Black
Forest/southwestern Germany. Comptes Rendus de
l’Académie des Sciences de Paris, Sciences de la
Terre et des Planétes, 330, 493–499.
Munnecke, A., Delabroye, A., Servais, T., Vanden-
broucke, T. R. A., & Marco Vecoli, M. (2012).
Systematic occurrences of malformed (teratological)
acritarchs in the run-up of Early Palaeozoic d13C
isotope excursions. Palaeogeography, Palaeoclima-
tology, Palaeoecology, 367–368, 137–146.
Nagovitsin, K. (2009). Tappania-bearing association of
the Siberian platform: Biodiversity, stratigraphic posi-
tion and geochronological constraints. Precambrian
Research 173, 137–145.
Ohmoto, H. (2004). Archean atmosphere, hydrosphere,
and biosphere. In P. G. Eriksson, W. Altermann, D.
R. Nelson, W. U. Mueller, & O. Catuneanu (Eds.), The
Precambrian Earth: Tempos and events. Develop-
ments in Precambrian geology (Vol. 12, pp. 361–368).
Pang, K., Tang, Q., Schiffbauer, J. D., Yao, J., Yuan, X.,
Wan, B., et al. (2013). The nature and origin of
2 Acritarchs
nucleus-like intracellular inclusions in Paleoprotero-
zoic eukaryote microfossils. Geobiology, 11, 499–510.
Peng, Y., Bao, H., & Yuan, X. (2009). New morpholog-
ical observations for Paleoproterozoic acritarchs from
the Chuanlinggou Formation, North China. Precam-
brian Research, 168, 223–232.
Reaugh, A. B. (1978). A new excystment mechanism in
the Silurian acritarchs Diexallophasis of Virginia.
Palaeontology, 21, 869–872.
Sarjeant, W. A. S. (1967). Observations on the acritarch
genus Micrhystridium (Deflandre). Revue de
Micropaléontologie, 9, 201–208.
Servais, T. (1996). Some considerations on acritarch
classification. Review of Palaeobotany and Palynol-
ogy, 93, 9–22.
Servais, T., Brocke, R., Fatka, O., Le Hérissé, A., &
Molyneux, S. G., (1997). Value and meaning of the
term acritarch. In O. Fatka, & T. Servais, (Eds.),
Acritarcha in Praha 1996. Acta Universitatis Caroli-
nae, Geologica, 40, 631–643.
Tappan, H. (1980). The paleobiology of plant profits
(pp. 1–1028). San Francisco, CA: Freeman.
Vidal, G., & Moczydlowska-Vidal, M. (1997). Biodiver-
sity, speciation, and extinction trends of Proterozoic
and Cambrian phytoplankton. Paleobiology, 23, 230–
246.
Wang, W., Servais, T., Yan, K., Vecoli, M., & Li,
J. (2014). The Ordovician acritarch Dactylofusa
velifera Cocchio 1982: A biostratigraphical and
palaeogeographical index species. Palynology, 39,
125–141.
Yan, Y. (1991). Shale-facies microflora from the
Changzhougou Formation (Changcheng System) in
Pangjiapu Region, Hebei, China. Acta Micropalaeon-
tologica Sinica, 8, 183–195.
Yin, L. M. (1997). Acanthomorphic acritarchs from
meso-neoproterozoic shales of the Ruyang Group,
Shanxi, China. Review of Palaeobotany and Palynol-
ogy, 98, 15–25.
Yin, L.-M., Xunlai, Y., Fanwei, M., & Jie, H. (2005).
Protists of the upper Mesoproterozoic Ruyang Group
in Shanxi Province, China. Precambrian Research,
141, 49–66.
Zhang, Z. (1986). Clastic facies microfossils from the
Chuanlinggou Formation (1800 Ma) near Jixian,
North China. Journal of Micropalaeontology, 5, 9–16.
Zhang, Z. (1997). A new Palaeoproterozoic clastic-facies
microbiota from the Changzhougou Formation,
Changcheng Group, Jixian, north China. Geological
Magazine, 134, 145–150.
 Pollen and Spores
3

awfully complex best noted by Traverse (2007,

1 Introduction
pp. 95–96), “They are very variable as to size, and
even as to morphological type in a single spo-
Fungi, algae, non-vascular plants, and vascular
rangium, or anther; in exceptional cases, monolete
non-seed-bearing plants present sporangia where
and trilete spores are found in one sporangium, or
the spores are produced, and spermatophytes
triporate, tetraporate, and pentaporate pollen in
(Pteridospermatophyta, gymnosperms, and angio-
one another, or (very commonly) pollen of very
sperms) produce pollens (see also Table 1). Spore
different size in one species.” For more details on
is a haploid cell derived from sporangium via
morphology, the readers are referred to excellent
meiosis, whereas pollen is an immature, endo-
works by Punt et al. (2007), Traverse (2007),
sporic male gametophyte derived from male
Hesse et al. (2009), and Halbritter et al. (2018).
spores (microspores) in seed plants. Due to their
(a) resistant wall (resistant to microbial attack and
to the effects of burial pressure and temperature),
(b) production in large numbers, (c) microscopic
2 Spore Morphology
nature, that enables them to travel large distances
The cryptospores (dispersed spores/spores pro-
by wind or water (and also their adaptations such
duced by embryophytes; although recent data
as Anemophily, Entomophily, or Hydrophily),
suggests that not all are produced by the latter),
and (d) the fact that since the last one billion years,
are the earliest evidence of the colonization of the
the palynomorphs of one group or another are
Earth by land plants. The embryophytes (Bryo-
always present in the sedimentary rocks of suitable
phyta, Pteridophyta, and seed plants) are plants
lithology, make them a valuable tool for bios-
with true embryos in their life cycle. The earliest
tratigraphic (particularly for the correlation of
record (of an assemblage of monads and tetrads)
continental and near-shore marine post-Silurian
comes from the Early Ordovician (*473–
strata), paleoecological, and paleoenvironmental
471 Ma) of Argentina (see Rubinstein et al.
studies (Traverse 2007). Here, it must be men-
2010). These are thought to be liverworts and
tioned that unlike all other groups dealt in this
predate the first vascular plants by almost 8–
book, these are not marine organisms but an
12 Ma (see Steemans et al. 2000). Previously,
allochthonous group that are products of conti-
the earliest hilate and trilete spores came from the
nental vegetation.
late Darriwilian age (Middle Ordovician; *463–
The chapter deals with pollen and tracheo-
461 Ma) of Saudi Arabia (Steemans et al. 2009)
phytes. Pollens form the bulk of the chapter and
and from the Czech Republic (Vavrdová 1990;
only major morphological characteristics are
Strother et al. 1996).
briefly dealt and illustrated as their terminology is

© Springer Nature India Private Limited 2020 43

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_3
44
Table 1 Differences between Pollen and Spore. The terms mentioned are detailed
Pollen
It is a container that houses male gametophyte of
phanerogams (i.e., seed-bearing plants; spermatophyte).
Reproduce by forming seeds. Examples:
Pteridospermatophyta, gymnosperms, and angiosperms
Aperturate pollen grains are colpate/porate, etc.
Apertures occur on the distal pole
Perine is absent in phanerogams (Perine is a loosely
organized sporopolleninous envelope covering the exine)
Division of exine into sexine and nexine is prominent
The trilete spore is the most basic spore type
and is characterized by a Y-shaped “mark” or
“suture” or “scar” (the Y-mark or trilete mark)
(Fig. 1(1)). These marks are called laesura (plu-
ral: laesurae; Fig. 1(1 and 3)) and are in the form
of raised ridges or fissures at the surface on the
proximal face of a spore (Fig. 1(2)). The laesurae
are formed by four equivalent reproductive cells
(spore tetrad or tetrahedral tetrads) that are pro-
duced by the spore mother cell by simultaneous
meiosis (Fig. 2); all four are in contact at a single
point forming the Y-mark or the trilete mark (as
in Dennstaedtiaceae). The three laesurae radiate
120° from the proximal pole (Fig. 1(1 and 3))
and thus, display radial symmetry; such spores
are heteropolar, i.e., with differently formed polar
faces. A laesura constitutes a Commissure (a slit
or line of dehiscence in the laesurae) and is
bordered by a Margo, a distinct thinning or
thickening of the ektexine (sometimes also
referred to as ectexine) bordering the aperture
(Fig. 1(4)). The equatorial contour is called the
amb (=the spore or pollen outline; see Fig. 1(3))
which displays varied shapes (see Fig. 3).
3 Pollen Morphology
The pollen morphology is very complex and
exhaustive (see Traverse 2007). It is briefly
explained under the following subheads: Pollen
Units, Shape, Symmetry, Size, Polarity,
3 Pollen and Spores
Spore
It is the resting spore and dispersal phase of
non-vascular plants and vascular non-seed-bearing
plants
Aperturate spores are monolete or trilete
Apertures occur on the proximal pole
Perine is present in the spores of Bryophyte and
pteridophyte
Exine and intine are absent
Apertures, Sporoderm stratification, and exine
ornamentation. The descriptive terms used below
are modified from several sources, including
Potonié (1934), Erdtman (1947, 1952), Harris
(1955), Potonié and Kremp (1955), Kremp (1968),
Grebe (1971), Punt et al. (1994, 2007), Traverse
(2007), and Halbritter et al. (2018).
3.1 Pollen Units
The pollen grains divide meiotically to form a
tetrad (Fig. 2) that dissociates at maturity into a
single grain called monad (Fig. 4(1)), into two,
dyad (Fig. 4(2)), four, tetrad (Figs. 4(3) and 5
(9)), eight, octad or many, polyad (Fig. 4(10)).
The tetrads (Figs. 4(3) and 5(9)) are formed
when four pollen grains are united and are the
unseparated product of meiosis (as in Acaci-
apollenites). Based on the arrangement of grains,
they are of the following types—tetrahedral tet-
rad (Fig. 4(3)), tetragonal tetrad (Fig. 4(4)),
rhomboidal tetrad (Fig. 4(5)), decussate tetrad
(Fig. 4(6)), T-shaped tetrad (Fig. 4(7)), linear
tetrad (Fig. 4(9)), and cryptotetrad or pseu-
domonad (Fig. 4(9)). In polyads (Fig. 4(10)), as
in Mimosaceae, the tetrad cell divides once or
twice or more, yielding a group of 8–64 cells that
remain together after maturity. In pollinia, the
whole contents of an anther are released as one
unit as in Orchidaceae (Fig. 4(11)).
3 Pollen Morphology 45

Fig. 1 Morphology of a
trilete spore (modified from
Erdtman 1943, 1952; Punt
et al. 2007; Halbritter et al.
2018). Spore can be
homosporous or
heterosporous. In club moss
(a primitive vascular plant), it
is homosporous (i.e., a single
type of spore is formed,
isospores), whereas the
heterosporous plants produce
male microspores and female
megaspores. See text for
further explanation

3.2 Polarity
Polarity is defined as the condition of having
distinct poles in a pollen grain and best detected
at the tetrad stage, i.e., prior to separation
(Fig. 3). The orientation of polarity is an
important defining character in species identifi-
cation and description and it is the polarity that
distinguishes a spore from pollen. To understand
polarity, it is important to first understand how a
grain is viewed (Fig. 5(1–3)). The polar and
distal views have varying focal planes; the darker
plane represents the equatorial focal plane (Fig. 5
(2 and 3)) as detailed in Fig. 5(4–9) by examples
of monolete (Fig. 5(4 and 5)) and trilete spores
(Fig. 5(6 and 7)) and monosulcate pollen grains
(Fig. 5(8 and 9)).
The spores have two poles, proximal and distal
(Fig. 5(4 and 6)). The proximal face/surface is the
one that faces inward/nearest or toward the center
of the tetrad (Fig. 5(6)). The distal face/surface
faces distal, i.e., away from the center of tetrad
and opposite the proximal part (Fig. 5(6)). An
imaginary straight line called the polar axis con-
nects the proximal pole to the distal pole (Fig. 5(4
and 6)). The equatorial axis/equatorial diameter
(=equator) is also an imaginary straight line that
occurs perpendicular to the polar axis and runs
through the middle of the pollen/spore (spore:
Fig. 5(4 and 6); pollen: Fig. 5(8)). The term
meridional (or longitudinal) is used to indicate the
surface of a pollen grain or spore that is oriented in
the pole-to-pole direction, and at right angles to
the equator (Fig. 5(4, 6, and 8)).
46 3 Pollen and Spores

Fig. 2 Spore mother cell. All

four reproductive cells (spore
tetrad or tetrahedral tetrads)
are produced by the spore
mother cell by simultaneous
meiosis. These four grains are
in contact with each other at a
single point, thus forming the
Y-mark or the trilete mark

Fig. 3 Amb shapes (spore

outline) (modified from
Erdtman 1943, 1952; Punt
et al. 2007). The equatorial
contour of a spore is called an
Amb, which displays varied
shapes
3 Pollen Morphology 47

Fig. 4 Pollen units (modified from Erdtman 1943, 1952; Punt et al. 2007). The pollen grains divide meiotically to form
a tetrad that dissociates at maturity into a single grain or many. See text for further explanation
48 3 Pollen and Spores

Fig. 5 Image generation (1–3), polarity (4–9) and views (10–17) (modified from Erdtman 1943, 1952; Punt et al.
2007; Halbritter et al. 2018). See text for further explanation

In the polar view (Fig. 5(10–13)), the polar
axis is directed toward the observer and the
aperture is viewed centrally and the zonal aper-
ture is then viewed in the circumference of the
pollen grain. Whereas, in the equatorial view
(Fig. 5(14–17)), the aperture is arranged merid-
ionally (i.e., pole to pole), and at right angles to
the equator. In this position, both the polar axis
and the equatorial diameter can be viewed and
measured.
3 Pollen Morphology 49

Fig. 6 Types of pollen grains based on polarity (1–9) and various axes discussed in the text (10) (modified from
Erdtman 1943, 1952; Punt et al. 2007). See text for further explanation
50 3 Pollen and Spores

Fig. 7 Simple (1–25) and compound aperture types (25–60) (modified from Erdtman 1943, 1952; Punt et al. 2007;
Halbritter et al. 2018). See text for further explanation
3 Pollen Morphology 51

Fig. 8 Pollen shapes (modified from Erdtman 1952). The pollen and spores are described by the shape of their outline,
either as non-angular or angular. See text for further explanation

A pollen grain can be either polar or apolar
(Fig. 6). Polar pollen grains can be isopolar or
heteropolar, depending upon the demarcation
between the two equal or unequal polar faces
(Fig. 6(1)). In isopolar grains (Fig. 6(2–4)), the
distal and proximal faces (above and below the
equatorial plane) are mirror images (as in Calluna,
Ericaceae), whereas in heteropolar grains (Fig. 6
(5 and 6)), the two faces are distinctly different,
either in shape, ornamentation or in the apertural
52
system (as in Irlbachia, Gentianaceae). Thus, one
face may have an opening (aperture) and the other
would be marked by its absence (as in Lilium and
Elaeis). The paraisopolar or subisopolar pollen
grains (Fig. 6(7 and 8); these are part of the polar
grain category and intermediate between isopolar
and heteropolar grains) are those that show minor
differences between the distal and proximal faces
such that one face (distal) is convex and the other
is concave or plane (proximal) or vice versa (as in
Diastella, Nivenia, and Ulmas).
In apolar pollen grains, the poles and regions
(proximal pole, distal pole, proximal region and
distal region) cannot be distinguished in indi-
vidual spores (monads), after the separation from
the tetrad. Such grains occur as linear tetrads and
exhibit no polarity such as in Periploca
ephedriformis and Typha.
The cryptopolar type (Fig. 6(9)) is noted in
some bryophyte spores (as in Calobryum denta-
tum) where the grain lacks the tetrad mark and
has different sculpturing on distal and proximal
faces. They appear to be apolar, i.e., poles and
polar region cannot be distinguished, but on
closer examination they do reveal a more or less
distinct polarity (see also Erdtman 1952).
A summary of varied axes and poles mentioned
above is given in Fig. 6(10).
The polarity of a pollen grain forms the basis
of the aperture terminology such that a circular
aperture that occurs equatorially or globally is
called porus. The circular aperture that occurs
distally is termed as Ulcus. An elongated aper-
ture that occurs equatorially or globally is called
colpus (furrow is a common word for colpus).
When an elongated aperture occurs distally it is
called sulcus (detailed below; see Fig. 7).
3.3 Apertures
An aperture (also treme—a Greek word for
aperture) is an opening or thinning of the exine
(outer layer); the intine (inner layer) is generally
not affected and remains thick (Sporoderm
stratification is detailed later in the chapter). In
3 Pollen and Spores
simple terms, it is a specialized region of the
sporoderm that is thinner than the remaining
sporoderm and differs in ornamentation and/or in
structure. Based on the position of this opening,
apertures can be grouped as polar, global, or
equatorial. Polar types are either monopolar
(positioned either in the proximal or distal pole)
or bipolar (both in the proximal and distal face).
The global apertures are uniformly distributed
over the surface, whereas the equatorial ones are
meridionally arranged. In some taxa, as in Canna
spp. of Cannaceae, there are no special apertures
and are thus called inaperturate or non-aperturate.
But, in reality, most of them are actually “om-
niaperturate,” i.e., the thin exine covers the grain
with a thick intine (or at least as thick as the
exine). The aperturate ones (i.e., those with
aperture), are of two types, either containing
pores (Porus, p1. Pori) or furrows (Colpus, p1.
Colpi or Sulcus, p1. Sulci) (see Fig. 7). The
apertures can be simple (Fig. 7(1–25)) or com-
plex (Fig. 7(26–60)). The simple ones have only
one type of aperture (i.e., monoaperturate) (see
Erdtman 1943, 1952) and include the following:
3.3.1 Lete
The aperture is slit-like and is situated at the
proximal end (as in the spores of pteridophytes).
Monolete (Fig. 7(1)) are with one slit (as in
Polypodium), and trilete (Fig. 7(2 and 3)) is with
a triradiate slit (as in Lycopodium).
3.3.2 Porus
This is an equatorially placed circular aperture
(see Fig. 7(4)) with a length/breadth ratio of ˂2;
grains with porus apertures are called Porate (as
in Urticaceae).
3.3.3 Periporus
A global-type circular aperture with a length/
breadth ratio of ˂2. They are called Periporate or
Pantoporate (as in Amaranthaceae and Mal-
vaceae) (see Fig. 7(5 and 39)). In the Periporate
type, the pores are scattered all over the pollen
grain surface, i.e., arranged other than on the
equator (Fig. 7(5)).
3 Pollen Morphology
3.3.4 Ulcrus
This is a simple circular aperture (Fig. 7(6 and
7)) with a length/breadth ratio of ˂2 (as in Typha,
Typhaceae). If placed distally, they are called
non-ulcerate and if proximally, then ulcerate.
3.3.5 Sulcus
This is a distally placed simple elongated
boat-shaped aperture, with tapering ends and a
length/breadth ratio ˃2 (Fig. 7(8 and 9)). The
grains with sulcus aperture are called sulcate (as
in Arecaceae).
3.3.6 Sulculus/Sulculi
These are sulcoid apertures, parallel to the equator
and often situated between the equator and the
distal pole (Fig. 7(10 and 11)) (as in Eupomatia).
3.3.7 Colpus
This is a meridionally placed simple, long
furrow-like aperture with a length/breadth ratio
˃2, facing the pole-to-pole direction (Fig. 7(12–
14)); they are called colpate, as in Lamiaceae.
3.3.8 Pericolpus
These are the global types with simple, long
furrow-like aperture with a length/breadth ratio
˃2. The pollen grains of this apertural type are
called Pantocolpate/Pericolpate (Fig. 7(47)), as
in Portulacaceae.
In addition, simple apertures are also noted in
angiosperms (Fig. 7(17–25)). These are:
3.3.9 Monoporate
Pollen grains provided with a single pore (Fig. 7
(16)).
3.3.10 Syncolpate
The colpi join at the polar region only (Fig. 7(17
and 18)) as in Bauhinia.
3.3.11 Parasyncolpate
In the polar region, colpi are bifurcated and the
adjacent branches meet with each other, leaving
an isolated apocolpial field of regular shape
(generally triangular) (Fig. 7(19 and 20)) as in
Nymphoides peltata (Menyanthaceae).
53
3.3.12 Monosulcate
These have a single germinal furrow or colpus or
sulcus (Fig. 7(21)).
3.3.13 Trichotomosulcate
These are simple, distally placed, trifurcated (the
sulcus is more or less triangular often simulating
a trilete laesurae) and as long as broad (Fig. 7(22
and 23)) as in Elaeis gidneensis (Arecaceae).
3.3.14 Spiraperturate
These spiral apertures are irregularly arranged
(Fig. 7(24 and 25)) as in Thunbergia
(Acanthaceae).
Composite apertures are of two types: colpo-
rate (Ora in Colpi) and pororate (Ora in Pore)
(Fig. 7(26–28)) and occur in varied combina-
tions, of the two (Fig. 7(29–57)). These are:
3.3.15 Colporus
These are meridional placed composite aperture
(Ora in Colpi) with ectoapertures (colpi) with a
length/breadth ratio >2. They are called colporate
(Fig. 7(26 and 27)), as in Asteraceae, Fabaceae,
and Solanaceae.
3.3.16 Pororus
These are meridional placed composite aperture
(Ora in Pore) with ectoaperture (pore) with a
length/breadth ratio >2. They are called pororate
and possess incongruent ectoaperture and
endaperture (Fig. 7(28)) as in Betula and
Casuarina. Congruent ecto and endospores are
called porate.
3.3.17 Tricolpate/Tricolporate/
Triporate
Three ectocolpi, three compound apertures or
three pores, respectively (Fig. 7(35–40)). Tri-
colpate (Fig. 7(35 and 36)) have three merid-
ionally placed colpi without pores, transverse
furrows or other such modifications. Tricolporate
(Figs. 7(37), 8, 9, 10, and 11(38)) have three
colpi with pores or other usually equatorial
modifications. The Triporate have three equato-
rial more or less isodiametric germinal apertures
(Fig. 7(39 and 40)).
54 3 Pollen and Spores

Fig. 9 Pollen symmetry. The pollen grains or spores are either symmetric (1–4) or asymmetric (5–9) (modified from
Erdtman 1943, 1952; Punt et al. 2007). See text for further explanation

3.3.18 Zonocolporate 3.3.20 Heterocolpate

(=Stephanocolporate)
These have more than three colpi, meridionally
arranged and provided with pores (Fig. 7(43 and
44)).
3.3.19 Zonoporate (=Stephanoporate)
These have more than three equatorial, equally
spaced pores on the equator (Fig. 7(45 and 46)).
Pollen grains with other intermediate types are
also noted, and they include:
These have both simple and compound colpi as in
Lythrum, Peplis (Lythraceae) (Fig. 7(50 and 51)).
3.3.21 Synorate or Svnclinorate
These have lalongate ora, anastoming latitudi-
nally (Fig. 7(52)), as in Solanum.
3.3.22 Multi Orate
These have two or more ora found latitudinally,
along the length of the colpus (Fig. 11(53)), as in
Viticipremna.
3 Pollen Morphology
Fig. 10 Spore size (modified
from Chaloner and Sheerin
1981; Traverse 2007).
0.1 mm = 100 lm. See text
for further explanation
3.3.23 Porocolpate
These have apertures in which colpi alternate
with pores around the equator (Fig. 7(54 and
55)), as in Pardoglossum (Boraginaceae).
3.3.24 Colporoidate
These have well-developed colpi but weakly
developed ora (=Oroids) (Fig. 11(56)) as in
Phalline lucida.
55
3.3.25 Colpoidorate
These have weakly developed colpi (=Colpoids)
but well-developed ora (Fig. 7(57)), as in Alan-
gium villosum.
3.3.26 Lophate/Fenestrate
These have window-like pattern of ridges
(=lophae) formed by an outer exine surrounding
window-like spaces or depressions. The fenestrate
pollen is characterized by the lack of tectum
(Fig. 7(58–60)).
56 3 Pollen and Spores

Fig. 11 Sporoderm stratification (1–5) and sculpturing, i.e., exine ornamentation (6–9) (modified from Erdtman 1969;
Faegri and Iversen 1964, 1975, 1989; Punt et al. 2007; Traverse 2007). The sporoderm (pollen wall) is stratified and
consists of two layers, the outer exine and the inner intine and with further subdivisions of these two. Perine is often
used in conjunction with sexine and nexine and often cannot be differentiated from sexine in the fossil material. See text
for further explanation

3.4 Shape
Shape (Fig. 8) is also a diagnostic character for
species identification. But it varies greatly, from
species to species, to within one grain type and
even within one species. Pollen and spores are
described by the shape of their outline, either as
non-angular or angular (Fig. 8). The outline or
circumference in polar view is called Amb. Based
on the ratio of polar axis (PA) and equatorial
diameter (ED) (Fig. 8, inset), eight shape classes
are noted (see Erdtman 1952). In the equatorial
view, the ratio between the PA and ED, multi-
plied by 100 gives the indication of the shape
such as prolate, prolate–spheroidal, spheroidal,
sub-prolate, perprolate, oblate, oblate–spheroidal,
sub-oblate, and peroblate (see Fig. 8).
3.5 Symmetry
Symmetry with relation to a pollen grain implies
similarity of halves on either side of a median line
or plane so far as aperture and ornamentation,
3 Pollen Morphology
etc., are concerned. The pollen grains or spores
can be either symmetric or asymmetric (Fig. 9).
In asymmetric ones, there is no plane of sym-
metry (as in Berberis darwinii, Papaver arge-
mone, and Myriophyllum alterniflorum). These
may be with fixed shape, called fixiform or
without any definite shape called non-fixiform.
Asymmetric pollen and spores are very rare.
The symmetric ones can be either radially
symmetrical (radiosymmetric; Fig. 9(1–4)) or
bilaterally symmetrical, i.e., having a single
plane of symmetry (Fig. 9(5–9)). In radially
symmetrical grains, any plane including the polar
axis, produces identical halves (Fig. 9(1–4)).
Hence, such grains possess more than two ver-
tical planes of symmetry (as in Centaurea); one
horizontal and two or more vertical. All the
planes are of equal length. In radially symmet-
rical heteropolar spores, there is no horizontal
plane of symmetry, but two or more vertical
planes of symmetry (as in Osmunda regalis,
Ophioglossum vulgatum, and Pteridium aquil-
inum) (Fig. 9(4)). Most pollen and spores are
symmetrical.
The bilateral isopolar spores (Fig. 9(7 and 8))
exhibit two vertical planes and one horizontal
plane of symmetry. All the planes are not of
equal length (as in Rungia grandis). In bilateral
heteropolar spores (Fig. 9(9)), there exist two
vertical planes of symmetry. The planes are
unequal in length and intersect each other at right
angles (as in Ephemerum serratum, Picea abies,
and Cycas revoluta).
3.6 Size
Size, like shape, varies greatly (Fig. 10). The
smallest is about 5  2.4 µm (Myosotis palus-
tris) whereas some are large, >200 µm in diam-
eter, as in Boraginaceae, Curcurbitaceae,
Nyctaginaceae, and Orectanthe ptaritepuiane
(Abolbodaceae). Microspore is a term used to
include all spores <200 lm in diameter (Fig. 10).
The wind pollinated and airborne pollen grains
normally range from 10 to 80 lm. It is impera-
tive to also measure exine elements (discussed in
57
the next section) by considering the thickness of
exine, sexine/nexine thickness ratio and the
thickness of the exine projections >0.5 µm, if
any. The different pollen size classes, based on
the size expressed as the length of the longest
axis, are given in Fig. 10 (see also Erdtman
1945).
4 Sporoderm Stratification
The Sporoderm (pollen wall) in mature pollen
(Fig. 11(1)) is stratified and consists of two lay-
ers, the outer exine and the inner intine (Fig. 11
(2–5)). Perine is the outermost layer of the
sporoderm that is mainly present in spores of
ferns and mosses in contact with the exine
(Fig. 11(3)), and is not always acetolysis resis-
tant. Perine is often used in conjunction with
sexine and nexine and often cannot be differen-
tiated from sexine in the fossil material (see
Erdtman 1943; Punt et al. 2007).
Exine is the outer layer of the wall of a
palynomorph (Fig. 11(4 and 5)). It is composed
largely of sporopollenin, a chemically inert bio-
logical polymer. Exine covers the entire pollen
surface except the apertures where it is either
absent or is very much reduced. Exine is divided
into two layers, an outer sculptured sexine and an
inner unsculptured nexine (Fig. 11(4 and 5)).
Sexine, in turn, is also made up of two layers, the
outer, ectosexine and the inner, endosexine
(Fig. 11(5)). Endosexine consists of columella,
baculum, and void spaces separating the bacula
from each other (Fig. 11(4)). The columella (pl.
columellae, adj. columellate) is a column-like
element of sexine that supports the tectum at the
top. A baculum is also a sexine element, which is
a slim, cylindrical rod-like structure and stands
on the nexine. Each baculum is more than 1 lm
in length and less than this, in diameter. Bacula
are straight and often end abruptly (Fig. 11(4)).
The sexine is made up of a set of radially directed
rods supporting a roof-like structure on the
periphery of a pollen grain called tectum or
tegillum, which may be partially perforated or
completely absent (Fig. 11(4)); when absent, it is
58
called atectate or intectate (Fig. 13(1)). The
rod-like elements of the ectexine/sexine sup-
porting a tectum are called columella/columellae
(Fig. 11(4 and 5)).
Intine is the innermost layer bordering the
surface of the cytoplasm and underlies the exine
(Fritzsche 1837). It is soft, fragile, and pliable
and remains tightly attached to the plasma
membrane (Fig. 11(2, 4 and 5)). It is composed
of cellulose and pectates and is normally absent
in fossil sporomorphs. Intine is not acetolysis
resistant, and hence, is often absent in conven-
tionally prepared fossil palynological material.
The thickness of intine is related to that of exine.
In the place where the exine gets thinner, the
layer of intine becomes thicker. Hence, thickened
portions of intine occur under pores.
4.1 Exine Ornamentation
There are two different types of exine ornamen-
tation, sculpturing (Fig. 11(6–9)) and structure or
texture (Fig. 13). The structure comprises of all
the internal (infratectal) baculae (singular = bac-
ulum; see Fig. 11(4)) with various form and
arrangements (Fig. 12(1–26)). A baculum is a
free-standing element of sculpturing, but a col-
umella is part of the structure (Punt et al. 2007)
(see Fig. 11(4)). All ektexine (also referred to as
Ectexine) (Fig. 11(4)) characters are structural
features, whereas sculpturing comprises all
external (supratectal) geometric features (Fig. 12)
without reference to their internal construction.
Tectum may be smooth i.e., psilate (see Fig. 12
(1)) or with processes of various kind like spin-
ules, spines, pila, verruca, gemma, clava, gran-
ules, etc. (see Fig. 12(1)). Tectum provided with
the processes are referred to their respective ter-
minology (see Fig. 12(1)). For example, tectum
with spinules is referred to as spinulate, with
spines called echinate, with pila called pilate, with
verruca called verrucate, with gemma called
gemmate, with clava called clavate, with granules
called granulate, etc. (see Fig. 12(1)).
The exine ornamentation can be diagnostic of
specific pollen types. Some of these are briefly
enumerated below and illustrated in Fig. 12.
3 Pollen and Spores
4.1.1 Fossulate
These are elongated, irregular grooves on the
surface (Fig. 12(1)).
4.1.2 Clavate
These are club-shaped elements higher than
1 lm (Fig. 12(1)).
4.1.3 Perforate
These are holes <1 lm and generally situated in
the tectum (Fig. 12(1)).
4.1.4 Psilate/Laevigate
These have smooth surface (Fig. 12(4 and 5)).
4.1.5 Gemmate
These are constricted at its base, higher than
1 lm, with same width and height; a sculpture
with more or less spherical projections (Fig. 12(6
and 7)).
4.1.6 Scabrate
Ornamentation of any shape but <1 lm in all
directions, giving it a rough appearance. Ele-
ments >1 lm are described according to their
shape, such as baculate, verrucate, and clavate.
(see below) (Fig. 12(8 and 9)).
4.1.7 Baculate
These are cylindrical, free-standing exine ele-
ments >1 lm in length; height is greater than the
basal diameter (Fig. 12(10 and 11)).
4.1.8 Verrucate
These are wart-like knobs or elevations >1 lm
wide, broader than its height and not constricted
at the base (Fig. 12(12 and 13)).
4.1.9 Clavate
These are club-shaped elements, higher than 1 lm,
with diameter smaller than its height, and thicker at
the apex than at the base (Fig. 12(14 and 15)).
4.1.10 Echinate/Spinate
These have ornamentation with spines >1 lm
where the height is at least twice the basal
diameter (Fig. 12(16)).
4 Sporoderm Stratification 59

Fig. 12 Sporoderm structure and various modifications (modified from Tschudy 1969). See text for further
explanation
60
4.1.11 Rugulate
These are elongated elements, >1 lm long,
arranged in an irregular pattern that is interme-
diate between striate and reticulate (Fig. 12(18
and 19)).
4.1.12 Reticulate/Reticulum
These have a network-like pattern consisting of
lumina or other spaces wider than 1 lm and
bordered by elements narrower than the lumina
(Fig. 12(20 and 21)). The lumina are pits
between the elevations of a reticulate sculptural
pattern (i.e., enclosed areas). Lumina can either
be rounded or polygonal in shape.
4.1.13 Striate
These are fine, elongated, parallel elements
(ridges) separated by grooves (Fig. 12(22 and
23)).
4.1.14 Faveolate
These are small (up to 2 µm), rounded depres-
sions that are more or less rounded in surface
view and widely spaced so as not to form a
reticulum (Fig. 12(24 and 25)).
4.1.15 Cicatricose
These are marked with long scars (more or less
parallel ridges) (Fig. 12(26)).
Nexine is the non-sculptured part of exine and
occurs below the sexine (Fig. 11(2, 4, and 5)).
Two subdivisions are noted, ectonexine and
endonexine on the basis of thickness and
refractive zone (Fig. 11(5)). Ectonexine is the
peripheral layer of nexine, exhibits more or less
thick and “not very refractive zone.” Endonexine
is the inner layer of nexine, more or less thin and
exhibits more refractive zone. Based on staining
characteristics, two layers are noted in nexine,
ectexine and endexine (Erdtman 1969). Ectexine
stains positively with basic fuchsin while
endexine remains relatively unstained. Ectexine,
the outer part of the exine also has higher elec-
tron density in conventional prepared TEM sec-
tions and includes the foot layer, if present
(Fig. 11(4 and 5)). Endexine, the inner part of the
exine, has a lower electron density in conven-
tionally prepared TEM sections (but this
3 Pollen and Spores
assertion is hotly debated); it does not include the
nexine which is considered part of the ectexine
(Fig. 11(4 and 5)).
5 Earliest Plants
Plant megafossils are rare, as they lack fossiliz-
able tissues. However, there is a large microfossil
record of dispersed spores (cryptospores) pre-
served (see Steemans 2000; Steemans et al.
2000).
Cryptospores are spore-like dispersed remains
of subaerial (and possibly, semi-aquatic) plants
that lack the haptotypic features (trilete marks)
and characterize vascular (tracheid-bearing) plant
spores (although rare non-vascular plants also
have spores with trilete mark). They are alete
(i.e., lacking a well-defined aperture), and rela-
tively thick-walled pseudomorphs that occur in
unusual configuration (monads, permanently
united dyads or tetrads) (Richardson et al. 1984;
Strother 1991; Richardson 1996; Steemans 1999,
2000; Strother 2000; Wellman and Gray 2000;
Strother and Beck 2000). The cryptospores are
the characteristic component of non-marine
palynomorph assemblages from the Palaeozoic
strata of pre-Devonian age. Because macroscopic
and mesoscopic plant remains do not occur prior
to the Homerian (430.5–427.4 Ma), and cryp-
tospores are used as a proxy for terrestrial plants
during the Early Paleozoic. The timing and
extent of the first terrestrial (subaerial) plant
cover is still a matter of much speculation, but
the cryptospore record indicates that plants of a
bryophyte grade were available to colonize sur-
face habitats from the Middle Cambrian
onwards.
A diverse cryptospore assemblage of five
genera of naked and envelope-enclosed monads
and tetrads are the earliest palynological evidence
dating back to earliest Ordovician (*473–
471 Ma) from Argentina (eastern Gondwana)
(see Rubinstein et al. 2010). These are: Cho-
motriletes? sp., Gneudnaspora (Laevolancis) di-
vellomedia or Laevolancis chibrikovae,
Sphaerasacus glabellus and Tetrahedraletes
cf. medinensis (see Rubinstein et al. 2010).
5 Earliest Plants 61

Fig. 13 Representative species diagrams

62 3 Pollen and Spores

Fig. 14 Representative species diagrams (continued…)

5 Earliest Plants
Fig. 15 Representative species diagrams (continued…)
Previously, the earliest embryophyte traces were
the late Darriwilian (Middle Ordovician; *463–
461 Ma) cryptospores from Saudi Arabia and
from the Czech Republic (western Gondwana)
(Vavrdová 1990; Strother et al. 1996). The anal-
ysis of wall ultrastructure of slightly younger
cryptospores from Oman indicated that they were
produced by early liverworts (Wellman et al.
2003). These evidences indicate that the embry-
ophytes first evolved in the Gondwana and that
the terrestrialization of plants might have begun
63
in the eastern part of Gondwana (see Rubinstein
et al. 2010). The Argentinean discovery, the
earliest evidence of plants on land, also suggests
an earlier origin of embryophytes, latest
Cambrian-Early Ordovician (488–472 Ma) or
even Cambrian (*541–488 Ma) (see Strother
and Beck 2000). The vascular plants are consid-
ered to have originated and adaptively radiated
from the Early Silurian on the basis of major
changes in spore assemblages (from cryptospore
to hilate/trilete spore dominated) and the
64
appearance of vascular plant megafossils (see
Steemans et al. 2009).
Some representative species are illustrated
with their respective time ranges (Figs. 13, 14
and 15).
References
Chaloner, W. G., & Sheerin, A. (1981). The evolution of
reproductive strategies in early land plants. In G. G. E.
Scudder, & J. L. Reveal (Eds.), Evolution today.
Proceedings of the Second International Congress of
Systematic and Evolutionary Biology, 93–100.
Erdtman, G. (1943). An introduction to pollen analysis
(p. 239). Waltham, MA: Chronica Botanica Company.
Erdtman, G. (1945). Pollen morphology and plant taxon-
omy. Labiatae, Verbenaceae and Avicenniaceae. Sven
Bot Tidskr, 39, 279–285.
Erdtman, G. (1947). Suggestions for classification of
fossil and recent pollen grains and spores. Svensk
Botanisk Tidskrift, 41, 104–114.
Erdtman, G. (1952). Pollen morphology and plant
taxonomy. Angiosperms (p. 539). Stockholm: Almq-
vist and Wiksell.
Erdtman, G. (1969). Handbook of palynology—An intro-
duction to the study of pollen grains and spores
(p. 486). Copenhagen: Munksgaard.
Faegri, K., & Iversen, J. (1964). Textbook of pollen
analysis (2nd ed., 237 pp). Copenhagen: Munksgaard.
Faegri, K., & Iversen, J. (1975). Textbook of pollen
analysis (3rd ed.). New York: Hafner.
Faegri, K., & Iversen, J. (1989). Textbook of pollen
analysis (4th ed., 328 pp). Chichester: John Wiley &
Sons, Inc.
Fritzsche, J. (1837). Über den Pollen. Mem Sav Etrang
Acad, 3: St. Petersburg, 649–769.
Grebe, H. (1971). A recommended terminology and
descriptive method for spores. Comm. Intern. Micro-
flore Palaeozoique, 4: Les Spores, 1, 7–34.
Halbritter, A. H., Halbritter, H., Ulrich, S., Grímsson, F.,
Weber, M., Zetter, R., et al. (2018). Illustrated pollen
terminology (486 p). Berlin: Springer.
Harris, W. F. (1955). A manual of the spores of New
Zealand Pteridophyta. A discussion of spore morphol-
ogy and dispersal with reference to the identification
of the spores in surface samples and as microfossils.
New Zealand Department of Scientific and Industrial
Research Bulletin, 116, 1–186.
Hesse, M., Halbritter, M., Zetter, R., Weber, M., Buchner,
R., Frosch-Radivo, A., & Ulirch, S. (2009). Pollen
terminology: An illustrated handbook. Vienna:
Springer.
Kremp, G. O. W. (1968). Morphologic encyclopedia of
palynology (2nd ed., 263 pp). Tucson: University of
Arizona Press.
3 Pollen and Spores
Potonié, R. (1934). I. Zur Morphologie der fossilen Pollen
und Sporen. Arb. Inst. Paläobotanik Petrographie
Brennsteine, 4, 5–24.
Potonié, R., & Kremp, G. O. W. (1955). Die Sporae
dispersae des Ruhrkarbons, ihre Morphographie und
Stratigraphie mit Ausblicken auf Arten anderer Gebi-
ete und Zeitabschnitte. Palaeontographica
Abteilung B, 98, 1–136.
Punt, W., Blackmore, S., Nilsson, S., & Le Thomas, A.
(1994). Glossary of pollen and spore terminology.
LPP Contributions Series 1, LPP Foundation, Labo-
ratory of Palaeobotany and Palynology, University of
Utrecht, Utrecht.
Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S., & Le
Thomas, A. (2007). Glossary of pollen and spore
terminology. Review of Palaeobotany and Palynology,
143, 1–81.
Richardson, J. B. (1996). Taxonomy and classification of
some new Early Devonian cryptospores from Eng-
land. Special Papers in Palaeontology, 55, 7–40.
Richardson, J. B., Ford, J. H., & Parker, F. (1984).
Miospores, correlation and age of some Scottish
Lower Old Red Sandstone sediments from the Strath-
more region (Fife and Angus). Journal of
Micropalaeontology, 3(2), 109–124.
Rubinstein, C. V., Gerrienne, P., de la Puente, G. S.,
Astini, R. A., & Steemans, P. (2010). Early Middle
Ordovician evidence for land plants in Argentina
(eastern Gondwana). New Phytologist, 188, 365–369.
Steemans, P., (1999). Paléodiversification des spores et
des cryptospores de l'Ordovicien au Dévonien infér-
ieur. Géobios, 32(2), 341–352.
Steemans, P. (2000). Miospore evolution from the
Ordovician to the Silurian. Review of Palaeobotany
and Palynology, 113, 189–196.
Steemans, P., Higgs, K. T., & Wellman, C. H. (2000).
Cryptospores and trilete spores from the Llandovery,
Nuayyim-2 Borehole, Saudi Arabia. In S. Al-Hajri &
B. Owens (Eds.), Stratigraphic palynology of the
Palaeozoic of Saudi Arabia (Special Vol. 1, pp. 92–
115). Manama, Bahrain: GeoArabia.
Steemans, P., Le Hérissé A., Melvin, J., Miller, M. A.,
Paris, F., Verniers, J., & Wellman, C. H. (2009).
Origin and radiation of the earliest vascular land
plants. Science, 324, 353.
Strother, P. K. (1991). A classification schema for the
Cryptospores. Palynology, 15, 219–236.
Strother, P. K. (2000). Cryptospores: The origin and early
evolution of the terrestrial flora. In R. A. Gastaldo &
W. A. DiMichele (Eds.), Phanerozoic terrestrial
ecosystems (Vol. 6, pp. 3–19). The Paleontological
Society Papers.
Strother, P. K., Al-Hajri, S., & Traverse, A. (1996). New
evidence for land plants from the lower Middle
Ordovician of Saudi Arabia. Geology, 24, 55–59.
Strother, P. K., & Beck, J. H. (2000). Spore-like
microfossils from Middle Cambrian strata: Expanding
the meaning of the term cryptospore. In M. M. Harley,
C. M. Morton, & S. Blackmore (Eds.), Pollen and
References
spores: Morphology and biology (pp. 413–424). Kew,
UK: Royal Botanic Gardens.
Traverse, A. (2007). Paleopalynology. In N. L. Landman
& D. S. Jones (Eds.), Topics in geobiology (2 ed.,
pp. 1–813). Dordrecht, The Netherlands: Springer.
Tschudy, R. H. (1969). The plant kingdom and its
palynological representation. In Aspects of palynology
(518 pp). Wiley-Interscience.
Vavrdová, M. (1990). Early Ordovician acritarchs from
the locality Myto near Rokycany (late Arenig,
65
Czechoslovakia). Casopis pro Mineralogii a Geologii,
35, 239–250.
Wellman, C. H., & Gray, J. (2000). The microfossil
record of early land plants. Philosophical Transac-
tions of the Royal Society of London, 355B, 717–732.
Wellman, C. H., Osterloff, P. L., & Mohiuddin, U. (2003).
Fragments of the earliest land plants. Nature, 425,
282–285.
 Dinoflagellates
4

(Fig. 1(1)). Class Dinophyceae is divided into

1 Introduction
three orders (and known only from fossil
record)—Dinophysiales, Gymnodiniales, and
Dinoflagellates (=whirling whips) are ubiquitous
Peridiniales (=the thecate dinoflagellate) (see
unicellular organisms of the aquatic ecosystems
Fig. 1(2–4), respectively) and are an important
and second only to diatoms as marine primary
constituent of the oceanic phytoplankton scene,
producers. Of the *2000 living species, 85% are
since the mid-Mesozoic. They are also one of the
marine and about 11% freshwater forms (Taylor
most primitive eukaryotes and are intermediates
et al. 2008). Almost half of the total species are
between prokaryotes and eukaryotes. Around
either photosynthetic or exclusively heterotrophic
4500 species assigned to about 550 genera are
(Gaines and Elbrächter 1987), hence, making
known, and of them, three quarters of the genera
them an important member of both marine and
and half of the species are fossil. Most dinocysts
freshwater phytoplankton and zooplankton
(the resting cysts that are resistant and occur in
ecosystems. Additionally, together with diatoms
fossil records, as opposed to the unmineralized
and coccolithophores (Calcareous nannofossils),
motile cells) are between 25 and 250 µm in
they are a major fraction of the eukaryotic phy-
diameter; they may be spherical to ellipsoid to
toplanktons of the world’s oceans (Delwiche
elongate in shape. The dinoflagellates are best
2007). The earliest fossil dinoflagellate cysts are
known for causing harmful algal blooms wherein
known from the Ladinian of the Middle Triassic
the dinoflagellates make up almost 80% of the
(Riding et al. 2010). The dinoflagellates are
toxic phytoplankton species (Cembella 2003).
excellent index fossils due to their restricted
These “red tides” kill fish and/or shellfish either
stratigraphic ranges and large spatial distributions
directly, because of the toxins they produce (di-
(Powell 1992; Stover et al. 1996; Poulsen and
noxanthin and peridinin), or due to the effects
Riding 2003; Traverse 2007; Riding 2012) (see
caused by the large numbers of dinoflagellate
Tables 1, 2, 3, 4, 5, 6, 7, 8, and 9).
cells that clog animal gills, and deplete oxygen
Traditionally, four classes constitute the algal
(Smayda 1997); the dinoflagellate toxins are also
Division Pyrrhophyta—Ebriophyceae, Ellobio-
among the most potent known biotoxins.
phyceae, Desmophyceae, and Dinophyceae

© Springer Nature India Private Limited 2020 67

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_4
68 4 Dinoflagellates

Table 1 Stratigraphic occurrences of important marker taxa (203–175 Ma)

 1
Introduction

Table 2 Stratigraphic occurrences of important marker taxa (173–164 Ma)

69
 70

Table 3 Stratigraphic occurrences of important marker taxa (163–146 Ma)

4
Dinoflagellates
 1
Table 4 Stratigraphic occurrences of important marker taxa (145–127 Ma)
Introduction
71
72 4 Dinoflagellates

Table 5 Stratigraphic occurrences of important marker taxa (126–101 Ma)

1 Introduction 73

Table 6 Stratigraphic occurrences of important marker taxa (100–67 Ma)

74 4 Dinoflagellates

Table 7 Stratigraphic occurrences of important marker taxa (64–42 Ma)

flagella (longitudinal and transverse) arising from

2 General Morphology
The dinoflagellates have large vacuoles (pusules;
Fig. 1(3)) that are connected to the exterior by a
delicate canal. The protoplast possesses two
flagellar pores in the cell wall, and whose loca-
tion is important in classification (Fig. 1(5–7)).
The whip-like longitudinal flagellum is directed
backwards along a longitudinal furrow (or de-
pression) called the sulcus; it is a groove in the
2 General Morphology
Table 8 Stratigraphic occurrences of important marker taxa (40–21 Ma)
ventral surface (Fig. 1(5–7)). The side bearing
the sulcus is ventral (Fig. 1(6)), while the oppo-
site side is dorsal (Fig. 1(7)). Many cells and
cysts are dorsoventrally compressed so that these
two views are the ones usually illustrated (as in
Genus Cyclonephelium). The transverse flagel-
lum that encircles the cell (once or several times)
in a transverse or spiral groove (=transverse
furrow or girdle; Fig. 1(4)) is called the cingulum
(Fig. 1(5)). If it encircles several times, the
75
cingulum is then twisted around the cell and is
defined as Torsion (see Fig. 2). The sulcus and
cingulum intersect near the mid-ventral point
(Fig. 1(6)). The cingulum divides the cell into an
anterior area called the episome (also called ep-
itheca or epicone), and a posterior area, the hy-
posome (Fig. 1(6)). The sulcus lies wholly or
partially in the hyposome (also called hypotheca
or hypocone) (Fig. 1(5 and 6)). The sulcus
extends in a posterior direction and may
76
Table 9 Stratigraphic occurrences of important marker taxa (22 Ma – Present)
terminate in a depression flanked by one or two
antapical horns (Fig. 1(4, 6, and 7)). The anterior
or apical end is often rounded, pointed, or pro-
duced into an apical horn (Fig. 1(7)). The ante-
rior portion of the episome is called the apex
(Fig. 1(5)). The posterior portion (excluding
spines and similar structures) of the hyposome is
called the antapex (Fig. 1(5–7)).
Around the epitheca, the apical and precin-
gular series occur (Figs. 3 and 4). In the
4 Dinoflagellates
cingulum, lies the cingular series while the
postcingular and antapical series occur on the
hypotheca (Figs. 3 and 4). Additional anterior
and posterior intercalary plates may also develop
at sites between the series, and the sulcus bears
small sulcal plates that are of taxonomic impor-
tance. The Intercalaries (the secondary series of
plates) never touch the apex, antapex, or cingu-
lum. The series and number of plates can be
expressed as the plate formula, using Kofoid’s
2 General Morphology 77

Table 10 Alphanumeric system of plate designation (after Kofoid 1907, 1909) (see also Figs. 3 and 4)
Vegetative Encysted Location Symbol Individual plate or
stage stage paraplate
Plate series Paraplate Position Symbol
series
Apical Apical Touching the apex but not the cingulum Second 2′
or paracingulum apical
Anterior Anterior Between the apicals and precingulars but a Third 3a
intercalary intercalary not touching the apex or cingulum anterior
(paracingulum); can be ventrally or intercalary
dorsally located
Precingular Precingular Immediately anterior to the cingulum or ″ Fifth 5″
paracingulum and not touching the apex precingular
Cingular Paracingular Cingulum or c or Fourth 4c
cingular
Or girdle Paracingulum pc Or 4pc
paracingular
Postcingular Postcingular Immediately posterior to the cingulum or ″′ Sixth 6″′
paracingulum and not touching the postcingular
antapex
Posterior, Posterior, Between the postcingulars and antapical p First 1p
intercalary intercalary (s) and not touching the cingulum posterior
(paracingulum) or antapex; usually intercalary
adjacent to the sulcus or parasulcus
Antapical Antapical Touching the antapex but not the ″″ First 1″
cingulum or paracingulum antapical
Sulcal Parasulcal Occupying a position on the s Anterior ls
sulcal
Sulcus or parasulcus ps Parasulcal lps

alphanumeric system (Table 1; Figs. 3 and 4).
Adjacent plates are firmly cemented together
along their margins where they overlap in a
tile-like fashion. Growth occurs in the sutural
areas between plates, in the form of intervening
bands, called the Intercalary bands (Fig. 3).
A series of articulated plates (composed of
cellulose or some other polysaccharide
microfibrils) makes the theca (alternatively also
called Amphiesma, Cell covering, or Cell wall);
the plate number and position constitutes the
Tabulation (i.e., counting the plates in a specific
designated series) (Figs. 3 and 4). Their number
and position are of taxonomic importance
(Figs. 3 and 4). This nomenclature is best
exemplified by the Peridiniales (Fig. 3(1–5)) as
first laid out by Kofoid (1907, 1909). In Peri-
diniales, the five-plate series are found to encircle
each cell, and each plate is numbered in a
counterclockwise direction. From apex to anta-
pex, these include apical (′), anterior intercalary
(a), and precingular series (″) of the epitheca, the
cingular series (c) composing the cingulum, and
the postcingular (‴), posterior intercalary (p) and
antapical series (″″) of the hypotheca (Fig. 3(1–
4)). The primary series of plates include—api-
cals, precingulars, cingulars, postcingulars,
antapicals and sulcals (see Table 10; see also
78 4 Dinoflagellates

Fig. 1 Characteristics of the Dinophyceae. 1: Broad classification of dinoflagellates; 2: Order Dinophysiales (ventral
view of Dinophysis collaris Kofoid and Michener); 3: Order Gymnodiniales (Gymnodinium); 4: Theca of Peridinium
leonis Pavillard; 5: Gymnodiniale, showing an extant “Naked” dinoflagellate (a gymnodinioid stage); 6: General
terminology for hypo- and epitheca (modified from Haq and Boersma 1998)
2 General Morphology
Fig. 2 Types of cingulum (modified from Sarjeant 1974)
Figs. 3 and 4). Each plate series has a superscript
designation or a letter. For example, 3′ means
that there are three apical (′) plates. More
recently, cingular (c) and sulcal (s) plate series
have been added to the plate formula as well as
components of the APC, i.e., Po, cp, and X, if
they occur. Thus, the plate formula is the com-
bined tabulations for the plate series—Po, 4′, 0a,
6″, 8c, 5 s, 5″, lp, 3″″. A plate formula is gen-
erally stable within a genus; however, a range in
number for a specific series can occur due to
splitting of plates or other factors.
79
2.1 Cyst Types
Cyst lengths are between 15 and 500 l, but
generally, they range between 60 and 120 l. The
presence or absence of processes enables to
recognize four major types of cysts, proximate,
chorate, cavate, and proximochorate (Fig. 5).
Proximate cysts are devoid of processes, and the
shape of the cyst closely approaches that of the
theca; it probably formed in close contact with
the theca (Fig. 5(1 and 2)). The tabulation, cin-
gulum, and sulcus are all reflected in the surface
sculpture of a proximate cyst (Fig. 5(2)). The
80 4 Dinoflagellates

Fig. 3 1–4: Tabulation and plate symbols in Peridinium leonis Pavillard (4′, 3a, 7″, 5″ ′ 2″ ″). The notation indicates four
apical plates, three anterior intercalary plates, seven precingular plates, five postcingular plates, and two antapical plates
(modified after Lebour (1925). 5: Plate shape variations. a: Rhombic apical plate (1′) and the pattern of its relationship with
bordering plates in Peridinium; a1: Orthoperidinium (ortho); a2: Metaperidinium (meta); a3: Paraperidinium (para), b: The
second anterior intercalary plate (2a) in the various sections of Orthoperidinium and Metaperidinium, b1 and b2, penta; b3,
hexa; b4, quadra (modified after Graham 1942); 6 and 7: Wetzeliella reticulata Williams and Downie (Early Eocene), showing
the “peridinioid” paratabulation of the genus; 6: Ventral view; 7: Dorsal view (modified after Evitt 1969)
2 General Morphology 81

Fig. 4 1–2: Theca of Gonyaulax spinifera (Claparede and Lachmann) Diesing, showing tabulation (4″, 6″, ?6c, 6′″ ′,
1″ ″, Ip); 1: Ventral view; 2: Lateral view; c = cingular; p = posterior intercalary; (modified after Wall and Dale, 1970);
3–8: Ceratium sp., showing tabulation of the theca (4′, 6″, 5, and 6c, 6″ ′, Ip, 1 ″ ″); 3: Apical view; 4: Ventral view
(with an offset sulcus); 5: Antapical view; 6: Right lateral view; Dorsal view; 8: Left lateral view. For details on plate
notation, see Fig. 2

chorate cysts (Fig. 5(3 and 4)) possess a main Hystrichosphaeridium (Fig. 6(1 and 2)). chorate
body bearing strongly developed ornamentation cysts usually exhibit no traces of a reflected
or processes (see also Fig. 6), and best noted in cingulum or sulcus. The inner and outer bodies in
82 4 Dinoflagellates

Fig. 5 Cyst stages. 1 and 2: Proximate cyst; 1: Peridinium (axial section); 2: Gonyaulacysta (1 and 2: modified after
Sarjeant 1974); 3 and 4: Chorate cyst; 3: Gonyaulax with detail of wall (axial section) (modified after Sarjeant 1974); 4:
Hystrichosphaeridium (modified after Sarjeant 1974); 5 and 6: Cavate cyst of Deflandrea; 5: Axial section; 7:
Proximochorate cyst of Spiniferites (modified after Evitt 1969)
2 General Morphology 83

Fig. 6 1–2: Paratabulation of a chorate cyst, Hystrichosphaeridium tubiferum (Ehrenberg) Deflandre showing the
number and position of intratabular processes. 1: Upper surface showing apical tetratabular archeopyle (shaded),
precingular processes (1”, etc.) and paracingular processes (pc); 2: Lower surface showing postcingular processes
(1” “, etc.), parasulcal processes (ps), single antapical process (1” “), and posterior intercalary processes (p) (modified
from Haq and Boersma 1998); 3-6: Process types in chorate dinocysts; 3: Spiniferites ramosus (Ehrenberg), Dorsal
view; 4: Oligosphaeridium complex (White) with one intratabular process per paraplate; 5: Hystrichokolpoma eisenacki
Williams and Downie with process differentiation; 6: Diphyes colligerum (Deflandre and Cookson) with enlarged
antapical process (black)
84
Fig. 7 Cardinal positions from which horns originate in
a dinocyst; the maximum is 5 (see text for explanation)
a cavate cyst (Fig. 5(5 and 6)) are only in limited
contact with each other and have an intervening
space between them. This group includes many
cysts of the peridinioid type such as Deflandrea
(Fig. 5(6)). The cyst wall (called Phragma) is
built of organic material and is resistant to bac-
terial decay. If the autocyst has a single layer, its
wall is called an autophragm. A two-layered cyst
with connections between the walls has an inner
layer, the autophragm, and an ectophragm, and
this condition is termed as holocavate. If the two
layers are not connected, the cyst is known as
cavate and the inner layer is called the en-
dophragm, and the outer layer, the periphragm,
which are partially separated, usually at the poles
(Fig. 5(5)). The cavities thus formed (pericoels)
promote buoyancy in the cyst (Fig. 5(5)).
Another cyst type, an intermediate between
4 Dinoflagellates
proximate and chorate, is the proximochorate and
is characterized by an elaborate ornamentation
(Fig. 5(7)). In this type, the tips of the processes
are joined by thin, filamentous trabeculae, thus,
giving the impression of an additional layer.
Some cysts have a smooth surface, while
others possess ornamentation that can be either
of minor elements such as granules or of major
elements like horns, septa or processes (see
Figs. 5 and 7). The horns may be apical,
antapical, or lateral (Fig. 7); these are useful
criteria for establishing the orientation of a
dinocyst. A dinocyst never has more horns than
its corresponding thecate stage and in both, the
maximum is five. The horns can be over 100 µm
in length but are generally less than 50 µm.
The processes are essentially columnar or
spine-like (Fig. 5(3)), whereas septa are mem-
braneous, linear projections arising perpendicu-
larly from the outer wall layer. Processes may be
located on or within paraplate boundaries. Pro-
cess complexes are processes on individual
paraplates that are united proximally, distally, or
along their length and can be plate centered or
form groups (Fig. 8). The ornament can be tab-
ular (sutural) if it defines plate boundaries, or
intertabular if it defines the central parts of plates
(Fig. 8). Processes that are situated at the inter-
section of paraplate boundaries are called Gonal
and those along boundaries are called Intergonal
(Fig. 8(13)). The processes of a dinocyst may be
either similar (see Fig. 6(1 and 2)) or they may
be differentiated (see Fig. 6(5)); a particular
paraplate series is often characterized by a diag-
nostic type of process. In other taxa, some of the
paraplates, commonly the cingular, are devoid of
processes (Fig. 6(4)). The preferred orientation
of septa and processes often permits one to
determine paratahulation in a dinocyst. Process
arrangements, shapes, and terminations are
important morphologic features in generic and
specific classification.
2 General Morphology 85

Fig. 8 1–6: Intratabular process complexes; 1: Annulate; 2: Soleate; 3: Arcuate; 4: Linear; 5: Simulate’ 6: Simulate
(adjacent processes joined); 7–14: Surface ornament; 7–8: General terminology (see text for explanation); 9:
Non-tubular; 10: Paratubular; 11: Peritubular; 12: Parasutural septae; 13: Gonal and Intragonal processes; 14:
Intratabular process group (modified after Evitt 1969)

2.2 Archeopyle apical series, the precingular series, an anterior

The function of the cyst is demonstrated by the
presence of an escape hole, called an archaeopyle
(from the Greek words arche: old and pyle: gate
or orifice) (Fig. 9). This regular-shaped opening
is called the operculum that results from the
complete or partial removal of one or more than
one paraplate. If the archeopyle is formed from
the rupture of the “suture” between adjacent
paraplates, then, in such cases the operculum is
absent. The plate removal is normally from the
intercalary plate or a combination of these.
Hence, this rupturing of plates (=archeopyle)
may occur along the girdle, along the line of
sutures between the apical and precingular series
of plates, or more commonly by the release of a
single plate of the epitheca. The form and posi-
tion of the archeopyle is constant within a genus,
a genetically determined feature and constant in
all known species. There are of three major types
of archeopyles—apical, intercalary, and precin-
gular (Fig. 9) (for details see Sarjeant 1974).
86 4 Dinoflagellates

Fig. 9 Archeophyle arrangements. 1: Archeophyle terminology; 2–3 and 6: Apical archeophyle; 2: Dorsal view; 3:
Apical view; 4 and 7–9: Intercalary archeophyle; 5 and 10–13: Precingular archeophyle; 14–17: Combination
archeophyle (modified after Evitt 1969)
3 Geological History
Fig. 10 Dinoflagellate species diversity, major radiation, and recovery patterns
3 Geological History
The earliest record of an equivocal peridinoid cyst
is Arpylorus antiquus Calandra from the Silurian
(Tunisia). It has been interpreted as possessing
tabulation, a cingulum and a precingular arche-
opyle. However, this has been strongly contested
(see Herisse Le et al. 2012) who do not consider
A. antiquus as a dinoflagellate. According to these
authors, based on biomarker record, the
dinoflagellates may have older Paleozoic or even
Proterozoic ancestors; the dinoflagellate tabula-
tion evolved only in the early Mesozoic.
The main dinoflagellate radiation began in the
Mid- to Late Triassic (Fig. 10) with the appear-
ance of genera such as Suessia or the Rhaeto-
gonyaulax from the Rhaetian (Late Triassic).
Proximate cysts were common throughout the
87
Jurassic (Gonyaulacysta jurassica), although
chorate and proximochorate cyst types had
appeared by the Middle Jurassic. Many Creta-
ceous forms are chorate (such as Hystrichos-
phaeridium) or proximochorate (such as
Spiniferites ramosus), and it was around this time
that the greatest diversity of dinoflagellate cysts
was reached (Fig. 10). The cavate peridinoid
dinoflagellate cysts began to flourish in Aptian–
Albian (Deflandrea and Wetzeliella) and domi-
nated many Tertiary assemblages until the Oli-
gocene, but almost died out in the Pliocene.
Proximate and chorate dinoflagellate cysts with
complex processes occur in the Eocene and
Oligocene, but simpler forms have prevailed
since then. Dinoflagellate cysts first appeared in
freshwater sediments during the Tertiary.
The representative dinoflagellate species are
given in Figs. 11, 12, 13, and 14.
88 4 Dinoflagellates

Fig. 11 Some representative chorate dinoflagellate species

3 Geological History 89

Fig. 12 Some representative dinoflagellate species (continued…)

90 4 Dinoflagellates

Fig. 13 Some representative dinoflagellate species (continued…)

3 Geological History 91

Fig. 14 Some representative dinoflagellate species

92
References
Cembella A. D. (2003). Chemical ecology of eukaryotic
microalgae in marine ecosystems. Phycologia, 42,
420–447.
Delwiche, C. F. (2007). The origin and evolution of
dinoflagellates. In P. G. Falkowski & A. H. Knoll
(Eds.), Evolution of primary producers in the sea
(pp. 191–205). Burlington, VT: Academic Press.
Evitt, W. R. (1969). Dinoflagellates and other organisms
in palynological preparations. In R. H. Tschundy & R.
A. Scott (Eds.), Aspects of palynology (pp. 439–481).
New York, NY: Wiley Interscience.
Gaines, G., & Elbrächter, M. (1987). Heterotrophic
nutrition. In F. J. R. Taylor (Ed.), The biology of
dinoflagellates (pp. 224–268). Blackwell: Oxford.
Haq, B. U., & Boersma, A. (1998). Introduction to marine
micropaleontology (2nd ed., 376 pp). Elsevier.
Herisse Le, A., Masure, E., Javaux, E. J., & Marshall, C.
P. (2012). The end of a myth: Arpylorus antiquus
paleozoic dinoflagellate cyst. Palaios, 27, 414–423.
Kofoid, C. A. (1907). The plates of Ceratium with a note
on the unity of the genus. Zoologischer Anzeiger, 32,
177–183.
Kofoid, C. A. (1909). On Peridinium stecni Jorgensen,
with a note on the nomenclature of the skeleton of the
Peridinidae. Archiv für Protistenkunde, 16(1), 25–47.
Poulsen, N. E., & Riding, J. B. (2003). The Jurassic
dinoflagellate cyst zonation of Subboreal Northwest
Europe. Geological Survey of Denmark and Green-
land Bulletin, 1, 115–144.
Powell, A. J. (1992). Dinoflagellate cysts of the Triassic
system. In A. J. Powell (Ed.), A stratigraphic index of
4 Dinoflagellates
dinoflagellate cysts (pp. 1–6)., British Micropalaeon-
tological Society Publications Series London: Chap-
man and Hall.
Riding, J. B. (2012). A compilation and review of the
literature on Triassic, Jurassic, and earliest Cretaceous
dinoflagellate cysts. American Association of Strati-
graphic Palynologists, 227pp. (Contributions Series,
46). http://www.palynology.org/aasp-store/4.
Riding, J. B., Mantle, D. J., & Backhouse, J. (2010).
A review of the chronostratigraphical ages of Middle
Triassic to Late Jurassic dinoflagellate cyst biozones
of the North West Shelf of Australia. Review of
Palaeobotany and Palynology, 162, 543–575.
Sarjeant, W. A. S. (1974). Fossil and living dinoflagel-
lates. London: Academic Press.
Smayda, T. J. (1997). Harmful algal blooms: their
ecophysiology and general relevance to phytoplankton
blooms in the sea. Limnology and Oceanography, 42,
1137–1153.
Stover, L. E., Brinkhuis, H., Damassa, S. P., De Verteuil,
L., Helby, R. J., Monteil, E., et al. (1996).
Mesozoic-Tertiary dinoflagellates, acritarchs and
prasinophytes. In J. Jansonius & D. C. McGregor
(Eds.), Palynology: Principles and applications
(pp. 641–750). Dallas, TX: American Association of
Stratigraphic Palynologists Foundation.
Taylor, F. J. R., Hoppenrath, M., & Saldarriaga, J. F.
(2008). Dinoflagellate diversity and distribution. Bio-
diversity and Conservation, 17, 407–418.
Traverse, A. (2007). Paleopalynology (2nd ed.). The
Netherlands: Springer.
 Conodonts
5

The conodont color alteration index

1 Introduction
Conodonts (=cone-tooth in Greek) are usually
microscopic (0.2–2 mm in length) organisms,
although, rare larger specimens up to 25 mm have
been recorded (Gabbott et al. 1995; Purnell 1995).
Conodonts are elongate, armorless, eel-shaped,
and large-eyed marine animals (Briggs et al. 1983;
Purnell 1995) (Fig. 1). They are soft-bodied,
except for their apparatus of tooth-like phosphatic
(bioapatite with a francolite-like structure) ele-
ments situated in the mouth and/or the pharynx
(Fig. 2(1)). These elements (of the feeding appa-
ratus), after death, are scattered in the sediment on
the seafloor and are termed as natural assemblages
or bedding plane assemblages, sometimes also
composed of fused assemblages (Fig. 2(2)).
The conodonts are putative jawless vertebrates
(Donoghue et al. 2000) that spanned from the Late
Cambrian to the latest Triassic, a duration
of *300 Ma (Sweet 1988; Aldridge et al. 1993).
Their tooth-like elements (Fig. 2) are the earliest
instance of a mineralized skeleton within the
vertebrate lineage (Sansom et al. 1992; Donoghue
and Sansom 2002; Murdock et al. 2013; Martí-
nez-Pérez et al. 2014a, b). Conodonts are con-
sidered as active swimmers of the water column.
The conodonts are an important microfossil
group for dating Paleozoic shallow-marine car-
bonates and have been extensively used in pale-
oecological and biogeographical studies (Sweet
and Donoghue 2001) (Tables 1, 2, 3, 4, 5, and 6).
(CAI) (Epstein et al. 1977; Rejebian et al. 1987)
has been applied for the interpretation of basinal
histories, regional metamorphic studies (Wiederer
et al. 2002; Zhang and Barnes 2007; Voldman
et al. 2009, 2010) and for hydrocarbon and min-
eral exploration (Marshall et al. 2001; Trotter
et al. 2007), as well as in evaluating Kimberlite
(the primary natural source of diamonds)
emplacement temperatures (Pell et al. 2015).
Additionally, bioapatite (the element that consti-
tutes the animal’s skeleton) archives information
on sea water chemistry, hence, their chemical
analyzes enables a better insight into ocean geo-
dynamics and past climates (Holmden et al. 1996;
Trotter et al. 1999; Wenzel et al. 2000).
2 General Morphology
Fossil evidences of soft parts are extremely rare,
and wherever they are well-preserved, are sug-
gestive of a chordate affinity for conodonts
(Mikulic et al. 1985; Aldridge et al. 1986, 1993;
Smith et al. 1987; Sansom et al. 1992; Aldridge
and Theron 1993; Gabbott et al. 1995). Complete
feeding apparatus with 15 or more elements
(Fig. 3) have now been recorded (Briggs et al.
1983). Functional modeling (Aldridge et al.
1987; Purnell 1993; Purnell and Donoghue 1998)
and growth studies favor a grasping and pro-
cessing function for the apparatus (Purnell 1994,

© Springer Nature India Private Limited 2020 93

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_5
94 5 Conodonts

and the variable basal body (Fig. 4(1)). The basal

body occupies an opening, the basal cavity
(Fig. 4(2–5)), within the crown that comprises of
hyaline lamella tissue with growth lines (Fig. 4
(1)) and an internal opaque tissue, called the
white matter (Fig. 4(5)). The latter is so called,
as, in translucent fossils, it appears white under
reflected light. However, in transmitted light, it
appears dark. The basal cavity (Fig. 4(2–5)) is an
excavation that increases in size with specimen
growth. Hence, its size is dependent on the size
of the specimen. Another taxonomically impor-
tant structure is the pit (Fig. 4(7)) which is
formed when growth proceeds laterally from or
above the initial point of secretion. Its size does
not increase with ontogenetic growth but ceases
after a few growth lamellae have been formed
(Fig. 4(7)). Hence, large specimens may have
small pits. The germ denticles (Fig. 4(5)) are
structures that have been suppressed during
ontogeny due to overgrowth by adjacent struc-
tures; these are best noted in translucent fossils in
transmitted light. It must be mentioned that the
electron microscopy has revealed that the
Fig. 1 Line drawing of the Early Carboniferous (Dinan- lamellar crown tissue (Fig. 4(1)) is possibly
tian, Mississippian) Clydagnathus cf. cavusiformis homologous to the enamel and the white matter
Rhodes et al. (1969) from the Granton Shrimp Bed
(Edinburgh, Scotland). Modified from Briggs et al. (1983) may be a form of the enamel, quite unique to
conodonts. In general, the conodonts grew by
polycyclic, appositional addition of layers of the
1995); the elements being the components of the crown enamel, so that the inner lamellae are the
oropharyngeal feeding apparatus (Aldridge et al. oldest (Fig. 4(1)). The lamellae, within the
2013) (Fig. 2). crown, based on their incremental patterns, are
termed as minor and major (Armstrong and
Smith 2001). In Protopanderodus varicostatus,
2.1 Elements the minor increments have a minimum thickness
of *1 lm (likely deposited in a day), although,
Most preCarboniferous conodonts elements are some minor increments have been noted to be as
constructed of two parts, the crown (readily thick as 7 lm and most likely represent growth
preserved and the more abundant component) episodes, lasting up to a week.
Fig. 2 The conodont animal. 1: Line drawing of the conodont animal showing its major morphological features. 2: A
natural assemblage of conodont elements of Scottognathus typicus (Rhodes) from Pennsylvanian rocks in Illinois
(Scottognathus is a junior synonym of Idiognathodus) (after Sweet and Donoghue 2001). 3: The postero-ventral view of
the head of the conodont animal showing the position of elements (apparatus) of the Lochkovian (earliest Devonian)
Pandorinellina rernscheidensis (Ziegler) (modified after Dzik 1991). 4: Illustration of conodont’s M, S and P elements
in relationship to one another within the animal’s exposed oral cavity. 5: Enlarged view of the two sets of P elements (P1
and P2) (modified from Purnell and Jones 2012)c
2 General Morphology 95
96
Table 1 Stratigraphic occurrences of important marker taxa (203–175 Ma)
2.2 Composite Elements
The conodont elements are varied in shape and
are of taxonomic relevance (see Sweet 1988 for
details). Two major categories are noted: com-
pound or composite and simple or single teeth.
The composite elements are differentiated into
four main groups (Fig. 4(5–10)). These are:
2.2.1 Blade-Type Conodonts
These are elongate, laterally compressed forms
with a row of denticles that are fused, except at
5 Conodonts
their tips (Fig. 4(5)). They functioned as cutting
teeth (Purnell and von Bitter 1992). These are
index fossils from Silurian to Triassic.
2.2.2 Bar-Type Conodonts
These are commonly branched, and with thin
bars (Fig. 4(6)). They are the most conspicuous
and commonly occurring element within
many associations. They are evolutionarily con-
servative and long ranging with stable mor-
phologies, thus, biostratigraphically not
important.
2 General Morphology 97

Table 2 Stratigraphic occurrences of important marker taxa (173–164 Ma)

98 5 Conodonts

Table 3 Stratigraphic occurrences of important marker taxa (163–146 Ma)

2 General Morphology 99

Table 4 Stratigraphic occurrences of important marker taxa (145–127 Ma)

100
Table 5 Stratigraphic occurrences of important marker taxa (126–101 Ma)
2.2.3 Platform-Type Conodonts
These highly differentiated forms (Fig. 4(7–10)),
through the development of broad flanges into
plates, probably evolved from blade- and bar-
type conodonts (Fig. 4(5 and 6)). Many of these
platform-type conodonts are excellent index fos-
sils for the Ordovician–Triassic duration. Mor-
phologically, a blade is the edge-like free portion
at the anterior end, and in many cases, it integrates
into a carina on the upper side of the platform
(Fig. 4(8)). The protrusions on one or both sides
5 Conodonts
of the platform are called Lobes (Fig. 4(8)). On
the lower side of the platform (as in Polygnathus;
Fig. 4(7 and 8)), the main elements are: (a) the pit
around the growth center, (b) the attachment area,
where the edges of growth lamellae are visible,
(c) the crimp, or the portion outside the basal
organ that does not expose the edges of growth
lamellae, and (d) the keel in the center, which
corresponds to the carina at the upper surface. The
lower side of some platforms (as in Gnathodus;
Fig. 4(9 and 10)) is entirely excavated and
2 General Morphology 101

Table 6 Stratigraphic occurrences of important marker taxa (100–67 Ma)

102 5 Conodonts

Fig. 3 Apparatus architecture comparisons. 1: Six-element apparatus of Archeognathus primus; 2: Fifteen-element

apparatus of Iowagnathus grandis (modified after Liu et al. 2017); 3: Fifteen-element apparatus of the ozarkodinid type
(modified after Aldridge et al. 2013); 4: Ninteen-element apparatus of Promissum pulchrum. Modified after Donoghue
et al. 2008)
2 General Morphology 103

Fig. 4 Morphological terminology of conodonts. 1: Cross section of the Late Devonian Palmatolepis showing the
platform with anisometric growth. The lamellae are more widely spaced in one direction (modified after Müller and
Nogami, 1971). 2–4: Basal cavity. 3: Spathognathodus (Ordovician–Permian): Internal structure of conodonts lamellar
structure barely masked by white matter; 4: Ozarkodina (Silurian–Devonian): Lamellar structure completely filled with
white matter; 5: Blade type. 6: Bar type; 7–10: Platform type; 7 and 8: Polygnathus; 9 and 10: Gnathodus
104
Fig. 5 Crown terminology
display the edges of growth lamellae on the entire
surface, termed as cup (Fig. 4(10)). In the center,
a furrow (Fig. 4(9)), homologous to the keel, is
developed.
2.3 Simple Elements
The simple coniform elements have two parts, a
broad base that encloses the basal cavity and a
solid and tapering unit (apex) called the cusp;
cusp can be straight or curved (Fig. 5). While
interpreting conodont taxonomy, the tip (apex) is
always placed up with the base, horizontal
(Fig. 5). Based on curvature, the coniform ele-
ments are divided into two, geniculate and
non-geniculate (Fig. 5; see also Fig. 6, inbox).
Based on the number of processes and their
relationship to cusp, the ramiform elements are
divided into seven types, alate, digyrate (exten-
siform and breviform), quadriramate, dolabrate,
bipennate, tertiopedate, and multiramate (Fig. 6
(1–7)). Based on the number and arrangement of
primary processes, the pectinform elements are
divided into five types—stellate, pastinate,
5 Conodonts
angulate, segminate, and carminate (Fig. 6(8–
12)). Element details are illustrated in Fig. 6 and
are briefly enumerated below.
2.3.1 Ramiform Elements
Alate
These elements are bilaterally symmetrical and
have a caudal and two lateral processes (Fig. 6(1)).
Digyrate
These elements are broadly similar in shape to
alate elements, but they are asymmetrical; the
caudal process is only rarely developed, and the
lateral processes are usually unequally developed
and variably twisted in opposite directions
(Fig. 6(2 and 3)).
Quadriramate
These elements have rostral, caudal, and two
lateral processes (Fig. 6(4)).
Dolabrate
These elements only have a caudal process and
are commonly pick-shaped (Fig. 6(5)).
2 General Morphology 105

Fig. 6 Element terminology

106 5 Conodonts

Fig. 7 Topological scheme of notations. 1: Comparative schemes put forward by Purnell et al. (2000) and Sweet
(1988). 2: Nicoraella apparatus architecture (after Huang et al. 2018)

Bipennate 2.3.2 Pectinform Elements

These elements have a caudal and rostral process; Elements with a dorsal process are again divided
the rostral is usually shorter and commonly into categories based on the number of primary
curves or is deflected inwards (Fig. 6(6)). processes (Fig. 6(8–12)).

Tertiopedate Stellate
These elements have a caudal process and lateral These elements have four primary processes that may
processes that are asymmetrically disposed about bifurcate to form secondary processes (Fig. 6(8)).
the cusp (Fig. 6(7)).
2 General Morphology
Pastinate
These elements have three primary processes,
dorsal, ventral, and a rostral or caudal process.
The processes may be adenticulate represented
only by a conspicuous flange (Fig. 6(9)).
Carminate and Angulate
These elements have dorsal and ventral pro-
cesses. In a carminate element, the aboral margin
is straight, whereas in angulate ones, this is
arched (Fig. 6(10 and 12)).
Segminate
These elements possess a dorsal process that can
bear one or more rows of nodes or ridges (Fig. 6
(11)).
2.4 Element Notation and Apparatus
The elements form a bilaterally symmetrical
apparatus (Fig. 7) located in the head region of
the conodont animal. Each conodont species is
reconstructed as having a certain number of
elements with various shapes, including the
anteriorly positioned M and S (Sa, Sb, Sc, and
Sd), and the posteriorly-positioned P (Pa and Pb)
elements (see Sweet 1988; Purnell et al. 2000)
(Fig. 7(1)). Broadly, the elements fall into two
distinct domains: a rostral domain of paired S
elements (Sb, Sc, and Sd; plus a single Sa ele-
ment on the midline), associated with a pair of
dorsolateral M elements, and a caudal domain
comprising up to four pairs of P elements (the Pa,
Pb, Pc, and Pd elements) (Fig. 7; see Sweet
1988). Later, Purnell et al. (2000) introduced new
terms where element locations are defined
according to the relationships between elements
107
with reference to the principal axes of the body,
confirmed in bedding plane assemblages (Fig. 7).
In the standard ozarkodinid apparatus, it takes the
form of letters with numeric subscripts (such as
P1, P2, S0–S4, M; the S locations are numbered
outwards from the central S0 (Fig. 7).
The three-dimensional architecture of rela-
tively few conodont apparatuses are known
(Aldridge et al. 1987; Purnell and Donoghue
1998) wherein, the apparatus of the ozarkodinids
is considered as the stereotypical apparatus for all
non-coniform species (Fig. 3). However, in
recent years, based on the find of discrete ele-
ments, fused clusters, bedding plane assem-
blages, and articulated assemblages associated
with the soft-tissue, several apparatus models
have been used to reconstruct the conodont ani-
mal (Purnell and Donoghue 1997, 1998; Orchard
2005; Goudemand et al. 2011, 2012; Agematsu
et al. 2017; Zhang et al. 2017). But, owning to
the animal’s disintegrated nature (after death),
large discrepancies remain among these models.
Additionally, even in well-preserved natural
assemblages (bedding plane assemblages and
fused clusters), the component elements are
masked due to one or a combination of the fol-
lowing factors. They are obscured (see Fig. 1
(2)): (1) by one another, (2) by the surrounding
matrix, and/or (3) by diagenetic mineral crusts.
For cases like these, recently, a non-destructive
tomographic technique, the synchrotron radiation
X-ray tomographic microscopy (SRXTM) has
been used successfully employed to better
understand (and document) the architecture of
the conodont apparatus (Fig. 8) (see also Jones
et al. 2012a, b; Murdock et al. 2013, 2014;
Martínez-Pérez et al. 2014, 2016; Suttner et al.
2017; Huang et al. 2018).
108 5 Conodonts

Fig. 8 Surface models of Middle Triassic 15-element Nicoraella showing apparatus architecture as revealed by
synchrotron X-ray microtomography (SRXTM) from the Luoping Biota (Yunnan Province, Southwest China).
Illustration shows “a single alate element located in the S0 position, flanked successively abaxially by pairs of breviform
digyrate S1 and S2 elements, bipennate S3 and S4 elements, and a pair of inwardly curved breviform digyrate M
elements. Carminate elements occupy the P1 and P2 positions” (after Huang et al. 2018)
2 General Morphology 109
 110 5 Conodonts

b Fig. 9 Conodont classification (after Sweet 1988). 1–7: Order Proconodontida: “uni-membrate skeletal apparatus
composed of relatively large, deeply excavated, smooth-surfaced, hyaline coniform elements that are subsymmetrically
oval transverse sections and keels on the anterior and/or posterior margins” (Sweet 1988); 1: Proconodontus (Late
Cambrian), the oldest known conodont; 2: Eoconodontus (latest Cambrian–Early Ordovician); 3: Corydylodus (Late
Cambrian–Early Ordovician); 4: Fryxellodontus; (Late Cambrian–Early Ordovician); 5: Ansella (Middle Ordovician); 6:
Belodella (Silurian–Devonian); and 7: Corydylodus 8–22: Order Prioniodontida: “conodonts in which one or both of the P
positions in the apparatus are occupied by pastinate coniform or pectiniform elements or their platformed equivalents”
(Sweet 1988); both the Early Ordovician Rossodus (8) and Tripodus (9) have the oldest prioniodontide skeleton (some
authors have assigned Tripodus to Acodus or Diaphorodus; 10: Pygodus (Ordovician); 11: Icriodella (Early Silurian); 12:
Pedavis (Early Middle Devonian); 13: Distomodus (Silurian; morphologically Distomodus is very close to Pedavis; Sweet
1988); 14: Multiostodus (Ordovician); 15: Prioniodus (Early Middle Ordovician; Whiterockian); 16 and 17: Oistodus
(Ordovician); 18: Phragmodus (Ordovician); 19: Plectodina (Late Ordovician); 20: Pterospathodus (Early Silurian); 21:
Polyplacognathus (Ordovician); 22: Rhipidognathus (Late Ordovician); 23–34: Protopanderodontida (Ordovician): this
includes “uni- to multi-membrate skeletal apparatuses composed of longitudinally striated, laterally unfurrowed coniform
elements” (Sweet 1988); 23–26: Apparatus of Protopanderotus (Ordovician); 27: Belodina (Early Middle Ordovician); 28:
Clavohamulus (Early Ordovician); 29: Cornuodus (Ordovician); 30: Dapsilodus (Ordovician–Silurian); 31: Parapan-
derodus (Early Ordovician); 32: Strachanognathus (Late Ordovician); 33: Drepanoistodus (Ordovician); 34: Oneotodus
(Ordovician); 35–45: Prioniodinida: “highly distinctive extensiform digyrate elements in the two P positions. Dentiles are
characterically discreet and peg-like; white matter is commonly of minor amount” (Sweet 1988); 35–41: Apparatus plan of
Periodon (Early Ordovician); 42: Erraticodon (Ordovician) is the oldest Prioniodinida member and appeared the late Early
Ordovician and became cosmopolitan; 43: Ellisonia (Atokan; Pennsylvanian, Late Carboniferous); 44: Oulodus (Late
Ordovician); 45: Bactrognathus (Mississippian, Early Carboniferous); 46: Merrillina (Permian); 47–56: Ozarkodinida
(Late Middle Ordovician–Earliest Triassic) “P positions occupied by carminate and angulate pectiniform elements or their
platformed analogues” (Sweet 1988); 47: Hindeodus (Early Mississippian; Kinderhookian); 48: Cavusgnathus (Early
Mississippian, Carboniferous); 49: Siphonodella (Late Devonian–Mississippian, Early Carboniferous); 50: Gnathodus
(Mississippian, Early Carboniferous); 51: Indiognathodus (Late Pennsylvanian, Carboniferous); 52: Kockelella (Early
Silurian); 53: Mestognathus (Mississippian, Early Carboniferous); 54: Palmatolepis (Late Devonian); Polygnathus (Early
Devonian to Early Carboniferous); 56: Sweetgnathus (Carboniferous–Permian)

3 Classification

The classification followed here is after Sweet
(1988) who recognized the following orders:
Proconodontida, Prioniodontida, Protopan-
derodontida, Prioniodinida, Panderodontida, and
Ozarkodinida (Fig. 9). Order Proconodontida is
best represented by the Late Cambrian Procon-
odontus (Fig. 9(1)). The skeletal apparatus is
composed of large, deeply excavated, smooth,
non-geniculate coniform elements (Fig. 9(1–7)).
Order Prioniodontida (Fig. 9(8–22)) possess
pastinate coniform or pectiniform elements or
their platformed equivalents (Sweet 1988). Order
Protopanderodontida (Fig. 9(23–34)) is best
exemplified by Protopanderodus (Fig. 9(23–26))
that lack white matter (i.e. hyaline) and have
non-geniculate coniform elements with longitu-
dinal striations. Order Prioniodinida (Fig. 9(35–
46)) is best represented by Periodon (Fig. 9(35–
41)). Order Panderodontida is similar to Pro-
topanderodontida but contains laterally furrowed
3 Classification 111

Fig. 10 Panderodus gracilis (Branson and Mehl) (modified from Sweet 1988)
112 5 Conodonts

Fig. 11 Representative conodont species (see also Fig. 9)

3 Classification
elements and is represented by Panderodus
(Fig. 10). Order Ozarkodinida (Fig. 9(47–56)) is
best known by genus Ozarkodina.
4 Geological History
The most primitive conodonts are single cones
that dominated during the Early Ordovician with
a peak in the Arenigian (late Early Ordovician);
the first platform-type conodonts occurred
around this time, as well. The earliest conodonts
are known from the Precambrian, but more
commonly in the Late Cambrian, with increased
diversity in the Ordovician and Devonian. Con-
odont diversity and abundance declined in the
Silurian. During the Early and Mid-Devonian
diversity gradually increased, reaching an acme
in the late Devonian. In the Early Carboniferous
conodonts remained abundant and widespread
but diversity decreased during the late Car-
boniferous. The conodonts survived the
Permo-Triassic boundary extinctions but became
extinct by late Triassic, coinciding with the
diversification of dinoflagellates and the first
appearance of calcareous nannofossils.
Representative conodont species, through
time, are illustrated in Fig. 11.
References
Agematsu, S., Uesugi, K., Sano, H., & Sashida, K.
(2017). Reconstruction of the multielement apparatus
of the earliest Triassic conodont, Hindeodus parvus,
using synchrotron radiation X-ray micro-tomography.
Journal of Paleontology, 91, 1220–1227.
Aldridge, R. J., Briggs, D. E. G., Clarkson, E. N. K., &
Smith, M. P. (1986). The affinities of conodonts—
New evidence from the Carboniferous of Edinburgh,
Scotland. Lethaia, 19, 279–291.
Aldridge, R. J., Briggs, D. E. G., Smith, M. P., Clarkson,
E. N. K., & Clark, N. D. L. (1993). The anatomy of
conodonts. Philosophical Transactions of the Royal
Society of London. Series B: Biological Sciences, 340,
405–421.
Aldridge, R. J., Murdock, J. E., Gabbott, S. E., & Theron,
J. N. (2013). A 17-element conodont apparatus from
the Soom Shale Lagerstätte (Upper Ordovician). South
Africa: Palaeontology, 56, 261–276.
113
Aldridge, R. J., Smith, M. P., Norby, R. D., & Briggs, D.
E. G. (1987). The architecture and function of
Carboniferous polygnathacean conodont apparatuses.
In R. J. Aldridge (Ed.), Palaeobiology of conodonts
(pp. 63–76). Chichester: Ellis Horwood.
Aldridge, R. J., & Theron, J. N. (1993). Conodonts with
preserved soft tissue from a new Upper Ordovician
Konservat-Lagetstätte. Journal of Micropalaeontol-
ogy, 12, 113–117.
Armstrong, H. A., & Smith, C. J. (2001). Growth patterns
in euconodont crown enamel: Implications for life
history and mode of life reconstruction in the earliest
vertebrates. Proceedings of the Royal Society, Series
B, 268, 815–820.
Briggs, D. E. G., Clarkson, E. N. K., & Aldridge, R. J.
(1983). The conodont animal. Lethaia, 16, 1–14.
Donoghue, P. C. J., Forey, P. L., & Aldridge, R. J. (2000).
Conodont affinity and chordate phylogeny. Biological
Reviews, 75, 191–251.
Donoghue, P. C. J., Purnell, M. A., Aldridge, R. J., &
Zhang, S. (2008). The interrelationships of ‘complex’
conodonts (vertebrata). Journal of Systematic
Palaeontology, 6(2), 119–153.
Donoghue, P. C. J., & Sansom, I. J. (2002). Origin and
early evolution of vertebrate skeletonization. Micro-
scopy Research and Technique, 59, 352–372.
Dzik, J. (1991). Evolution of oral apparatuses in the
conodont chordates. Acta Palaeontologica Polonica,
36(3), 265–323.
Epstein, A. G., Epstein, J. B., & Harris, L. D. (1977).
Conodont color alteration-an index to organic meta-
morphism. Geological survey professional paper 995,
1–27.
Gabbott, S. E., Aldridge, R. J., & Theron, J. N. (1995). A
giant conodont with preserved muscle tissue from the
Upper Ordovician of South Africa. Nature, 374, 800–
803.
Goudemand, N., Orchard, M. J., Tafforeau, P., Urdy, S.,
Brühwiler, T., Brayard, A., et al. (2012). Early
Triassic conodont clusters from South China: Revision
of the architecture of the 15 element apparatuses of the
superfamily Gondolelloidea. Palaeontology, 55,
1021–1034.
Goudemand, N., Orchard, M. J., Urdy, S., Bucher, H., &
Tafforeau, P. (2011). Synchrotron-aided reconstruc-
tion of the conodont feeding apparatus and implica-
tions for the mouth of the first vertebrates.
Proceedings of the National Academy of Sciences,
USA, 108, 8720–8724.
Holmden, C., Creaser, R. A., Muehlenbachs, K.,
Bergström, S. M., & Leslie, S. A. (1996). Isotopic
and elemental systematics of Sr and Nd in 454 Ma
biogenic apatites: Implications for paleoseawater
studies. Earth and Planetary Science Letters, 142,
425–437.
Huang, J.-Y., Martínez-Pérez, C., Hu, S.-X., Donoghue,
P. C.J., Zhang, Q.-Y., Zhou, C.-Y., Wen, W., Benton,
M. J., Luo, M., Yao, H.-Z., & Zhang, K.-X. (2018).
Middle Triassic conodont apparatus architecture
revealed by synchrotron X-ray microtomography.
114
Palaeoworld. https://doi.org/10.1016/j.palwor.2018.
08.003.
Jones, D. O., Evans, A. R., Rayfield, E. J., Siu, K. K., &
Donoghue, P. C. J. (2012a). Testing micro structural
adaptation in the earliest dental tools. Biology Letters,
8, 952–955.
Jones, D. O., Evans, A. R., Rayfield, E. J., Siu, K. K., &
Donoghue, P. C. J. (2012b). The sharpest tools in the
box? Quantitative analysis of conodont element
functional morphology. Proceedings of the Royal
Society B: Biological Sciences, 279, 2849–2854.
Liu, H. P., Bergström, S. M., Witzke, B. J., Briggs, D. E.
G., McKay, R. M., & Ferretti, A. (2017). Exception-
ally preserved conodont apparatuses with giant
elements from the Middle Ordovician Winneshiek
Konservat-Lagerstätte, Iowa, USA. Journal of Paleon-
tology, 91(3), 493–511.
Marshall, C. P., Mar, G. L., Nicoll, R. S., & Wilson, M.
A. (2001). Organic geochemistry of artificially
matured conodonts. Organic Geochemistry, 32,
1055–1071.
Martínez-Pérez, C., Plasencia, P., Jones, D., Kolar-
Jurkovšek, T., Sha, J., Botella, H., et al. (2014). There
is no general model for occlusal kinematics in
conodonts. Lethaia, 47, 547–555.
Martínez-Pérez, C., Rayfield, E. J., Botella, H., &
Donoghue, P. C. J. (2016). Translating taxonomy into
the evolution of conodont feeding ecology. Geology,
44, 247–250.
Martínez-Pérez, C., Rayfield, E. J., Purnell, M. A., &
Donoghue, P. C. J. (2014). Finite element, occlusal,
microwear and microstructural analyses indicate that
conodont microstructure is adapted to dental function.
Palaeontology, 57(5), 1059–1066.
Müller, K. J., & Nogami, Y. (1971). Über den Feinbau der
Conodonten. Memoirs of the Faculty of Sciences of the
Kyoto University, Series of Geology and Mineralogy, 38,
1–88.
Mikulic, D. G., Briggs, D. E. G., & Kluessendorf, J.
(1985). A Silurian soft-bodied biota. Science, 228,
715–717.
Murdock, D. J. E., Rayfield, E. J., & Donoghue, P. C. J.
(2014). Functional adaptation underpinned the evolu-
tionary assembly of the earliest vertebrate skeleton.
Evolution and Development, 16, 354–361.
Murdock, D. J. E., Sansom, I. J., & Donoghue, P. C. J.
(2013). Cutting the first ‘teeth’—A new approach to
functional analysis of conodont elements. Proceedings
of the Royal Society B: Biological Sciences, 280,
20131524.
Orchard, M. J. (2005). Multielement conodont appara-
tuses of Triassic Gondolelloidea. Special Papers in
Palaeontology Series, 73, 73–101.
Pell, J., Russell, J. K., & Zhang, S. (2015). Kimberlite
emplacement temperatures from conodont geothermome-
try. Earth and Planetary Science Letters, 411, 131–141.
Purnell, M. A. (1993). The Kladognathus apparatus
(Conodonta, Carboniferous): Homologies with ozar-
kodinids and the prioniodinid Bauplan. Journal of
Paleontology, 67, 875–882.
5 Conodonts
Purnell, M. A. (1994). Skeletal ontogeny and feeding
mechanisms in conodonts. Lethaia, 27, 129–138.
Purnell, M. A. (1995). Large eyes and vision in
conodonts. Lethaia, 28, 187–188.
Purnell, M. A., & Donoghue, P. C. J. (1997). Architecture
and functional morphology of the skeletal apparatus of
ozarkodinid conodonts. Philosophical Transactions of
the Royal Society of London. Series B, 352, 1545–
1564.
Purnell, M. A., & Donoghue, P. C. J. (1998). Skeletal
architecture, homologies and taphonomy of ozarkodi-
nid conodonts. Palaeontology, 41, 57–102.
Purnell, M. A., Donoghue, P. C. J., & Aldridge, R. J.
(2000). Orientation and anatomical notation in con-
odonts. Journal of Paleontology, 74, 113–122.
Purnell, M. A., & Jones, D. (2012). Quantitative analysis
of conodont tooth wear and damage as a test of
ecological and functional hypotheses. Paleobiology,
38(4), 605–626.
Purnell, M. A., & von Bitter, P. H. (1992). Blade-shaped
conodont elements functioned as cutting teeth. Nature,
359, 629–631.
Rejebian, V. A., Harris, A. G., & Huebner, J. S. (1987).
Conodont color and textural alteration: An index to
regional metamorphism, contact metamorphism, and
hydrothermal alteration. Geological Society of Amer-
ica Bulletin, 99, 471–479.
Rhodes, F. H. T., Austin, R. L., & Druce, E. C. (1969).
British Avonian Carboniferous conodont faunas, and
their value in local and intercontinental correlation.
Bulletin of the British Museum (Natural History)
Geology, (Suppl, 5), 1–313.
Sansom, I. J., Smith, M. P., Armstrong, H. A., & Smith,
M. M. (1992). Presence of the earliest vertebrate hard
tissues in conodonts. Science, 256, 1308–1311.
Smith, M. P., Briggs, D. E. G., & Aldridge, R. J. (1987).
A conodont animal from the lower Silurian of
Wisconsin, U.S.A., and the apparatus architecture of
panderodontid conodonts. In R. J. Aldridge (Ed.),
Palaeobiology of conodonts (pp. 91–104). Chichester:
Ellis Horwood.
Suttner, T. J., Kido, E., & Briguglio, A. (2017). A new
icriodontid conodont cluster with specific mesowear
supports an alternative apparatus motion model for
Icriodontidae. Journal of Systematic Palaeontology,
16, 909–926.
Sweet, W. C. (1988). The conodonta: Morphology,
taxonomy, paleoecology, and evolutionary history of
a long-extinct animal phylum. Oxford: Clarendon
Press.
Sweet, W. C., & Donoghue, P. C. J. (2001). Conodonts:
Past, present, future. Journal of Paleontology, 75(6),
1174–1184.
Trotter, J. A., Fitz, J. D., Kokkonen, G. H., & Barnes, C.
R. (2007). New insights into the ultrastructure,
permeability, and integrity of conodont apatite deter-
mined by transmission electron microscopy. Lethaia,
40, 97–110.
Trotter, J. A., Korsch, M. J., Nicoll, R. S., & Whitford, D.
J. (1999). Sr isotopic variation in single conodont
References
elements: Implications for defining the Sr seawater
curve. Bollettino della Societa Paleontologica Itali-
ana, 37(2–3), 507–514.
Voldman, G. G., Albanesi, G. L., & Ramos, V. A. (2009).
Ordovician metamorphic event in the carbonate plat-
form of the Argentine Precordillera: Implications for
the geotectonic evolution of the proto-Andean margin
of Gondwana. Geology, 37, 311–314.
Voldman, G. G., Bustos-Marún, R. A., & Albanesi, G. L.
(2010). Calculation of the conodont Color Alteration
Index (CAI) for complex thermal histories. Interna-
tional Journal of Coal Geology, 82(1–2), 45–50.
Wenzel, B., Lécuyer, C., & Joachimski, M. M. (2000).
Comparing oxygen isotope records of Silurian calcite
and phosphate–δ18O compositions of brachiopods and
conodonts. Geochimica et Cosmochimica Acta, 64
(11), 1859–1872.
115
Wiederer, U., Königschof, P., Feist, R., Franke, W., &
Doublier, M. P. (2002). Low grade metamorphism in
the Montagne Noire (S–France): Conodont Alteration
Index (CAI) in Paleozoic carbonates and implications
for the exhumation of hot metamorphic core complex.
Schweizerische Mineralogische und Petrographische
Mitteilungen, 82, 393–407.
Zhang, S., & Barnes, C. R. (2007). Late Ordovician-early
Silurian conodont biostratigraphy and thermal matur-
ity, Hudson Bay Basin. Bulletin of Canadian Petro-
leum Geology, 55, 179–216.
Zhang, M., Jiang, H., Purnell, M. A., & Lai, X. (2017).
Testing hypotheses of element loss and instability in
the apparatus composition of complex conodonts:
Articulated skeletons of Hindeodus. Palaeontology,
60, 595–608.
 Radiolarians
1 Introduction
Radiolarians are microscopic (20–300 µm) mar-
ine organisms with an exclusive planktonic mode
of life. They are also the most abundant micro-
zoolankton within the marine realm with a very
wide distribution extending from pole to equator
and from shallow to abyssal depths (Anderson
1983; Afanasieva et al. 2005; Boltovskoy et al.
2010). Radiolarians have a continuous fossil
record since the earliest Cambrian (Fig. 1) and
are also one of the earliest groups of microor-
ganisms capable of inhabiting various regions of
the ocean (the ecological plasticity of Afanasieva
et al. 2005). The early forms had a spherical
skeleton with a radial symmetry (Braun et al.
2007; Maletz 2011) (Fig. 1). But it must be kept
in mind that the Early Cambrian material is
poorly preserved and at times fragmentary to
merit positive identifications, thus, leaving the
question of the earliest radiolarian record, still
open (see Maletz 2011 but also see Cao et al.
2014; Chang et al. 2018).
© Springer Nature India Private Limited 2020
S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_6
6
2 Morphology
The cell cytoplasm is divided into an outer ec-
toplasm and an inner endoplasm, separated by a
perforate organic membrane called the central
capsule (Fig. 2(1)). The ectoplasm secretes the
skeleton/test and bulk of it lies within it only.
The nucleus is within the endoplasm (Fig. 2).
From the central capsule, pseudopodia radiate
outwards as central rods of fibers called axopo-
dia; if they are thread-like, they are called fil-
ipodia. Calymma, a zone of frothy, gelatinous
bubbles occurs within ectoplasm (Fig. 2(1)). The
mineral skeleton lies inside the cytoplasm and is
composed of opal (98% amorphous SiO2
nH2O;
Hurd and Takahashi 1981; Petrushevskaya 1981;
Takahashi and Honjo 1981) with small amounts
(1–4%) of Mg, Ca, Al, and Na (Anderson 1983).
The mineralized skeleton consists of either radial
or tangential elements, or both. The radial ele-
ments include loose spicules, external spines or
internal bars, whereas the tangential elements
consist of a porous lattice shell of variable
117
118 6 Radiolarians

Fig. 1 Early Cambrian radiolarian records (chronostratigraphy based on Peng 2003; Geyer and Landing 2004; Babcock
and Peng 2007). 1: Braun et al. (2007), Ningqiang, Shaanxi (uppermost Precambrian or basal Cambrian); 2: Danelian and
Bengtson (2009), Australia (Nemakit/Daldynian); 3: Braun et al. (2007), Xintangwu, Zhejiang (Meishucunian); 4: Hao and
Shu (1987), Zhenba, Shaanxi (Eoredlichia Zone); 5: Obut and Iwata (2000), Gorny Altai (Botomian); 6: Won and Below
(1999), Georgina Basin, Queensland, Australia (Templetonian, Floran, Undillian); 7: White (1986), Utah (Middle
Cambrian, Ptychagnostus atavus Zone); 8: Dong et al. (1997), Paibi, Hunan Province, China (basal Hunanian); 9: Won and
Iams (2002), western Newfoundland (Franconian, Trempealeauan). Arrows marks exact ages. Gray arrows indicate
material not considered to be radiolarians. For more details see Maletz (2011), image curtsey Jörg Maletz and with
permission from Elsevier (Paleoworld)

morphology, such as spheres, spindles, and cones
(Fig. 2(2)).
Geologists are concerned with only one radi-
olarians group, the polycystines (Class Poly-
cystinea) that includes the radially symmetrical
spumellarians (Order Spumellaria Ehrenberg)
(Fig. 3) and the helmet-shaped nassellarians
(Nassellaria Ehrenberg) (Fig. 4). Both are
important because of their solid opaline skeleton
which is more resistant than those of silicoflag-
ellates and diatoms but somewhat susceptible to
dissolution as seawater is undersaturated relative
to silica.
The spumellarians possess a radial symmetry
along a central point (spherical symmetry)
(Fig. 3(2)), whereas the nassellarians has a radial
symmetry along the long axis (non-spherical,
usually bilateral symmetry) (Fig. 4(3)). Both
have a solid skeleton made of amorphous opaline
silica (SiO2
nH2O). One other minor order
(Order Phaeodarea) that has no fossil record has a
skeleton made by the admixture of organic mat-
ter (dominant constituent) and 20% opaline sil-
ica. Although the radiolarians occur as a simple
cell but some spumellarians occur as colonies of
numerous individuals each bearing its own
skeleton. These colonies are as large as 250 mm.
The radiolarian skeleton is composed of a
network of two types of elements, the elongate
elements attached at one end only called spines
or connected at both ends called bars (Fig. 3(2)).
In some polycystines, the entire skeleton is made
up of an association of simple spines called spi-
cules (Fig. 3(1)); the simplest skeleton of both
spumellarians and nassellarians is the spicule.
The spicules range in size from 100 to 400 lm
and in some rare cases, up to 2 mm. The skeletal
geometry and perforations on the capsular
membrane distinguish these two groups—
spumellarians and nassellarians.
2 Morphology 119

Fig. 2 A typical radiolarian

cell (Thalassicola). Cross
section of a spumellarian
displaying Nucleus,
Endoplasm and Ectoplasm
(1) with respect to the
concentric lattice shells and
radial spines (2) (modified
after Westphal 1976). See text
for explanation

The Spumellaria are characterized by spheri-
cal skeletal shapes (Spheres), where the radial
spines (=Main spine) extends from the surface of
the sphere (Fig. 3(2)). The skeleton generally
consists of two or more nested spheres that are
concentric and connected by radial bars (Fig. 3
(2)). The main outer shell is called the cortical
shell and the inner one is the medullary shell
(Fig. 3(2)). The innermost spherical shell is
generally very small but is characteristic of
radiolarians. The latticed shell wall is a mesh-
work of bars that forms closely spaced pores;
they are mostly hexagonal in shape (Fig. 3(3–7)).
The shape and the distribution of pores are
120
Fig. 3 Spumellarian skeletal
morphology (modified from
Haq and Boersma 1998)
consistently same in a species, hence, both
characters are of taxonomic importance. The
latticed skeleton is equally varied in shape
(Fig. 3(8–13)).
The nassellarian spicule is usually facetted
and has a distinct organization of spines (Fig. 4
(1–4); the various types of spine include A:
Apical spine; D: Dorsal spine; PL: Primary lat-
eral spine; V: Vertical spine; SL: Secondary
lateral spine). Although the spicules are similar
to those of the co-occurring sponges but the
6 Radiolarians
former is solid and does not possess a central
space. In nassellarians, the Median bar is of
taxonomic importance due to its distinctive
structure enabling categorization into various
groups (Fig. 4(1)). The arrangement of the
median bar and spines is collectively called the
collar structure that rests between the head
(cephalus) and the body (thorax) (Fig. 4(3)). The
spines divide the cephalus into two parts, the
larger eucephalic lobe (EUC) with the median
bar at its base and the smaller anicephalic lobe
2 Morphology 121

Fig. 4 Nassellarian skeletal morphology (modified from Haq and Boersma 1998)

(Fig. 4(3)). The spines may also be associated
with the Tube (Fig. 4(4)). The median bar has
axostyle supports which are downward projec-
tions (see (Fig. 4(4)). A lumber structure (Fig. 4
(3)) separates the thorax with the abdomen
(Fig. 4(3)); those lying below the abdomen are
called postabdominal segments that has an open
or a constricted aperture (Fig. 4(3)) or a peris-
tome (a basal ring; Fig. 4(6)). The peristome may
be without pores (unpored; Fig. 4(6)) or ragged
122 6 Radiolarians

(Fig. 4(7)). That on which a terminal segment 3 Geological History

rests is called feet (singular = foot) (Fig. 4(5–8));
the subterminal appendages are called wings or
shoulders (Fig. 4(8)). The cephalus (Fig. 4(3))
has varied morphological shapes, from spherical
to elongate to subcylindrical and contains pro-
cesses such as horns or spines (Fig. 4(9–11)).
The shape of shell segments (Fig. 4.3) is of
taxonomic value and occur in different shapes
(Fig. 4(12–19)). Even the pores (Fig. 4(21 and
22)) within the segment are arranged differently
and therefore are of taxonomic value, as well; the
pores also occur in different shapes (Fig. 4
(22–25)).
The Permian/Triassic crisis was a period of
marked radiolarian extinctions (of orders Albail-
lellaria and Latentifistularia), followed by the
spectacular diversification of Spumellaria and
Nassellaria in Early-Mid Triassic (Fig. 5).
Broadly, the radiolarian diversity gradually
increased from Cambrian to Jurassic, remained
stable until the Cretaceous, and thereafter,
decreased slightly (see De Wever et al. 2001,
2003) (Fig. 5). The Phanerozoic records three
major mass extinction events—at the
Permian/Triassic boundary, at the Triassic/

Fig. 5 Radiolarian diversity (Family), extinctions (Species and Genera), and polycystine diversity of radially
symmetrical spumellarians (Order Spumellaria Ehrenberg) and helmet-shaped nassellarians (Nassellaria Ehrenberg),
along with Order Albaillellaria. Modified after Anderson (1983), De Wever et al. (2001) and Afanasieva et al. (2005)
3 Geological History 123

Fig. 6 Ordovician radiolarian biostratigraphy (after Maletz 2011). 1: Basal Tremadocian, western Newfoundland,
except Archeoentactinia reedae from Won et al. (2002); 2: Protoproventocitum sp. (Won et al. 2007); 3: Maletz and
Bruton (2007); 4: Cow Head Group, St. Paul’s Inlet, western Newfoundland (Won and Iams 2002); 5: Maletz and
Bruton (2008); 6: Viola Limestone, Katian, Alabama, except Protoceratoikiscum (Noble and Webby 2009); 7: After
Noble and Webby (2009); Graptolite biostratigraphy based on Maletz (2007). Dapin. = Dapingian; Hir. = Hirnantian.
(after Maletz 2011; curtsey Jörg Maletz and with permission from Elsevier (Paleoworld)

Jurassic boundary, and at the Cretaceous/Tertiary
(K/T) boundary (Lethiers 1998; De Wever et al.
2003) (see Fig. 5).
A note on the chronostratigraphic usefulness
of the Ordovician radiolarian merits special
mention (Fig. 6) (Maletz 2011). These are
associated with well-dated graptolite faunas
(Maletz 2007) that allows very precise dating of
the radiolarian faunas and provides an oppor-
tunity to establish and calibrate the radiolarian
biostratigraphy with various other macrofossils
groups (Maletz 2011). An updated Permian and
Triassic radiolarian biostratigraphy are provided
in Tables 1 and 2 and representative radiolarian
species through time are illustrated in Figs. 7, 8,
9, and 10.
124 6 Radiolarians

Table 1 Permian radiolarian markers

3 Geological History 125

Table 2 Triassic radiolarian markers

126 6 Radiolarians

Fig. 7 Representative Albaillellarian (1) and Spumellarian species. 1: Albaillellarian. 1: Albaillella Deflandre
(Carboniferous); 2–14: Spumellarian. 2: Haplentactina (Devonian); 3 and 4: Paronaella Pessagno (Carboniferous–
Cretaceous); 5 and 6: Alievium Pessagno (Triassic); 7: Capnuchosphera De Wever (Triassic); 8: Cenosphera (Eocene–
Recent); 9: Ommatartus Haeckel (Miocene–Recent); 10: Cannartus Haeckel (Oligocene–Miocene); 11: Collosphera
Müller (Miocene–Recent); 12: Actinomma Haeckel (Miocene–Recent);13: Amphirhoplanum Haeckel (Pliocene–Recent);
14: Saturnalis Haeckel (Pleistocene–Recent); 15–17: Nassellarian. 15–17: Dorcadospyris Haeckel (Eocene–Miocene);
15: Adult form; 16: Front view with cephalic constriction; 17: View from below
3 Geological History 127

Fig. 8 Representative Spumellarian species. 1: Heliosphaera clavate Hinde (Devonian); 2: Theodiscus hastus Hinde
(Devonian); 3: Cenosphaera affinis Hinde (Devonian); 4: Lithocampe andersoni Campbell and Clarke (Cretaceous); 5:
Saturnalis lateralis Campbell and Clarke (Cretaceous); 6: Hexadoridium magnificum Campbell and Clarke
(Cretaceous); 7: Lithomelissa armata Campbell and Clarke (Cretaceous); 8: Trochodiscus nicholsoni Riist
(Carboniferous); 9: Porodiscus (Trematodiscus) charlestonensis Campbell and Clarke (Eocene); 10: Calocyclas
semipolita Campbell and Clarke (Eocene); 11: Tripilidium clavipes advena Campbell and Clarke (Eocene); 12:
Roplacodictyum malagaense Campbell and Clarke (Miocene); 13: Saturolonche aculeata Campbell and Clarke
(Miocene); 14: Pterocorys splendens Campbell and Clarke (Miocene)
128 6 Radiolarians
 3 Geological History 129

b Fig. 9 Representative Nassellarian species. 1 and 2: Mirifusus Pessagno (Jurassic–Cretaceous); 3: Saturniforma

Pessagno (Jurassic–Cretaceous); 4–6: Dictyomitra Zittel (Cretaceous); 5: Apical section; 6: Transverse section at the
level of a diaphragm; 7: Holocryptocapsa Tan Sin Hok (Cretaceous); Axial section; 8 and 9: Theocampe Haeckel
(Cretaceous–Recent), 8: Axial section; 9: Apical view of Cephalis; 10: Calocyclas Ehrenberg (Eocene); 11:
Pterocanium Ehrenberg (Eocene–Recent); 12: Anthocyrtidium Haeckel (Oligocene–Recent); 13–15: Giraffospyris
Haeckel (Eocene–Recent); 16: Zygocircus Butschli (Eocene–Recent); 17: Lychnocanoma Haeckel (Eocene–Recent);
18: Callimitra Haeckel (Eocene–Recent); 19: Phormostichoastus Haeckel (Oligocene–Recent); 20: Stichocorys Haeckel
(Miocene–Pliocene); 21: Cyrtocapsa Haeckel (Oligocene–Miocene); 22: Peripyramis Haeckel (Oligocene–Recent); 23:
Botryopyle Haeckel (Eocene–Recent); 24: Corocalyptra Haeckel (Pleistocene–Recent)

Fig. 10 Representative Nassellarian species. 1: Anthocyrtis mespihis Ehrenberg (Tertiary); 2: Lychocanium lucerna
Ehrenberg (Tertiary); 3: Dictyomitra mespilus Ehrenberg (Tertiary); 4: Eucyrtidium elegans Ehrenberg (Tertiary); 5:
Pterocodon compana Ehrenberg (Tertiary); 6: Podocyrtis schomburgi Ehrenberg (Tertiary); 7: Cyrtocalpis amphora
Haeck (Recent); 8: Bothryocampe hexathalamia Haeck (Recent); 9: Petalospyris foreolata Ehrenberg (Tertiary)
130
References
Afanasieva, M. S., Amon, E. O., Agarkov, Yu V, &
Boltovskoy, D. S. (2005). Radiolarians in the geolog-
ical record. Paleontological Journal, 39(3), 135–392.
Anderson, O. R. (1983). Radiolaria. New York, NY:
Springer, 365p.
Babcock, L. E., & Peng, S. (2007). Cambrian chronos-
tratigraphy: current state and future plans. Palaeo-
geography, Palaeoclimatology, Palaeoecology, 254,
62–66.
Boltovskoy, D., Kling, S. A., Takahashi, K., & Bjorklund,
K. (2010). World atlas of distribution of living
radiolaria. Palaeontologia Electronica, 13(3), 1–230.
Braun, A., Chen, J., Waloszek, D., & Maas, A. (2007).
First Early Cambrian Radiolaria. Geological Society,
London, Special Publications 286(1), 143–149.
Cao, W. C., Feng, Q. L., Feng, F. B., & Ling, W. (2014).
Radiolarian Kalimnasphaera from the Cambrian Shui-
jingtuo Formation in South China. Marine Micropa-
leontology, 110(2), 3–7.
Chang, S., Feng, Q., & Zhang, L. (2018). New siliceous
microfossils from the Terreneuvian Yanjiahe forma-
tion, South China: The possible earliest radiolarian
fossil record. Journal of Earth Science, 29(4), 912–
919.
De Wever, P., Dumitrica, P., Caulet, J. P., Nigrini, C., &
Caridroit, M. (2001). Radiolarians in the sedimentary
record (p. 533). Amsterdam: Gordon and Breach
Science Publishers.
De Wever, P., O’Dogherty, L., Caridroit, M., Dumitrica,
P., Guex, J., Nigrini, C., et al. (2003). Diversity of
radiolarian families through time. Bulletin de la
Société géologique de France, 174(5), 453–469.
Dong, X., Knoll, A. H., & Lipps, J. R. (1997). Late
Cambrian Radiolaria from Hunan, China. Journal of
Paleontology, 71(5), 753–758.
Geyer, G., & Landing, E. (2004). A unified lower–middle
Cambrian chronostratigraphy for West Gondwana.
Acta Geologica Polonica, 54, 179–218.
Hao, Y. C., & Shu, D. G. (1987). The oldest
well-preserved Phaecodaria (Radiolaria) from south-
ern Shaanxi. Geoscience, 1, 301–310.
Haq, B. U., & Boersma, A. (1998). Introduction to marine
micropaleontology (2nd ed., pp. 376). Elsevier.
Hurd, D. C., & Takahashi, K. (1983). On the estimation of
minimum mechanical loss during an in situ biogenic
silica dissolution experiment. Marine Micropaleontol-
ogy, 7, 441–447.
Lethiers, F. (1998). Evolution de la biosphère et événe-
ments géologiques. (p. 321). Paris: Gordon and
Breach.
6 Radiolarians
Maletz, J. (2011). Radiolarian skeletal structures and
biostratigraphy in the early Palaeozoic (Cambrian–
Ordovician). Palaeoworld, 20, 116–133.
Maletz, J. (2007). The Ordovician Beothuka terranova
(Radiolaria) assemblage in western Newfoundland.
Paläontologische Zeitschrift, 81(1), 71–82.
Maletz, J., & Bruton, D. L. (2007). Lower Ordovician
(Chewtonian to Castlemainian) radiolarians of Spits-
bergen. Journal of Systematic Palaeontology, 4, 245–
288.
Maletz, J., & Bruton, D. L. (2008). The middle Ordovi-
cian Proventocitum procerulum radiolarian assem-
blage of Spitsbergen and its biostratigraphic
correlation. Palaeontology, 51(5), 1181–1200.
Noble, P. J., & Webby, B. D. (2009). Katian (Ordovician)
radiolarians from the Malongulli Formation, New
South Wales, Australia, a reexamination. Journal of
Paleontology, 83, 548–561.
Obut, O. T., & Iwata, K. (2000). Lower Cambrian
Radiolaria from Gorny Altai (southern West Siberia).
Lower Cambrian Radiolaria from the Gorny Altai
(southern West Siberia). Journal of Geology and
Geophysics 41 (Suppl.News of Paleontology and
Stratigraphy Nos. 2–3), 33–37.
Peng, S. (2003). Chronostratigraphic subdivision of the
Cambrian in China. Geologica Acta, 1, 135–144.
Petrushevskaya, M. G. (1981). Radiolarians of the Order
Nassellaria of the World Ocean. Leningrad: Nauka.
(in Russian).
Takahashi, K., & Honjo, S. (1981). Vertical flux of
Radiolaria: A taxon-quantitative sediment trap study
from the western Tropical Atlantic. Micropaleontol-
ogy, 27, 140–190.
Westphal, A. (1976). Protozoa (325p). Glasgow: Blackie.
White, R. D. (1986). Cambrian radiolaria from Utah.
Journal of Paleontology, 60, 778–780.
Won, M. Z., & Below, R. (1999). Cambrian Radiolaria
from the Georgina Basin, Queensland, Australia.
Micropaleontology, 45, 325–363.
Won, M. Z., Iams, W. J., & Reed, K. M. (2007). Late
Tremadocian radiolarian faunas and biostratigraphy of
the Cow Head Group, western Newfoundland,
Canada. Journal of the Korean Earth Science Society,
28(4), 497–540.
Won, M. Z., Blodgett, R. B., & Nestor, V. (2002).
Llandoverian (Early Silurian) radiolarians from the
Road River Formation of East-Central Alaska and the
new family Haplotaeniatumidae. Journal of Paleon-
tology, 76(6), 941–964.
Won, M. Z., & Iams, W. J. (2002). Late Cambrian
radiolarian faunas and biostratigraphy of the Cow
Head Group, western Newfoundland. Journal of
Paleontology, 76, 1–33.
 Marine Diatoms
7

25% of the oxygen release and carbon fixation in

1 Introduction
the world. Diatoms alone account for around
40% of the phytoplankton on Earth (= *20–
Diatoms are very small (10–200 lm) plant-like
25% global net primary production), more than
organisms (phytoplanktons) that float. Of all the
the primary production of all tropical rainforests
algae phyla, diatoms are the most numerous,
combined! They also play an important role in
with *10,000 known species, inhabiting both
the silica cycle and contribute to other biogeo-
marine (*75%) and fresh water and other
chemical cycles, especially in carbon fixation
habitats (*25%). Like plants, the diatoms pho-
(carbon cycle) by converting CO2 into biomass
tosynthesize sunlight (autotrophs), but unlike
(Smol and Stoermer 2010; see also Lane et al.
terrestrial plants their light absorbing pigments
2005; Falkowski and Raven 2007; Leblanc et al.
are brown, not green. As they are light-dependent
2018). Their contribution to global productivity
(sunlight for photosynthesis), they generally live
and nutrient cycling is significant (Rau et al.
within the photic zone (i.e. depths <200 m) in
1996; Falkowski et al. 1998; Mann 1999; Sme-
the oceans and fresh water bodies. Some diatom
tacek 1999; Boyd et al. 2000; Leblanc et al.
species float in the water currents near the sur-
2018). Additionally, individual species are often
face, whereas others attach themselves to larger
restricted to specific ecological conditions, thus,
floating objects or to the seafloor. The individual
making them excellent proxies for environmental
diatoms are solitary or sometimes form colonies.
(paleo) analyzes. Also, the assemblages of dia-
They are mostly non-motile but, some benthic
tom species are limited to a particular geo-
diatoms have a specialized raphe system that
graphical region, thus, reflecting the prevailing
secretes mucilage to enable them to attach or
characteristic of physical, chemical, and biolog-
glide along a surface. They are also known to
ical conditions (Stoermer and Smol 1999; Smol
form biofilms, i.e., layers of tightly attached cells
and Stoermer 2010).
of microorganisms. When diatoms die, they
slowly sink to the seabed. The buildup of trillions
of these dead shells forms a crumbly white sed-
iment called diatomite (Diatomaceous earth),
2 Morphology
which is used in manufacturing pool filters, mild
The results from ultrastructural, biochemical,
abrasives, insulators, mineral fillers, mechanical
molecular phylogenetic, and comparative geno-
insecticides, anticaking agents, and including the
mic approaches have revealed that diatoms
everyday toothpaste (see Harwood 1999).
belong to the heterokontophytes, a group of algae
Diatoms are also a major phytoplankton group
with golden-brown to brown chloroplasts
within aquatic ecosystem and account for 20–

© Springer Nature India Private Limited 2020 131

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_7
132
originating from a red algal endosymbiont. These
single-celled, eukaryotic organisms, have their
genetic information sequestered into subcellular
compartments called Nuclei. This character dis-
tinguishes this group from other single-celled
photosynthetic aquatic organisms, like the
blue-green algae that do not possess nuclei and
are thus, more closely related to bacteria. Dia-
toms are distinct as they secrete complex outer
cell walls called Frustule (Fig. 1(1–3)). The
Frustule, makes up *20% (can be as low
as <1% in slightly silicified species) of the dia-
tom dry weight, and consists of two overlapping
thecae, each in turn, consisting of a valve and a
number of hoop-like or segmental girdle bands,
fitting together like a pillbox (Fig. 1(1)). The
larger valve is called the epitheca and the smaller
valve is called the hypotheca (Fig. 1(1–3)). The
overlap area is called a Girdle (a circular band
frequently noted in the fossil record, but of no
stratigraphic relevance) (Fig. 1(2 and 3)) and the
longitudinal valve slit is called the Raphe (Fig. 1
(4 and 5)). Both valves possess a valve plate and
a cingulum (Fig. 1(4)). The singular algal diatom
cell lives protected inside the frustule halves like
a pair of shoes snuggled within a shoe box.
Frustule is bilamellar, very ornate, and with
intricate pore patterns. The pores (also called
Punctae) perforate the frustule thereby allowing
gases, nutrients, and metabolic waste products to
be exchanged between the watery environment
and the algal cell. Not just pores, the frustule is
also riddled with spikes, spines, and other fea-
tures that are of taxonomic significance. The
brittle frustule cell wall is made of opaline silica
(glass; SiO2) and pectin. The glass frustule pro-
vides diatom, the structure, and a mechanism for
defense.
The frustules either exhibit bilateral (Fig. 1(4
and 5)) or radial symmetry (Fig. 1(6)) (for details
see Round et al. 1990). The symmetry is defined
by three axes: the apical axis (parallel to the
length of the valve), the transapical axis (at right
angles to the apical axis), and the pervalvar axis
(perpendicular to the center of the valve face)
7 Marine Diatoms
(Fig. 1(7)). In bilaterally symmetric forms, the
halves are mirror images of each another; these
are elongated forms (Fig. 1(4 and 5)). In them,
the frustules are elliptical or rectangular in valve
view (Fig. 1(4)), and bilaterally symmetrical
about a central line. The latter (central line) is a
longitudinal groove in the middle of the valve
face (median, Fig. 1(8 and 9); can also be mar-
ginal, Fig. 1(10 and 11)) and is called a raphe
(=elongate fissure in a valve). Some diatoms do
not possess this groove but only a silicified clear
area of punctae, it is then called a pseudoraphe
(in some literature it is also called a rapheless
valve) (Fig. 1(12 and 13)). The central nodule
divides the raphe into two; those at the extremi-
ties are called polar nodules (Fig. 1(4)). The
central nodule is a thickly silicified area located
between the ends of the proximal raphe. The
central nodule is the place for the first silica
deposition during valve formation. The raphe or
pseudoraphe are used for taxonomic subdivision
within the pennate diatoms (Fig. 1(4 and 5)); the
suborder Araphidineae only has a pseudoraphe.
The radially symmetric frustules (Fig. 1(6)) have
numerous mirror images. No matter from where
a shell is divided, each half is always a mirror
image of the other. These radially symmetric
diatoms are round and flattened (Fig. 1(6)). The
combination of this symmetry and the perfora-
tion patterns of diatom frustules are useful in
identifying different species. Because they are
composed of silica, a largely inert material, dia-
tom frustules remain well preserved over long
periods of time within geologic sediments and
are known from fossil records since the Early
Albian (Early Cretaceous) (Harwood and
Gersonde 1990; Sims et al. 2006).
The valves of most diatoms are covered with a
honeycomb structure called areolae that are
vertical-sided chambers within the valve wall
(Fig. 1(14)). Their geometry and spacing are
diagnostic features for species identification (i.e.
the pores are species-specific). Sometimes, a finely
perforated plate is also noted that may cover the
areolae and it is then called the sieve plate.
2 Morphology 133

Fig. 1 Diatom terminology. 1–3: Frustule terminology; 4–6: Examples of two major orders of diatom based on frustule
shape—elliptical (pennate) (4 and 5) or circular (centric) (6) in valve view; these make up the two orders of diatoms
(Pennales and Centrales, respectively). 4 and 5: Pinnate diatom; 4: Valve view; 5: Connective view. 6: Centric diatom.
7: Symmetry axes. 8–11: Diatom species showing Raphe. 12 and 13: Diatom species showing pseudoraphe. 14:
Structural details of the cell wall. For further explanation, see text
134
3 Terminology
Additional terms used in describing a diatom are
enumerated below and illustrated in Fig. 1.
3.1 Gross Morphology
3.1.1 Apical Axis
It is the long axis of the valve face and passes
through the apices following the midline of the
valve (Fig. 1(7)). It may be curved, as in
Cymbella.
3.1.2 Apical Plane
A plane perpendicular to the transapical axis
(Fig. 1(7)).
3.1.3 Band or Segment
A single element of the girdle (Fig. 1(3 and 5)).
3.1.4 Cingulum
A portion of the girdle that is associated with a
single valve (Fig. 1(5)).
3.1.5 Girdle
A part of a frustule between epi- and hypo-valves
consisting of epi- and hypo-cingula (Fig. 1(2 and
3)).
3.1.6 Septum
A sheet or ridge in the valvar plane projecting
from a girdle band into the interior of the frus-
tule, often with several openings.
3.1.7 Valve View
A frustule seen from top or bottom (Fig. 1(4)).
3.2 Cell Wall Structure
3.2.1 Areola
A regularly repeated perforation through the
valve wall. These are pores in a row that form a
stria on a valve (plural is areolae) (Fig. 1(14)).
7 Marine Diatoms
3.2.2 Cribrum
A velum perforated by regularly arranged pores
or is reticulate (Fig. 1(14)).
3.2.3 Foramen
The passage through the constriction at the sur-
face opposite the velum; a large opening in one
wall of an areola (Fig. 1(14)).
3.2.4 Velum
A thin perforated layer of silica occluding an
areola (pores 30 lm or more in diameter) (Fig. 1
(14)).
4 Geological History
Diatoms have a long record based on fossil silica
frustules and on biochemical markers (Gersonde and
Harwood 1990; Sinninge Damststé et al., 2004;
Sims et al. 2006). The earliest diatoms have been
recorded from the Early Albian (Early Cretaceous)
from the Weddell Sea, Antarctica as also from
Korea, Northern Germany, Australia, and the
Antarctic margin (Harwood and Gersonde 1990;
Gersonde and Harwood 1990; Nikolaev and
Harwood 2000; Sims et al. 2006) (Figs. 2 and 3).
However, few suspect occurrences of diatoms or
diatom relatives have also been reported from the
Precambrian and Triassic. Older records are doubted
as silica recrystallizes under pressure; hence any
older diatom fossils are likely to be destroyed (see
Kwiecinska 2000; Sieminska 2000). Abundant well
preserved forms first appeared in the Campanian
and first freshwater forms appeared in the Pale-
ocene, but only diversified until the Oligocene. The
scarcity of preCretaceous diatom fossils could lar-
gely be a function or combination of: (a): the early
diatoms probably were rare, (b) lived in habitats
adverse to fossilization, (c) their frustules were
lightly silicified, and at least for the Paleogene
(Cenozoic) diatoms, (d) in coastal habitats, the
frustules had a high probability of being dissolved,
pyritized, or crushed and diagenetically transformed
from biogenic opal into chert or minerals, such as
4 Geological History 135

Fig. 2 Diatom classification (after Simonsen 1979)

136 7 Marine Diatoms

Fig. 3 Species diversity, fossil record, and biomarkers. The Centric diatoms are either Radial (with petri dish-shaped
valves) or Multipolar (bi-, tri-, etc., referring to the polarity of shape) with a radial pore organization; their cell form is
elongate, triangular or star-like, thus, exhibiting polarity. The pennate ones are elongate with a midrib from which striae
and interstriae extend at right angles. Pennate diatoms are of two types, Raphid (those with a slit-shaped process, called
a Raphe), and Araphid (that lack a raphe, but have apical pore fields) (see text and Fig. 1 for further explanation). The
fossil record is after Katz et al. (2005). The presence of biomarkers in crude oil is believed to be linked to the rise of
diatoms (see Moldowan and Jacobson 2000)

zeolites (see von Rad et al. 1977). There are prob-

ably well over ca. 105 species (Mann and Droop
1996) but “only” *10,000 species inhabit the
marine plankton (Sournia 1995), i.e., the major
portion of the diatom biodiversity is not planktonic,
but benthic.
5 Classification
Simonsen’s (1979) classification is followed
here. It divides the Centric diatoms into three
suborders, each characterized by the shape of the
cells, the polarity, and the arrangement of the
5 Classification 137

Fig. 4 Representative diatom species, through time. 1: Minidiscus trioculatus (Taylor). Along with M. comicus, these two
species are considered the smallest known centric diatoms (* 2–5 µm; Leblanc et al., 2018) of genus Minidiscus; 2:
Minidiscus comicus Takano; 3: Minidiscus chilensis Rivera; 4–6: Planktoniella blanda (Schmidt); 7: Planktoniella sol
(Wallich); 8-9: Asteromphalus sp.; 8: Valve view; 9: Girdle view; 10: Asterolampra marylandica Ehrenberg; 11 and 12:
Actinoptychus senarius (Ehrenberg); 12: Girdle view; 13: Planktoniella muriformis (Loeblich, Wight and Darley); 14 and
15: Rhizosolenia formosa Peragallo; 14: Ventral view; 15: Lateral view; 16: Rhizosolenia robusta Norman; 17: Guinardia
cylindrus Cleve; 18: Dactyliosolen phuketensis (Sundström)
138 7 Marine Diatoms

Fig. 5 Representative diatom species, through time. 1: Chaetoceros Ehrenberg; broad girdle view; 2–6: Bacteriastrum
furcatum Shadbolt; 2: Terminal parts of chain in girdle view; 3: Terminal; 4 and 5: Intercalary valves in valve view; 6:
Middle valve; 7 and 8: Nanoneis hasleae Norris; 7: Stepped chain in girdle view; 9: Bacteriastrum delicatulum Cleve;
Intercalary valve in valve view; 10: Rhaphoneis amphiceros (Ehrenberg); 11 and 12: Fragilariopsis oceanica (Cleve);
11: Ribbon with resting spores; 12: Valve showing isopolarity; 13–15: Achnanthes taeniata Grunow; 13: Chain with
resting spores and chloroplasts; 14: Valve with Raphe; 15: Rapheless valve
5 Classification 139

Fig. 6 Representative Mesozoic diatom species. 1 and 2: Coscinoconus (Cretaceous–Recent); 1: Valve view; 2:
Connective view; 3 and 4: Actinoptychus Ehrenberg (Cretaceous–Recent); 3: Valve view; 4: Connective view; 5–7:
Melosira Agardh (Cretaceous–Recent); 6: Valve view; 7: Connective view; 8 and 9: Stephanopyxis Ehrenberg
(Cretaceous–Recent); 8: Valve view; 9: Connective view; 10 and 11: Triceratium Ehrenberg (Cretaceous–Recent); 10:
Valve view; 11: Connective view; 12 and 13: Hemiaulus Heiberg (Cretaceous–Recent); 12: Row of colonial cells; 13:
Diagonal view of a valve; 14: Trinacria Heiberg (Cretaceous–Oligocene); Connective view; 15–17: Biddulphia Gray
(Cretaceous–Recent); 15: Connective view; 16: Section of a valve; 18: Valve view
140 7 Marine Diatoms

Fig. 7 Representative Cenozoic diatom species. 1: Rhizosolenia Ehrenberg (Oligocene–Recent); Connective view; 2–4:
Chaetoceros Ehrenberg (Miocene–Recent); 2: Valve view with colonial form; 3: Connective view of a valve. The genus is
characterized by frustules that are very lightly silicified with long and thin spines; the latter overlap to join cells in colonies; 5:
Strangulonema Greville (Eocene–Recent); Connective view; 6: Asterolampra Ehrenberg (Eocene–Recent); Valve view; 7:
Cymbella Agardh (Miocene–Recent); Valve view. The genus is characterized by valves that are slightly to strongly asymmetric
with respect to the apical axis; 8 and 9: Denticula Kützing (Miocene–Recent); 8: Connective view; 9: Internal view of a valve.
The genus is characterized by having an eccentric raphe; 10 and 11: Pleurosigma Smith (Miocene–Recent); 10: Valve view; 11:
Connective view. The genus is characterized by sigmoid-shaped valve and raphe; 12 and 13: Gomphonema Agardh (Pliocene–
Recent); 12: Connective view; 13: Valve view. The genus is characterized by club-shaped valve outline and uniseriate striae

processes (process pattern) (see Fig. 2). The
pennate diatoms are divided into two suborders,
one including diatoms without a raphe, the other
one with a raphe (Fig. 2). The further differen-
tiation of the centric diatoms into families
follows Simonsen (1979) in gross features,
whereas the classification of the pennate diatoms
is partially from Round et al. (1990).
Representative diatom species, through time,
are illustrated in Figs. 4, 5, 6, 7 and 8.
5 Classification 141

Fig. 8 Representative Cenozoic diatom species. 1 and 2: Fragilaria Lyngbye (Eocene–Recent); 1: Valve view; 2:
Connective view. The genus is characterized by frustules that are rectangular to lanceolate in shape (girdle view) and are
joined by spines to form colonies; 3 and 4: Cocconeis Ehrenberg (Oligocene–Recent); Valve views. The genus is
characterized by having one valve with a raphe, and the other without it; the valve shape is elliptical; 5 and 6: Surirella
Turpin (Oligocene–Recent); 5: Valve view; 6: Connective view. The genus is characterized by a raphe that runs along the
entire valve margin; 7 and 8: Campylodiscus Ehrenberg (Miocene–Recent); 7: Profile view; 8: Valve view. The genus is
characterized by having a saddle-shaped valve; the raphe runs along the entire valve margin; 9 and 10: Grammatophora
Ehrenberg (Miocene–Recent); 9: Valve view; 10: Connective view; 11 and 12: Epithemia Kützing (Pliocene–Recent);
11: Valve view; 12: Connective view. The genus is characterized by having an eccentric raphe and complex areolae; 13–
15: Amphiprora (Miocene–Recent); 13: Connective view; 14: Valve view; 15: Section of a valve; 16 and 17: Eunotia
Ehrenberg (Miocene–Recent); 16: Valve view; 17: Submarginal raphe. The genus is characterized by having a very short
raphe and uniseriate striae
142
References
Boyd, P. W., Watson, A. J., Law, C. S., Abraham, E. R.,
Trull, T., Murdoch, R., et al. (2000). A mesoscale
phytoplankton bloom in the polar Southern Ocean
stimulated by iron fertilization. Nature, 407, 695–702.
Falkowski, P. G., & Raven, J. A. (2007). Aquatic
Photosynthesis (2nd ed., pp. 484). Princeton, New
Jersey: Princeton University Press.
Falkowski, P. G., Barber, R. T., & Smetacek, V. (1998).
Biogeochemical controls and feedbacks on ocean
primary production. Science, 281, 200–206.
Gersonde, R., & Harwood, D. M. (1990). Lower Creta-
ceous diatoms from ODP Leg 113 site 693 (Weddell
Sea) part 1: Vegetative cells. In Proceedings of the
Ocean Drilling Program, Scientific Results (Vol. 113,
pp. 365–402).
Harwood, D. M. (1999). Diatomite. In E. F. Stoermer &
J. P. Smol (Eds.), The diatoms: Applications for the
environmental and earth sciences (pp. 436–443).
Cambridge, UK: Cambridge University Press.
Harwood, D. M., & Gersonde, R. (1990). Lower Creta-
ceous diatoms from ODP leg 113 site 693 (Weddell
Sea). Part 2: Resting spores, chrysophycean cysts, an
endoskeletal dinoflagellate, and notes on the origin of
diatoms. In P. F. Barker & J. P. Kennett et al. (Eds.),
In: Proceedings of the Ocean Drilling Program,
Scientific Results (Vol. 113, pp. 403–415).
Katz, M. E., Finkel, Z. V., Grzebyk, D., Knoll, A. H., &
Falkowski, P. G. (2005). Evolutionary trajectories and
biogeochemical impacts of marine eukaryotic phyto-
plankton. Annual Review of Ecology, Evolution, and
Systematics, 35, 523–556.
Kwiecinska, B. (2000). How the diatoms were found in
the Proterozoic marbles at Przeworno. In A. Wit-
kowski & J. Sieminska (Eds.), The origin and early
evolution of diatoms: Fossil, molecular and biogeo-
graphical approaches (pp. 281–289). Cracow,
Poland: W. Szafer Institute of Botany, Polish
Academy of Sciences.
Lane, T. D., Saito, M. A., George, G. N., Pickering, I. J.,
Prince, R. C., & Morel, F. M. M. (2005). A cadmium
enzyme from a marine diatom. Nature, 435, 42.
Leblanc, K., Quéguiner, B., Diaz, F., Cornet, V.,
Michel-Rodriguez, M., Durrieu de Madron, X., Bowler,
C., Malviya, S., Thyssen, M., Grégori, G., Rembauville,
M., Grosso, O., Poulain, J., de Vargas, C., Pujo-Pay,
M., & Conan, P. (2018). Nanoplanktonic diatoms are
globally overlooked but play a role in spring blooms
and carbon export. Nature Communications, 9, 953.
https://doi.org/10.1038/s41467-018-03376-9.
Mann, D. G. (1999). The species concept in diatoms.
Phycologia, 38, 437–495.
7 Marine Diatoms
Mann, D. G. & Droop, S. J. M. (1996). Biodiversity,
biogeography and conservation of diatoms. Hydrobi-
ologia, 336, 19–32.
Moldowan, J. M., & Jacobson, S. R. (2000). Chemical
signals for early evolution of major taxa: Biosigna-
tures and taxon-specific biomarkers. International
Geologiy Review, 42, 805–812.
Nikolaev, V. A., & Harwood, D. M. (2000). Diversity and
system of classification in centric diatoms. In A.
Witkowski, & J. Sieminska (Eds.), The origin and
early evolution of the diatoms: Fossil, molecular and
biogeographicai approaches (pp. 37–53). Cracow,
Poland: W. Szafer Institute of Botany, Polish
Academy of Sciences.
Rau, G. H., Riebesell, U., & Wolf-Gladrow, D. A. (1996).
A model of photosynthetic 13C fractionation by
marine phytoplankton based on di!usive molecular
CO2 uptake. Marine Ecology Progress Series, 133,
275–285.
Round, F. E., Crawford, R. M., & Mann, D. G. (1990).
The diatoms. Biology and morphology of the genera
(747pp). Cambridge: Cambridge University Press.
Sieminska, J. (2000). The discoveries of diatoms older
than the Cretaceous. In A. Witkowski & J. Sieminska
(Eds.), The Origin and Early Evolution of Diatoms:
Fossil, Molecular and Biogeographical Approaches
(pp. 55–74). Cracow, Poland, Szafer Institute of
Botany, Polish Academy of Sciences.
Simonsen, R. (1979). The diatom system: Ideas on
phylogeny. Bacillaria, 2, 9–71.
Sims, P. A., Mann, D. G., & Medlin, L. K. (2006).
Evolution of the diatoms: Insights from fossil,
biological and molecular data. Phycologia, 45(4),
361–402.
Smetacek, V. (1999). Diatoms and the ocean carbon
cycle. Protist, 150, 25–32.
Smol, J. P., & Stoermer, E. F. (2010). The diatoms.
Applications for the environmental and earth sciences
(2nd ed.). Cambridge: Cambridge University Press.
Sournia, A. (1995). Red tide and toxic marine phyto-
plankton of the world ocean: an inquiry into biodi-
versity. In P. Lassus, G. Arzul, E. Erard-Le-Denn,
P. Gentien, & C. Marcaillou-Le-Baut (Eds.), Harmful
Marine Algal Blooms. Paris: Technique et Documen-
tation, Lavoisier. pp. 103–112.
Stoermer, E. F., & Smol, J. P. (Eds.). (1999). The
diatoms: Applications for the environmental and earth
sciences. Cambridge: Cambridge University Press.
Von Rad, U., Riech, V., & Rosch, H. (1977). Silica
diagenesis in continental marginal sediments off
Northwest Africa. In Y. Lancelot, E. Seibold, et al.,
(Eds.), Initial reports of the deep sea drilling project
(41, pp. 879–905). Washington, DC: US Government
Printing Office.
 Ostracods
1 Introduction
Of all the arthropods, the bivalved tiny ostracods
(0.20–2 mm, rarely up to 32 mm in size) have
the most complete fossil record, extending for
about 450 million years (Ma) from the ordovi-
cian to the present (Martens 1998, 2008; Siveter
et al. 2003; Martens et al. 2003). The ostracods
have a wide distribution from intertidal to hadal
depths (up to 7000 m), in coastal estuaries and
marshes, in most fresh waters, and in a few ter-
restrial habitats. Besides, the fact that the ostra-
cods thrive in a diverse habitat, their valves also
preserve a “snapshot” of the ambient water at the
time of shell formation, thus, making them an
excellent proxy for accessing changes in the
paleoenvironment (see Whatley 1988; Smith and
Horne 2002; De Deckker 2002).
2 General Morphology
2.1 The Carapace and Associated
Structures
The bivalved ostracod body (Figs. 1(1) and 2(2))
is covered by a continuous cuticle secreted by the
two lateral folds of epidermis (Fig. 1(3)), called
the duplicatures (flap-like outgrowths) which
with growth become hardened, leaving only few
parts (the joints) uncovered to allow for move-
ment. The carapace (Fig. 1(4)) is a vital part of
© Springer Nature India Private Limited 2020
S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_8
8
the cuticle and develops as a single cuticular fold
originating on the head region that ultimately
covers the whole body. The ovate, kidney-
shaped or bean-shaped carapace (Fig. 1(4)),
generally measures between 0.5 to 3 mm in
length, but in some Paleozoic species, 80 mm
long ones have also been noted.
On the dorsal part, the two valves are united by
a narrow strip of soft cuticle called the Ligament
(Fig. 1(3)). The adductor muscles (Figs. 13 and 1
(4)) connect the right and left valves and traverse
the soft body in the media region (Fig. 1(3)). In
Podocopa one valve is usually larger than the other
and overlaps the smaller valve, whereas in Myo-
docopa this overlap is less obvious.
Hinge, the articulation of valves, is developed
in the dorsal margin of the two valves (Fig. 1(4)).
The hinges (Figs. 2 and 3) are composed of
variously shaped bars, grooves, teeth, and sock-
ets, thereby articulating (interlocking) the two
valves together. Explained through a series of
complex terminology (Fig. 2), three main cate-
gories of hinges are noted (Fig. 3). These are:
2.1.1 Adont (or Unipartite)
Hinge type with no subdivision into terminal and
median elements (Fig. 3(1); see 3.4 for details).
It is the simplest type and lacks both teeth and
sockets. However, it often has a single groove
along the margin of the larger valve and a cor-
responding ridge on the smaller valve as noted in
Paleozoic and freshwater ostracods.
143
144 8 Ostracods

Fig. 1 Ostracod terminology Modified from van Morkhoven (1962–1963)

2 General Morphology
Fig. 2 Hinge terminology Modified from van Morkhoven (1962–1963)
2.1.2 Merodont
This type has terminal teeth in one valve only
(Fig. 3(2); see 3.4 for details) and is character-
ized by elongate and strongly crenulated terminal
elements on the right valve; the median elements
may be smooth or crenulated.
2.1.3 Amphidont
This type that has teeth and sockets in both
valves (Fig. 3(2); see 3.4 for details). It has short
terminal elements and well-developed teeth on
the right valve that may be crenulated, divided,
or smooth. The median element comprises of an
anterior socket (it can be smooth or divided) and
a median groove that is usually smooth.
2.2 Lamellae and Cardinals
A valve has two lamellae (Fig. 4(1)). The Outer
lamella is calcified and forms the outer surface of
the valve and bends at the valve margin into the
non-calcified inner lamella (Fig. 4(1)). The space
in-between the two is called the Vestibulum (valve
cavity) (Fig. 4(1)) that contains various organs
and is connected in the dorsal region with the main
soft body. The marginal structures (spines,
145
denticles, or ridges) develop near the free edge of
the valves and are characteristic of Cenozoic
ostracods. Among them, the most important one is
the Selvage (Figs. 2(1 and 3) and 3(1))—a chiti-
nous ridge on the duplicature that aids in the clo-
sure of valves along the ventral margin. Its base is
calcified and can be noted on fossil valves as a
continuous crest (Fig. 4(1)). The contact margin of
each valve is subdivided into the cardinal margin
(hinge margin) and the extracardinal margin (free
margin); the latter is composed of the anterior,
ventral, and posterior margins (Fig. 4(2)).
2.3 Pore Canals
Ostracod valves are pierced by diversely shaped
holes called pore canals (Fig. 4(1)). According to
their position, two categories of pore canals are
noted, the normal (lateral) and the radial (mar-
ginal). The lateral ones traverse the outer lamel-
lae, usually at a right angle, and display a large
variety of shapes. Two types of lateral pore
canals are noted, open (simple) and sieve-type
(Fig. 4(1)). The open type has a simple outer
opening (pore), while the sieve-type has vari-
ously shaped sieve-plates. Most species have
146 8 Ostracods

Fig. 3 Hinge types. Modified from van Morkhoven (1962–1963)

2 General Morphology
Fig. 4 1: Sectional details of a podocopid ostracod valve (modified from Kesling 1951). 2: Cardinals. 3–4:
Musculature terminology
more than one type of lateral pore canals so that
both open and sieve-types may occur on the
same species. Most pore canals possess hair-like
setae (sensory structures) which may be long and
thick while others are more delicate. Some
sieve-type pore canals have pigment cells that
function as photoreceptors.
2.4 Muscle Scars
On the internal surface of an ostracod valve,
there are small spots of a somewhat different
147
shell structure, confined to two areas that,
according to their position on the valve, are ter-
med either as the central muscle scar field or the
dorsal muscle scar field (Fig. 4(3 and 4)). Each
contain a large number of scars, and the majority
of them are attachments of short sinews that
connect with muscles. The central muscle scar
field includes three groups of scars:
2.4.1 Adductor Muscle Scar Group
This represents the imprints of closing, and of
adductor muscles. It is generally located in front
of the mid-length (Fig. 4(3 and 4)).
148 8 Ostracods

2.4.2 Mandibular Group

This lies in front of the ventral part of the ad-
ductor muscle scar group. Its scars are formed by
chitinous support rods (and not by muscles)
(Fig. 4(3 and 4)).

2.4.3 Frontal Group

This group is associated with the mandibular
muscles and is situated above the mandibular
scars. It is an area for the attachment of chitinous
rods that help support the mandibulae (Fig. 4(3
and 4)).
The most distinctive group and also the most
resistant to fossilization is the adductor group
(Fig. 4(3 and 4)) that at the familial level is used
for generic or specific identifications. The ad-
ductor muscle scar group of adult ostracods is,
with very few exceptions, situated in front of the
valve mid-length (Fig. 4(3)). Frontal and
mandibular scars (Fig. 4(4)), if present, always
mark the position of the anterior end (Fig. 4(3
and 4)). When muscle scars are obliterated by
fossilization, their position may be derived from
associated features, such as the position of the
median sulcus (elaborated below; see Fig. 5), the
lower portion of which corresponds to the posi-
tion of the adductor scar group or sculptural
features which are often arranged concentrically
or radially around the central muscle scars.

Fig. 5 Orientation and carapace views

2.5 Carapace Orientation

The carapace has the following views (Fig. 5):
2.5.1 Lateral or Side View
It is normal to the contact (sagittal) plane of the
valves (Fig. 5(1 and 2))
2.5.2 Dorsal View
It is the view on the hinge margin, with line of sight
in the sagittal plane of the carapace (Fig. 5(3)).
2.5.3 Ventral View
It is the appearance of the carapace in line of the
sagittal plane seen from below (Fig. 5(4)).
2.5.4 Frontal View
The carapace seen from the anterior end in line of
the sagittal plane (Fig. 5(5)).
2.5.5 Posterior View
The carapace seen from the posterior end in the
sagittal plane (Fig. 5(6)).
The term “carapace length” (Fig. 5(8 and 9))
is not uniformly applied. In straight-backed
ostracods, it is defined as the maximum dimen-
sion of the carapace in the direction parallel to
the hinge line (Fig. 5(8)). In specimens with an
arched dorsum, the length is the maximum
2 General Morphology
distance of end points of the carapace measured
parallel to the basal line (Fig. 5(9)). The height is
measured as the maximum distance perpendicu-
lar to the length (Fig. 5(9)). The width is the
maximum distance of the carapace outline per-
pendicular to the sagittal plane (Fig. 5(8)).
2.6 Valve Closure
The box-like closure as in some platycopins
occurs when the cardinal and extracardinal
regions are not morphologically differentiated
into a true hinge region. In this case, the simple
edge of one valve fits into the marginal groove of
the other and is then called the holosolenic type.
In this case, all the elements of the cardinal and
extracardinal margins are either raised (positive)
or depressed (negative). In the other type of
closure, the hemisolenic, the median part of the
hinge is raised in valves with otherwise de-
pressed features of the contact margin and vice
versa.
2.7 Sculpture
The features of the outer valve surface are con-
sidered as sculpture (Fig. 6). The simplest
sculpture consists of an undifferentiated fine
Fig. 6 Basic valve sculpture
terminology
149
reticular network (Fig. 6). Based on the relative
thickening of walls between neighboring meshes
of this primitive type, varied meshes, ridges, tu-
bercles, spines, and other sculptural features are
derived (Fig. 6); more detailed types and asso-
ciated terminology are illustrated in Fig. 7.
3 Sexual Dimorphism, Ontogeny,
and Morphology of the Carapace
The ostracod male and ostracod female are sep-
arate and have carapaces of different size and
shape; size difference between larval and adult
valves characterizes dimorphism in ostracods
(Fig. 8(1)). Considerable differences in sculpture
between larvae and adults are seen, especially in
species with highly differentiated adult sculpture
(see Fig. 8(1)). Strong ridges or nodes develop
gradually during ontogeny (Fig. 8(1)). Valves at
different ontogenetic stages differ also in their
shapes (Fig. 8(2–4)). Major changes from larval
to adult stages are enumerated in Fig. 8(5).
Dimorphism occurs not only in shape and
function of the soft body parts, size, shape, or
sculpture of the carapace, but may also manifest
itself in finer structural details, such as the pattern
of muscle scars (Fig. 8(3)). Besides dimorphic
features (enumerated above), the other criterion
for sex recognition is the sex ratio; more often
150 8 Ostracods

Fig. 7 Types of sculptures (after Rodriguez-Lazaro and Ruiz-Munoz (2012) and with permission from Elsevier)
3 Sexual Dimorphism, Ontogeny and Morphology of the Carapace 151

Fig. 8 Sexual dimorphism, ontogeny and carapace morphology. 1: Dimorphism and difference in sculpture in
Rehacythereis?kodyami (Pokorný), modified after Pokorný (1967); 2: Lateral outlines of successive instars of
Cypridopsis vidua. 3: Dimorphic shells are typically similar in everything but not in size. Male-shell is generally longer
than the female-shell. 4: Ontogenetic development from the fifth to the ninth (adult) instar of Cytherella posterspinosa
Herrig, modified after Herrig (1966)

the percentage of females is higher than that of

males. The two sexes differ in their lateral outline
or length/height ratio (Fig. 8(3 and 4)); it is
smallest in the earliest larval stages, as these have
a small number of appendages and consequently
a shorter body (Fig. 8(3 and 4)). In Palaeocopida,
dimorphism is of taxonomic value where female
forms are called heteromorphs and differ from
the tecnomorphs (males and juveniles) in pos-
sessing a more inflated posterior region, pro-
nounced ventral lobes, prominent hemispherical
bulges (brood pouches or crumina), or wide frills
extending beyond the free edges of the valves.
4 Classification
The classification of class ostracoda (Figs. 9 and
10) is based on general shape, valves overlap-
ping, presence of ornamentation, hinge charac-
teristics, and male and female structures.
5 General History
Over 65,000 living and fossil taxa have been
described (Ikeya et al. 2005) but only half of the
20,000 living species have been formally
described, and the majority of them have come
from marine and transitional waters; some 2000
species are known from non-marine waters
(Martens et al. 2008). Hence, the diversity of
ostracods is based on broad estimates (see Kempf
1996, 1997; Rodriguez-Lazaro and Ruiz-Munoz
2012) (see Fig. 11).
The earliest known ostracod was probably
marine, an active crawler and possibly a swimmer
(Horne 2003; Rodriguez-Lazaro and Ruiz-Munoz
2012). The first ostracods appeared in the Early
Cambrian; mostly archeocopids and rare leperdi-
tiids (Late Cambrian). A Silurian planktonic
ostracod represents the oldest known record of a
male metazoan (Siveter et al. 2003). The
152 8 Ostracods

Fig. 9 Ostracod classification (after Rodriguez-Lazaro and Ruiz-Munoz (2012) and with permission from Elsevier)
5 General History 153

Fig. 10 Major characteristics of ostracod subclasses (after Rodriguez-Lazaro and Ruiz-Munoz (2012) and with
permission from Elsevier)
154
Fig. 11 Ostracod species diversity
Palaeocopida dominated the Paleozoic, but were
almost wiped out at the end of the Permian
(Fig. 11); they are now represented by the lone
enigmatic genus Manawa. The Leperditicopida
became extinct by the end of the Devonian
(Fig. 11). Both the Myodocopa and Podocopa
were well-established since the Ordovician (see
also Rodriguez-Lazaro and Ruiz-Munoz 2012)
(see Fig. 11). The first major ostracod radiation
occurred in the Early Ordovician and coincided
with a global transgression marked by the
appearance of the first Palaeocopida, Leiocopida,
Podocopida, and Leperditicopida. The Ordovician
was the acme of the Palaeocopida which declined,
thereafter until their extinction in the Permian
(Fig. 11). The Silurian saw the development of the
beyrichiaceans; all members of this group were
restricted to the Silurian–Early Carboniferous.
Due to their restricted occurrence, they are of
biostratigraphic value. During the Mesozoic, the
Triassic saw the beginning of the dominance of
podocopids (Fig. 11); this was also a time of
intense adaptive radiation of the cytheraceans.
Additionally, the Triassic and Permian assem-
blages were characterized by their bizarre spinos-
ity (Kozur 1971) as also by the sculptured
8 Ostracods
cytherellids. The Early Jurassic is marked low
diversity assemblages in which the healdiid genus
Ogmoconcha and the progonocytherid Pro-
cytheridea dominated. Thereafter, in the late part
of the Early Jurassic, the Ogmoconcha disap-
peared and Procytheridea dominated. By Middle
Jurassic, the progonocytherids and trachyleberi-
dids became important as also the smooth deep
water bairdiids and cytherelhds. The Late Jurassic
and Early Cretaceous saw the expansion of limnic
(non-marine deposits, including deltaic,
marginal-lagoonal settings) ostracod faunas.
These rich and highly diverse faunas are used in
intercontinental correlation (Horne 1995). The
Late Jurassic Lophocythere and Protocythere are
useful Late Jurassic stratigraphic markers. By
Cretaceous, the marine cytheraceans suffered a
minor decline. In the Cenozoic, since the Pale-
ocene, the diversity of ostracods has increased
(Fig. 11). Strongly calcified and often richly
sculptured valves of the trachyleberidids, the
cytherettids, and the hemicytherids, became
important elements of Cenozoic assemblages.
Representative ostracod species, through time,
are illustrated in Figs. 12, 13, 14, 15, 16, 17, 18,
19, 20, 21, 22, 23, 24, 25 and 26.
5 General History 155

Fig. 12 Earliest ostracods. 1–2: Bradoria Matthew; 4–5: Indiana Matthew; 6–7:? Mononotella Ulrich and Bassler; 8–
9: Beyrichana Matthew; 10: Sellula Wiman; 11–12: Aluta Matthew; 13:? Cambria Neckaja and Ivanova; 14–16:
Walcottella Ulrich and Bassler; 17:? Eremos Moberg and Segerberg (modified from Moore 1961)
156 8 Ostracods

Fig. 13 Representative species of some Ordovician ostracods. 1: Ctenobolbina ciliate (Emmons); Right valve; 2–3:
Raymondatia goniglypta Kay; 2: Left valve; 3: Dorsal view; 4: Isochilina bulbosa Harris; 5: Eukloedenella
richmondensis Spivey; 5: Right valve; 6–7: Dicranella bicornis Ulrich; 6: Left valve; 7: Ventral view; 8: Euprimitia
labiosa (Ulrich); Left valve; 9: Primitiella constricta Ulrich (Right valve); 10: Primitia tumidula Ulrich (Left valve);
11–12: Sacellatia arcuamuralis Kay; 11: Left valve; 12: Ventral view; 13: Milleratia cincinnatiensis (Miller); Right
valve; 14: Bellornatia tricollis Kay; Left valve; 15–16: Eurychilina reticulata parvifrons Kay; 15: Left valve; 16:
Ventral view; 17: Leperditella rex (Coryell and Schenck); Left valve; 18–19: Schmidtella crassimarginata Ulrich; 18:
Ventral view; 19: Left valve; 20–21: Bairdia (?Ordovician; Carboniferous-Permian); 20: Interview of left valve; 21:
Muscle scar; 22: Bollia suaequata Ulrich; Left valve
5 General History 157

Fig. 14 Representative species of some Silurian ostracods. 1: Eurychilina (Ordovician-Silurian); Right valve; 2–3:
Drepanella (Ordovician-Silurian) (Right valve); 4: Aparchites (Ordovician-Devonian) (Right valve); 5–6: Drepanellina
clarki Ulrich and Bassler; 5: Left valve (Male); 6: Left valve (Female); 7: Zygobolbina conradi Ulrich and Bassler; 8–9:
Zygobeyrichia regina Ulrich and Bassler; 8: Right valve (Female); 9: Right valve (Male); 10: Zygobolba decora
Billings (Left valve); 11: Aparchites obliquatus Ulrich and Bassler; Left valve; 12: Plethobolbina ornata Ulrich and
Bassler; Right valve; 13–14: Mastigobolbina intermedia Ulrich and Bassler (Right valves); 15–16: Bonnemaia transita
Ulrich and Bassler (Right valves); 17: Zygosella mimica Ulrich and Bassler (Left valve); 18–19: Welleria obliqua
Ulrich and Bassler (Right valves); 18: Male; 19: Female; 20: Kloedenella rectangularis Ulrich and Bassler; Left valve;
21: Paraechmina postica Ulrich and Bassler; Left valve; 22–24: Beyrichia (Silurian); 22: Right valve (Female), 23:
Right valve (Male); 24: Lateral view; 25: Entomozoe (Silurian-Permian). (Modified from Treatise on Invertebrate
Paleontology, Part Q: Arthropoda 3-Crustacea / Ostracoda)
158 8 Ostracods

Fig. 15 Representative species of some Devonian ostracods. 1–2: Dizygopleura recta Roth; 1: Left valve; 2: Dorsal
view; 3–4: Tetradella cicatricosa Warthin; 3: Left valve; 4: Ventral view; 5: Parahealdia pecorella Coryell and
Cuskley; Right valve; 6: Thlipsurella putea Coryell and Cuskley; Left valve; 7: Hyphasmaphora textiligera Van Pelt;
Left valve; 8: Thlipsura furcoides Bassler; Left valve; 9: Octonaria loculosa Ulrich; Left valve; 10: Ctenoloculina
acanthophora Swartz and Oriel; Left valve; 11: Quasillites lobatus Swartz and Oriel; Right valve; 12: Octonariella
bifurcata Bassler; Left valve; 13: Eustephanus catastephanes Swartz and Swain; Left valve; 14: Kirkbyella verticolis
Coryell and Cuskley; Right valve; 15: Entomis rugatulus Van Pelt; Left valve; 16: Cytherellina punctulifera (Hall); Left
valve; 17: Hollina devonica Van Pelt; Left valve; 18–20: Monoceratina (Devonian-Recent); 18: Lateral view; 19:
Ventral view; 20: Dorsal view; 21–22: Quasillites (Devonian-Carboniferous); 21: Lateral view; 22: Ventral view; 23:
Leperditia (Silurian-Devonian); Left valve
5 General History 159

Fig. 16 Representative species of some Devonian ostracods. 1–2: Condracypris binoda Roth; 1: Right valve; 2:
Dorsal view; 3: Halliella pulchra Bassler; Left valve; 4: Ranapeltis typicalis Bassler (Left valve); 5-6: Tubulibairdia
windomensis Swartz and Oriel; 5: Right valve; 6: Dorsal view; 7: Aechmina cuspidata Jones and Holl (Right valve); 8:
Parabolbina parvinoda Swartz and Swain (Left valve); 9–10: Favulella favulosa Swartz and Swain; 9: Left valve; 10:
Ventral view; 11: Euglyphella numismoides Swartz and Oriel; 12: Graphiodactylus catenulatus Van Pelt (Left valve);
13: Bicornella tricornis Coryell and Cuskley (Right valve); 14: Stibus kothornostibus Swartz and Swain (Left valve);
15–16: Beecherella bloomfieldensis Swartz and Swain; 15: Right valve; 16: Ventral view; 17: Euglyphella (Devonian);
Muscle scar
160 8 Ostracods

Fig. 17 Representative species of some Mississippian (Carboniferous) ostracods. 1–2: Jonesina insculpta Croneis and
Funkhouser; 1: Left valve; 2: Dorsal view; 3–4: Bairdiolites procerus Cooper; 3: Right valve; 4: Dorsal view; 5–6:
Paracavellina elliptica Cooper; 5: Right valve; 6: Dorsal view; 7–8: Kirkbyella quadratus Croneis and Gutke; 7: Right
valve; 8: Dorsal view; 9: Verrucosella golcondensis Croneis and Gutke; Right valve; 10–11: Glyptopleurina vetula
Cooper; 10: Left valve; 11: Dorsal view; 12–13: Venula striata (Croneis and Funkhouser); 12: Left valve; 13: Dorsal
view; 14–15: Geffenina marmerae Coryell and Sohn; 14: Right valve; 15: Dorsal view; 16: Balantoides quadrilobatus
Morey (Left valve); 17–18: Deloia sulcata Croneis and Funkhouser; 17: Right valve; 18: Dorsal view
5 General History 161

Fig. 18 Representative species of some Mississippian (Carboniferous) ostracods. 1–2: Beyrichiopsis brynhildae
Coryell and Jhonson; 1: Left valve; 2: Dorsal view; 3–4: Beyrichia contracta Cooper; 3: Left valve; 4: Dorsal view; 5–
6: Acratia mucronata Cooper; 5: Left valve; 6: Ventral view; 7–8: Incisurella prima Cooper; 7: Right valve; 8: Dorsal
view; 9–10: Carboprimitia depressa Croneis and Funkhouser; 9: Left valve; 10: Dorsal view; 11–12: Micropara-
parchites spinosus Corneis and Gale; 11: Right valve; 12: Dorsal view; 13–14: Tetratylus ellipticus Cooper; 13: Left
valve; 14: Dorsal view; 15–16: Perprimitia turrita Croneis and Gutke; 15: Right valve; 16: Dorsal view; 17:
Glyptopleura alata Corneis and Funkhouser; Right valve; 18–19: Sargentina allani Coryell and Jhonson; 18: Right
valve; 19: Dorsal view
162 8 Ostracods

Fig. 19 Representative species of some Pennsylvanian (Carboniferous) to Permian ostracods. 1: Bairdia beedei Ulrich
and Bassler (Right valve); 2–3: Cavellina nebrascensis (Geinitz); 2: Left valve; 3: Dorsal view; 4–5: Sansabella laevis
(Warthin); 4: Right valve; 5: Dorsal view; 6–7: Hastifaba robusta Cooper; 6: Left valve; 7: Dorsal view; 8–9: Moorites
minutus (Warthin); 8: Left valve; 9: Dorsal view; 10: Discoidella convexa Scott and Borger (Right valve); 11:
Seminolites truncatus Coryell (Right valve); 12–13: Roundyella simiplicissima (Knight); 12: Left valve; 13: Dorsal
view; 14: Kellettina montosa (Knight) (Right valve); 15–16: Healdia colonyi Coryell and Booth; 15: Right valve; 16:
Dorsal view; 17–19: Darwinulina (Carboniferous-Recent); 19: Muscle scar; 20: Glyptopleura (Carboniferous-Permian)
5 General History 163

Fig. 20 Representative species of some Pennsylvanian (Carboniferous) to Permian ostracods. 1–2: Cornigella
tuberculopsina (Jones and Kirkby); 1: Left valve; 2: Dorsal view; 3: Paraparchites magnus Kellett; Left valve; 4:
Hollinella digitata Kellett; Left valve; 5: Sulcella sulcata Coryell and Sample; Left valve; 6: Amphissites centronotus
(Urich and Bassler); Left valve; 7–8: Ectodemites sullivanensis (Payne); 7: Right valve; 8: Dorsal view; 9–10:
Coryellites lowelli Cooper; 9: Right valve; 10: Dorsal view; 11–12: Fabalicypris wetumkaensis Cooper; 11: Right
valve; 12: Dorsal view; 13–14: Geisina gallowayi (Bradfield); 13: Left valve; 14: Dorsal view
164 8 Ostracods

Fig. 21 Representative species of some Triassic to Jurassic ostracods. 1: Estheriella (Triassic–Cretaceous) (Right
valve); 2–6: Ogmoconcha (Triassic–Lias); 2–3: Left valve; 4–5: Section of the ventral edge; 6: Muscle scar; 7:
Paracypris Sars (Jurassic–Recent); 8–11: Cypridea (Jurassic–Cretaceous); 8: Ventral view; 9: Left valve; 10:
Anterioventral rostrum; 11: Muscle scar; 12–16: Macrodentina (Jurassic–Cretaceous); 15: Hinge; 17–19: Procytheridea
(Jurassic); 20: Cytheropteron Sars (Late Jurassic–Recent); 21–23: Protocythere (Jurassic–Cretaceous); 24–27:
Ilyocypris (Jurassic–Recent); 24: Internal view (Left valve); 25: External view (Left valve); 26: Dorsal view; 27:
Section of the ventral edge of the left valve
5 General History 165

Fig. 22 Representative species of some Jurassic ostracods. 1–4: Cypridea (Jurassic–Cretaceous); 1: Ventral view; 2:
Anterioventral rostrum; 3: Left valve; 4: Muscle scar; 5–9: Macrodentina (Jurassic–Cretaceous); 10–12: Procytheridea
(Jurassic); 13: Paracypris Sars (Jurassic–Recent); 14–16: Protocythere (Jurassic–Cretaceous); 17: Cytheropteron Sars
(Late Jurassic–Recent); 18–21: Ilyocypris (Jurassic–Recent); 18: Internal view (Left valve); 19: External view (Left
valve); 20: Dorsal view; 21: Section of the ventral edge of the left valve
166 8 Ostracods

Fig. 23 Representative species of some Cretaceous ostracods. 1–2: Cytherella ovata (Roemer); 1: Left valve; 2: Dorsal
view; 3: Eucytherura brown-stowensis Alexander; Right valve; 4: Cytherura taxana Alexander (Right valve); 5:
Loxoconcha cretacea Alexander (Right valve); 6–7: Brachycythere goodlandensis Alexander; 6: Right valve; 7: Dorsal
view; 8: Eucythere brown-stowensis Alexander (Right valve); 9–10: Cytheridea evertti Berry; 9: Right valve; 10:
Dorsal view; 11–12: Macrocypris graysonensis Alexander; 11: Right valve; 12: Dorsal view; 13–14: Monoceratina
umbonata (Williamson); 13: Right valve; 14: Dorsal view; 15–16: Bythocypris goodlandensis Alexander; 15: Right
valve; 16: Dorsal view; 17: Cytherelloidea goodlandensis Vanderpool; Left valve
5 General History 167

Fig. 24 Representative species of some Cretaceous ostracods. 1: Cytheropteron trinitiensis (Vanderpool); Right valve;
2–3: Paracypris siliqua Jones and Hinde; 2: Right valve; 3: Dorsal view; 4: Haplocytheridea Stephenson; 5–6:
Orthonotacythere scrobiculata Alexander; 5: Right valve; 6: Dorsal view; 7: Cythereis burlesonensis Alexander (Right
valve); 8–9: Paracytheridea Müller; 10: Brachycythere Alexander; 11: Eocytheropteron Alexander; 12–13: Argilloecia
Sars; 14: Clithrocytheridea Stephenson; 15: Archicythereis van den Bold; 16–17: Cytherella Jones
(Cretaceous-Recent); 16: Section of the ventral edge of the left valve; 17: Muscle scar; 18–19: Xestoleberis
(Cretaceous-Recent); 18: Left valve; 19: Dorsal view
168 8 Ostracods

Fig. 25 Representative species of some Paleocene to Eocene ostracods. 1–2: Alatacythere Hirschmann (Paleocene–
Recent); 3–4: Buntonia Howe (Eocene–Recent); 5–8: Cypris (Eocene–Recent); 5: External view (Left valve); 6:
Internal view (Left valve); 7: Muscle scar; 8: Ventral view; 9: Cytheromorpha Hirschmann (Paleocene–Recent); 10:
Hemicythere Sars (Eocene–Recent); 11: Cytheretta Müller (Eocene–Recent); 12: Buntonia Howe (Eocene–Recent); 13–
17: Cytheridea (Eocene–Recent); 13: Dorsal views of left valve (Female and Male); 14: Hinge; 15: External view of left
valve; 16: Internal view of left valve; 17: Muscle scar; 18–22: Trachyleberis (Eocene–Recent); 18: Left valve; 19:
Hinge; 20: Dorsal view; 21: Internal view (Left valve); 22: Muscle scar. 23–24: Cytheretta (Paleocene–Recent); 24:
Hinge; 25: Laxoconcha (Paleocene–Recent)
5 General History 169

Fig. 26 Representative
species of some Oligocene to
Recent ostracods. 1–3:
Hemicythere (Oligocene–
Recent); 2: Hinge; 4:
Anomocytheridea Stephenson
(Miocene-Recent); 5:
Perissocytheridea Stephenson
(Miocene); 6–7: Cyprideis
(Miocene-Recent); 6: Dorsal
view with tubercles;
Section of the ventral edge of
both Left and Right valves; 8–
9: Cypridina Milne Edwards
(Recent); 8: Lateral view; 9:
Front view; 10–11:
Cytheridella Daday (Recent)

References Kempf, E. K. (1997). Index and bibliography of

De Deckker, P. (2002). Ostracod Palaeoecology.
In J. A. Holmes & A. R. Chivas (Eds.), The
Ostracoda: Applications in quaternary research
(pp. 121–134). Ser.: AGU Mon.
Herrig, E. (1966). Ostracoden aus der Weissen Schreibk-
reide (Unter-Maastricht) der Insel Riigen. Paldontol.
Abh., A, 2, 693–1024.
Horne, D. J. (1995). A revised ostracod biostratigraphy
for the Purbeck-Wealden of England. Cretaceous
Research, 16, 639–663.
Horne, D. J. (2003). Key events in the radiation of the
Ostracoda. In: Park, L. E. & Smith, A. J. (Eds.),
Bridging the gap: Trends in the Ostracode biological
and geological sciences (Vol. 9, pp. 181–201). The
Paleontological Society Papers.
Kempf, E. K. (1996). Index and bibliography of marine
Ostracoda, vols. 1–9. Geologisches Institut der
Universitat Koeln, Sonderveroffentlichungen 50–53,
88, 102, 103. (2002, 2004, 2008, CD version).
non-marine Ostracoda, vols. 1–6. Geologisches Insti-
tut der Universitaet zu Koeln, Sonderveroffentlichun-
gen 35–38, 77, 109–112. (2001, 2006, CD version).
Kesling, R. V. (1951). Terminology of ostracode cara-
paces. Contributions (Vol. 9, pp. 93–171). Museum of
Paleontology, University of Michigan.
Kozur, H. (1971). Die Bairdiacea der Trias. Teil II:
Skulpturierte Bairdiidae aus mitteltriassischen Tief-
schelfablagerungen. Geol. Paldontol. Mitt. Innsbruck,
1, 1–21.
Martens, K. (Ed.). (1998). Sex and parthenogenesis.
Evolutionary ecology of reproductive modes in
non-marine ostracods (p. 335). Leiden: Backhuys.
Martens, K. (2008). Ancient asexuals: darwinulids not
exposed. Nature, 453, 587.
Martens, K., Rossetti, G., & Horne, D. J. (2003). How
ancient are ancient asexuals? Proceedings of the Royal
Society of London, 270, 723–729.
Martens, K., Schön, I., Meisch, C., & Horne, D. J. (2008).
Global diversity of ostracods (Ostracoda, Crustacea)
in freshwater. Hydrobiologia, 595, 185–193.
170
Moore, R. C. (Ed.). (1961). Treatise on Invertebrate
Paleontology, Part Q, Arthropoda 3 (PP. 74–92).
Boulder Colorado and Lawrence, Kansas: Geological
Society of America and University of Kansas Press.
Pokorny, V. (1967). New Cythereis species (Ostracoda,
Crustacea) from the lower Turonian of Bohemia,
Czechoslovakia. Acta Universitatis Carolinae, Geo-
logica, 1967, 365–378.
Rodriguez-Lazaro, J., & Ruiz-Munoz, F. (2012). A gen-
eral introduction to ostracods: morphology, distribu-
tion, fossil record and applications. Developments in
Quaternary Science, 17 (Elsevier). https://doi.org/10.
1016/b978-0-444-53636-5.00001-9.
Siveter, D., Sutton, M. D., Briggs, D. E. G., & Siveter, D.
(2003). An ostracode crustacean with soft parts from
the lower Silurian. Science, 302, 1749–1751.
8 Ostracods
Smith, A., & Horne, D. J. (2002). Ecology of marine,
marginal-marine and non-marine ostracods.
In J. A. Holmes & A. R. Chivas (Eds.), The
ostracoda: Applications in quaternary research
(pp. 37–64). Ser.: AGU Mon.
van Morkhoven, F. (1962–1963). Post-palaeozoic ostra-
coda. Their morphology, taxonomy and economic use,
vol. 1 (1962, general); vol. 2 (1963, generic descrip-
tions). Elsevier, Amsterdam.
Whatley, R. (1988). Ostracoda and palaeogeography. In:
De Deckker, P., Colin, J.-P., & Peypouquet, J.-
P. (Eds.), Ostracoda in the Earth Sciences (pp. 103–
123). Elsevier, Amsterdam.
 Benthic Foraminifera
9

The benthic foraminifera (Fig. 1(6–14)), on the

1 Introduction
other hand, live on, in, or near the bottom of the
ocean (floor) and are equally widely distributed
Foraminifera are single-celled protozoans with a
from tidal pools along the foreshore, out to the
very wide distribution pattern ranging from the
continental shelf, and down to abyssal depths.
paralic (littoral) to abyssal depths and with a
They move around a little with their pseudopodia
geological history spanning since the Early
(Fig. 1(13 and 14)) or have a planktonic phase in
Cambrian (see Murray 2006). Three major
their life cycle. More than 5000 recent for-
groups of foraminifera are noted, planktic (Fig. 1
aminiferal species are known (in contrast to *40
(1–5)), smaller benthic (Fig. 1(6–9)), and larger
of planktic species), thus, making them excellent
benthic; the latter are identified by their larger
proxies for present and past environments and
size and complex interiors, visible in thin sec-
their changes (see Jones 2014; Jain and Farouk
tions (Fig. 1(10–12)). The foraminiferal test
2017).
(Fig. 1(13)) encloses the cytoplasm (soft tissue)
of the foraminiferan cell with an outer layer
called the ectoplasm and an inner layer, the
endoplasm; the latter gives rise to sticky pseu-
2 External Morphological Features
dopodia that traps food for the organism (Fig. 1
The morphological test features (test shape,
(14)). The endoplasm contains the nucleus, cell
chamber arrangement, chamber form, position of
bodies and organelles (such as mitochondria,
primary aperture, aperture form, wall
golgi apparatus, and ribosomes) (Fig. 1(13)).
microstructure, sutures and keel, and ornamen-
This chapter primarily deals with benthic
tation) are of prime importance for group clas-
foraminifera with minor reference to planktic
sification (Fig. 2). These features (Fig. 2(1)) are
foraminifera (for some examples of test features,
extremely varied, even in a morphologically
primarily to show types of microperforations).
well-constrained group of Foraminifera, such as
The planktic foraminifera (Fig. 1(1–5)) are pho-
the lagenids (see Fig. 3). More than 1000 species
tosynthetic symbionts, cosmopolitan, and live as
have been identified for this unilocular
drifting in the open ocean. They are excellent
(monothalamous) group traditionally containing
proxies of ocean currents and climate changes, as
Lagena, Oolina, Fissurina, and Parafissurina
their distribution correlates with ocean dynamics
(see Loeblich and Tappan 1964; Haynes 1981).
(such as salinity and water–temperature). They
Later, this grouping was modified using identi-
are, at places, abundant in modern oceans that
fiable characters of taxonomic value such as the
their tests, after death, sink to form thick sedi-
nature and position of the aperture, the presence
mentary layers such as the Globigerina ooze.

© Springer Nature India Private Limited 2020 171

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_9
172 9 Benthic Foraminifera

Fig. 1 Major groups of foraminifera groups. Figure 1–55: Planktic foraminifera; 6–9: Smaller benthic foraminifera;
10–12: Larger benthic foraminifera; 13: The foraminiferan test which encloses the cytoplasm (soft tissue) of the
foraminiferan cell with an outer layer called ectoplasm and an inner layer, endoplasm; the latter gives rise to sticky
pseudopodia that trap food for the organism (Fig. 1(14)). All photographs are from Kender et al. (2008), courtesy
Michael A. Kaminski and with permission from Micropaleontology
2 External Morphological Features 173

Fig. 2 Morphological features of benthic foraminifera explained and illustrated in this chapter
174 9 Benthic Foraminifera

Fig. 3 Morphological variability in lagenids. Modified after Mikhalevich and Debenay (2001), Revets (2005)
2 External Morphological Features
or absence of an entosolenian tube, as well as the
overall shape of the test (see Jones 1984; Revets
2005). Revets (2005) noted that despite the
simplicity of the single-chambered test, about 20
characters can be used to classify the lagenids
(Fig. 3). These characters fall into broad four
groups: the overall shape of the test; the nature of
the apertural complex, including details of the
entosolenian tube; the nature of the test wall; and
ornamentation of the test (Fig. 3) (see also
Mikhalevich and Debenay 2001). Hence, large
variability and complexity exit for the description
of foraminifera. Due to this variability and
complexity of morphological features, this
chapter restricts itself in illustrating basic features
relevant to benthic foraminiferal studies. Most
illustrations are by means of photographs (cour-
tesy Professor Anthony A. Kaminski and with
permission from the Mircopaleontology journal),
rather than using line diagrams, as done for other
chapters. Wherever possible, examples are given
at the species level, if not, then at the generic
level.
2.1 Test Shape
Test consists either of a single chamber (uniloc-
ular) (Fig. 3) or multiple chambers (multilocular)
(Fig. 4). The chambers are generally <1 mm
across and are interconnected by one or through
several openings called foramen; the opening in
the last chamber is called the aperture (Fig. 2(1)).
Ten basic types of test shapes are enumerated
below and illustrated in Fig. 4; these also reflect
the same pattern of large variability. Hence, in a
coiled spirally test shape (Fig. 4(1–5)), there can
be varied combinations, from tight spiral coiling
(Fig. 4(1)) to forms that exhibit relative open
coiling (Fig. 4(5)).
2.1.1 Coiled Spirally
The test shape is coiled along its horizontal axis
with very less rate of translation, i.e., along the
vertical axis (Fig. 4(1–5)).
175
2.1.2 Elongate
The test shape is coiled along its vertical axis
(Fig. 4(6–10)).
2.1.3 Globose-Ovate
The test shape is globular to ovate in shape
(Fig. 4(11–13)).
2.1.4 Round, Flat, and Discoidal
The test shape is coiled along its horizontal axis
with no rate of translation, i.e., along the vertical
axis (Fig. 4(14–16)).
2.1.5 Tubular
The test shape is tube-like (Fig. 4(17)).
2.1.6 Conical to Subconical
The test is cone-shaped (Fig. 4(18–21)).
2.1.7 Fusiform
The test shape is spindle-shaped (Fig. 4(22–25)).
2.1.8 Leaf-like
The test shape is leaf-like (Fig. 4(26–28)).
2.1.9 Irregular
The test is very irregular in shape (Fig. 4(29–
30)).
2.1.10 Dendroid
The test shape is dendritic in nature (Fig. 4(31)).
2.2 Chamber Arrangement
In many primitive foraminifera, the test is
unilocular (=single chambered or monothalam-
ous; Fig. 5(1–4)). In more advanced forms, the
test is multilocular (=multi-chambered; Fig. 5(5–
42)) such as tubular/enrolled with 1–2 chambers
(Fig. 5(5 and 6)), uniserial (Fig. 5(7 and 8)),
biserial (Fig. 5(9)), triserial (Fig. 5(10–14)), a
combination of triserial/biserial/uniserial (Fig. 5
(15–19)), trochospiral (Fig. 5(20–26)), planispi-
ral (Fig. 5(27–32)), milioline/quinquiloculine
176 9 Benthic Foraminifera

Fig. 4 Major types of test

shape. 1–5: coiled spirally; 6–
10: Elongate; 11–13:
Globose-ovate; Fig. 4(14–
16): Round, flat, and
discoidal; 17: Tubular; 18–21:
concal to subconical; 22–25:
fusiform; 26–28: Leaf-like;
29–30: Irregular; 31:
Dendroid. All photographs
are from Kender et al. (2008),
courtesy Michael A.
Kaminski and with
permission from
Micropaleontology
2 External Morphological Features 177

Fig. 5 Major types of chamber arrangement: 1–4: Unilocular; 5–42: Multilocular; 5–66: Tubular/enrolled with 1–2
chambers; 7–8: Uniserial; 9: Biserial; 10–14: triserial; 15–19: A combination of triserial/biserial/uniserial; 20–26:
Trochospiral; 27–32: Planispiral; 33–38: Milioline/quinquiloculine; 36–40: Irregular streptospiral; 41–42: Spiral to
serial. All photographs are from Kender et al. (2008), courtesy Michael A. Kaminski and with permission from
Micropaleontology
178
(Fig. 5(33–38)), irregular streptospiral (Fig. 5
(36–40)), spiral to serial (Fig. 5(41 and 42)), or
several combinations of these.
2.2.1 Uniserial
The chamber arrangement is in a single row
(Fig. 5(7 and 8)).
2.2.2 Biserial
The chambers are arranged in two rows (Fig. 5
(9)).
2.2.3 Triserial
The chambers are arranged in a trochospire (i.e.
3 chambers in each whorl, placed *120°
between median planes of consecutive chambers)
(Fig. 5(10–14)).
2.2.4 Trochospiral (Trochoid,
Rotaliform)
The chambers are arranged spirally, i.e., in a
helical coil, involute on one side, and evolute on
the other, and the whorl rate of translation is
always greater than zero. The spiral and umbili-
cal sides are dissimilar (Fig. 5(20–26)).
2.2.5 Planispiral
In this arrangement, the whorl rate of translation
is zero, i.e., there is no growth in the third axis.
With regard to the plane of bilateral symmetry,
the spiral and umbilical sides are symmetrical, if
not identical (Fig. 5(27–32)).
2.2.6 Miliolid
In this arrangement, the growth is in varying
planes along a vertical axis with externally visi-
ble chambers (Fig. 5(33–38)). A note must be
made here about the Milioline coiling; it is best
explained by 25 “…porcelaneous foraminifera:
bilocular coils where all terminal apertures are
positioned on a single common axis (apertural
axis); with coiling axis normal to apertural axis,
and rotating in such a manner that between the
median planes of consecutive chambers various
angles are produced, e.g., 72° (quinqueloculine;
[Fig. 5(33–35)]), 120° (triloculine; [Fig. 5(36
and 37)]) or 180° (spiroloculine or biloculine;
[Fig. 5(38)]) or that chambers are positioned in
9 Benthic Foraminifera
an s-shaped curve in one plane as seen in section
(sigmoiline).”
2.2.7 Irregular Streptospiral
The chambers are irregularly enrolled (Glomo-
spira, Ammosphaeroidina, and Paratrochammi-
noides; Fig. 5(36–40)).
2.2.8 Spiral to Serial
The chamber arrangement is either an open spire
(i.e., the whorl rate of translation is zero) or the
initial whorls are coiled (Fig. 5(41–42)) (Fig. 6).
2.3 Chamber Form
The forms of chambers are equally varied
(Fig. 7). These include: spherical–globular
(Fig. 7(1–5)), pyriform oval convex (Fig. 7(5–
8)), biconvex (Fig. 7(9 and 10)), round and flat
(Fig. 7(11 and 12)), tabular (Fig. 7(13 and 14)),
triangular to trapezoid (Fig. 7(15–18)), semicir-
cular cyclical (Fig. 7(19 and 20)), brick (Fig. 7
(21–23)), A or V-shaped, 1–2 chambers (Fig. 7
(24–27)), crescentic (Fig. 7(27–31)), and irregu-
lar (Fig. 7(32 and 33)).
2.4 Position of Primary Aperture
In the wall of the final chamber, there is an
opening called aperture; the primary aperture is
the main opening (see Fig. 2(1)). Aperture varies
considerably both in its position (Fig. 7) and
forms (see Fig. 8). The aperture connects the
external pseudopodia with the internal ectoplasm
and endoplasm and gives passage to food, con-
tractile vacuoles, cytoplasm, nuclei, excretory
products, and enables the release of daughter
cells. Throughout life, the position of the aper-
ture remains constant and each chamber is linked
to the next by a single or multiple foramen. With
regard to the position of the primary aperture,
nine types are noted (Fig. 7). These are: terminal
(Fig. 7(1–3)), umbilical (Fig. 7(4–8)), extraum-
bilical (Fig. 7(9 and 10)), basal–extraumbilical
(Fig. 7(11 and 12)), umbilical–extraumbilical:
(Fig. 7(13–20)), basal interiomarginal (Fig. 7
2 External Morphological Features 179

Fig. 6 Major types of chamber form. 1–4: Spherical–globular; 5–8: Pyriform oval convex: 9–10: Biconvex; 11–12:
Round and flat; 13–14: Tabular; 15–18: Triangular to trapezoid; 19–20: Semicircular cyclical; 21–23: Brick; 24–27: A
or V-shaped, 1–2 chambers; 28–31: Crescentic; 32–33: Irregular. All photographs are from Kender et al. (2008),
courtesy Michael A. Kaminski and with permission from Micropaleontology

(21–27)), areal (Fig. 7(27–34)), no primary but 2.4.2 Umbilical

multiple apertures (Fig. 7(35)), and aperture not See Fig. 7(4–8).
visible (Fig. 7(36–38)).
2.4.3 Extraumbilical
2.4.1 Terminal The opening is in the final chamber and not
See Fig. 7(1–3). connected to the umbilicus, commonly sutural
180 9 Benthic Foraminifera

Fig. 7 Position of primary aperture. 1–3: Terminal; 4–8: Umbilical; 9–10: Extraumbilical; 11–12: Basal–extraum-
bilical; 13–20: Umbilical–Extraumbilical; 21–27: Basal interiomarginal; 28–34: Areal; 35: No primary but multiple
apertures; 36–37: Aperture not visible. All photographs are from Kender et al. (2008), courtesy Michael A. Kaminski
and with permission from Micropaleontology

(interiomarginal) midway between umbilicus and 2.4.4 Basal-Extraumbilical

periphery; Fig. 7(9–10)). See Fig. 7(11 and 12).
2 External Morphological Features 181

2.4.5 Umbilical–Extraumbilical 2.4.9 Aperture not Visible

See Fig. 7(13–20). See Fig. 7(36–37).

2.4.6 Basal Interiomarginal

See Fig. 7(21–27). 2.5 Aperture Form

2.4.7 Areal The shape of aperture can be round–oval–reni-

Within front wall (aperture is in the face of the
last chamber, not at its base nor at the shell
margin, may be single or multiple; Fig. 7(28–
34)).
2.4.8 No Primary but Multiple
Apertures
See Fig. 7(35).
form (Fig. 8(1–5)), arcuate lunear–arch (Fig. 8
(6–8)), slit (Fig. 8(9–12)), loop-shaped (Fig. 8
(13–15)), cribrate (Fig. 8(16 and 17)), pores in
rows (Fig. 8(18 and 19)), radiate (Fig. 8(20 and
21)), lip (Fig. 8(22)), neck (Fig. 8(23–25)), tooth
(Fig. 8(26–29)), and special (Fig. 8(30–31)).
2.5.1 Round–Oval–Reniform
See Fig. 8(1–5).

Fig. 8 Major types of aperture form. 1–5: Round–oval–reniform; 6–8: Arcuate Lunear–arch; 9–12: Slit; 13–15: Loop-
shaped; 16–17: cribrate; 18–19: Pores in rows; 20–21: Radiate; 22: Lip; 23–25: Neck; 26–29: Tooth; 30–31: Special.
All photographs are from Kender et al. (2008), courtesy Michael A. Kaminski and with permission from
Micropaleontology
182 9 Benthic Foraminifera

2.5.2 Arcuate Lunear-Arch 2.5.8 Lip

See Fig. 8(6–8). See Fig. 8(22).

2.5.3 Slit 2.5.9 Neck

See Fig. 8(9–12). See Fig. 8(23–25).

2.5.4 Loop-Shaped 2.5.10 Tooth

See Fig. 8(13–15). See Fig. 8(26–29).

2.5.5 Cribrate 2.5.11 Special

Sieve-like; perforated with multiple round holes; See Fig. 8(30 and 31).
Fig. 8(16 and 17).

2.5.6 Pores in Rows 2.6 Wall Structure and Test

See Fig. 8(18 and 19). Composition

2.5.7 Radiate The foraminiferal wall material is largely of three

See Fig. 8(20 and 21). types: calcareous (porcelaneous; Fig. 9(1);
microgranular: Fig. 9(2 and 3); hyaline: Fig. 9

Fig. 9 Wall structure and test composition. 1–4: Calcareous; 1: Porcelaneous; 2–3: microgranular; 4: Hyaline: 5:
Agglutinated; 6: Organic
2 External Morphological Features
(4)), Agglutinated (arenaceous and argillaceous;
Fig. 9(5)), and rarely organic (Fig. 9(6)).
2.6.1 Porcelaneous
The test wall is calcareous, shiny, and pearly to
white (Fig. 9(1)). The wall is composed of opti-
cally cryptocrystalline lathes and rods (arranged
randomly) or needles (arranged in a tile-roof
pattern) of high magnesium calcite. The wall is
generally imperforate, but may possess pits
(finely perforate proloculus).
2.6.2 Microgranular
The test wall is composed of calcareous ele-
ments, of minute calcite crystals that are sub-
spherical and equidimensional, held together
with very little cement (Fig. 9(2 and 3)). The
latter were originally granular (secreted by the
organism) but later recrystallized. The granules
are aligned in rows perpendicular to the outer
wall, thus, forming a fibrous structure. In com-
parison with porcelaneous, the microgranular
wall is much more stable and resistant to diage-
netic recrystallization.
2.6.3 Hyaline
These appear glassy in reflected light; the test is
thick and perforate (Fig. 9(4)). The c-axis of the
calcite framework shows a preferred growth
orientation for certain foraminiferan groups; the
wall is penetrated by fine pores and hence, ter-
med perforate.
2.6.4 Agglutinated
The test wall is composed of foreign particles
picked up from the ambient environment and
glued together by the organic or biomineralized
cement, produced by the cell; the accumulated
grains are selected on the basis of specific grav-
ity, shape, or size (Fig. 9(5)). The test wall may
be arenaceous (wall is composed of sand or other
foreign particles) or argillaceous (composed of
clay or mud).
2.6.5 Organic
The organic wall is composed of protinaceous
mucopolysaccharides (Fig. 9(6)). The wall is
183
thick in earlier growth stages but thins out in the
ultimate and penultimate chambers.
There are also few oddities, as well. The sub-
order Spirillinina has a test constructed of an
optically single crystal of calcite, the Suborder
Silicoloculinina as the name suggests has a test
composed of silica. Another group (the Suborder
Involutina) have a two-chambered test composed
of aragonite. The Robertinina also have a test
composed of aragonite and the Suborder Carterina
is believed to secrete spicules of calcite which are
then weakly cemented together to form the test.
2.7 Sutures
A suture is the line of attachment between two
adjacent chambers or between two whorls in
varied combination (Fig. 10). These can be de-
pressed (Fig. 10(1–4)), raised (Fig. 10(5–7)),
straight (Fig. 10(8–10)), curved (Fig. 10(11–
14)), thickened (Fig. 10(15)), ponticuli (Fig. 10
(16 and 17)), and keeled (Fig. 10(18–20)).
2.7.1 Depressed
The suture forms a depression (Fig. 10(1–4)),
2.7.2 Raised
The suture is raised like a ridge (Fig. 10(5–7)),
2.7.3 Straight
The suture is straight (Fig. 10(8–10)),
2.7.4 Curved
The suture is curved, often backward (Fig. 10
(11–14)),
2.7.5 Thickened
The suture is marked by ridges (Fig. 10(15)),
2.7.6 Ponticuli
The suture is characterized by small holes
(Fig. 10(16 and 17)) and
2.7.7 Keeled
The suture is marked by elevated ridges along the
venter (Fig. 10(18–20)).
184
Fig. 10 Major types of sutures. 1–4: Depressed; 5–7: Raised; 8–10: Straight; 11–14: Curved; 15: Thickened; 16–17:
ponticuli; 20: Keeled. All photographs are from Kender et al. (2008), courtesy Michael A. Kaminski and with
permission from Micropaleontology
2.8 Ornamentation
The external surface of the test bears numerous
types of protrusions that may be in the form of
spines (termed spinose), keels (carinate), rugae
(rugose), fine striae (striate), coarser costae
(costate), granules (granulate) or a reticulate
sculpture, or a combination of these (Fig. 11).
The major types of ornamentation include Hispid
to Pustulose (Fig. 11(1)), Costae (Fig. 11(2–4)),
Striate (Fig. 11(5–9)), Spinose projections
(Fig. 11(10 and 11)), Reticulate (Fig. 11(12 and
13)), Spinose (Fig. 11(14–16)) and Fistulose
(Fig. 11(17)).
2.8.1 Hispid to Pustulose
The test is covered with minute pustules or
pseudospines (Fig. 11(1)).
9 Benthic Foraminifera
2.8.2 Costae
The test is covered with long ridges (Fig. 11(2–
4)).
2.8.3 Striate
The test is covered with thin costae (Fig. 11(5–
9)).
2.8.4 Spinose Projections
The test is covered with very small spinose
projections (Fig. 11(10 and 11)).
2.8.5 Reticulate
The test is criss-crossed with small ridges in a net
or network-like pattern (Fig. 11(12 and 13)).
2.8.6 Spinose
The test has acicular spines, commonly noted in
planktic foraminifera (Fig. 11(14–16)).
2 External Morphological Features 185

Fig. 11 Major types of

ornamentation. 1: Hispid to
pustulose; 2–4: Costae; 5–9:
Striate; 10–11: Spinose
projections; 12–13:
Reticulate; 14–16: Spinose;
12.17: Fistulose. All
photographs are from Kender
et al. (2008), courtesy
Michael A. Kaminski and
with permission from
Micropaleontology

2.8.7 Fistulose period the Fusulinina (larger benthic for-

The covered with fistula, a reed-like pattern
(Fig. 11(17)).
3 Geological History
Rare agglutinated tubes resembling Bathysiphon
have been recorded since the earliest Cambrian
(Ehrenberg 1843) (see Fig. 12). Agglutinated
foraminifera became more abundant in the
Ordovician but true multichambered forms did
not appear until the Devonian, during which
aminifera) began to flourish, culminating in the
complexly constructed tests of the Fusulinacea in
Late Carboniferous and Permian times (not dis-
cussed in this chapter). This super-family died
out at the end of the Palaeozoic. Miliolina and
Lagenina (smaller benthic foraminifera) first
appeared in the Early Carboniferous. Important
Mesozoic events include the appearance and
radiation of the Rotaliina (largely from endo-
thyracean stock), Miliolina and complex Textu-
lariina in the Jurassic, soon followed by the
appearance of the first unquestionably planktonic
186 9 Benthic Foraminifera

Fig. 12 Geological history

3 Geological History 187

Fig. 13 Some representative species of foraminifera with their major taxonomic characteristics. Age and major
taxonomic characteristics are after Loeblich and Tappan (1988). 1–3: Alabamina Toulmin; 4–5: Ammonia Brünnich; 6:
Amphistegina d’Orbigny; 7–10: Archaeosepta Wernli; 11–13: Angulogavelinella Hofker; 14: Ammodicus Reuss; 15:
Biloculinella Wiesner; 16–18: Bigenerina d’Orbigny; 19: Bolivina d’Orbigny; 20–21: Bolivinopsis Yakovlev; 18:
Microspheric; 19: Megalospheric; 22–23: Brizalina Costa; 24: Buliminoides Cushman; 25–28: Cassidulina d’Orbigny;
29: Cassidella Hofker; 30: Cerobertina Finlay
188 9 Benthic Foraminifera

Fig. 14 Some representative species of foraminifera with their major taxonomic characteristics. Age and major
taxonomic characteristics are after Loeblich and Tappan (1988). 1: Cancris de Montfort; 2: Citharina d’Orbigny; 3:
Chilostomella Reuss; 4–5: Cyclammina Brady; 6–7: Clavulina d’Orbigny; 8: Choffatella Schlumberger; 9: Cyclogyra;
10–11: Cruciloculina d’Orbigny; 12–13: Eggerella Cushman; 14–16: Ehrenberginia Reuss; 17: Fissurina Reuss; 18:
Frondicularia Defrance; 19–26: Fursenkonina Loeblich and Tappan; 27: Flabellammina Cushman; 28–29: Gyroidella
Saidova; 30–31: Gyroidina d’Orbigny
3 Geological History 189

Fig. 15 Some representative foraminifera species with their major taxonomic characteristics. Age and major
taxonomic characteristics are after Loeblich and Tappan (1988). 1: Geosella; 2: Gyroidinoides Brotzen; 3:
Haplophragmoides Cushman; 4–6: Haplophragmium Reuss; 7: Hopkinsina Howe and Wallace; 8–9: Hoeglundina
Brotzen; 10: Jadammina Bartenstein and Brand; 11: Jullienella Schlumberger; 12–13: Kurnubia Henson; 14–15:
Lagena Popescu; 16: Trochulina d’Orbigny; 17–18: Lenticulina Lamarck; 19–22: Laticarinina Galloway and Wissler;
23: Loxostomum Ehrenberg; 24: Neocorbina; 25–26: Nonionella Voloshinova; 27–28: Pyrgo Defrance
190 9 Benthic Foraminifera

Fig. 16 Some representative foraminifera species with their major taxonomic characteristics. Age and major
taxonomic characteristics are after Loeblich and Tappan (1988). 1–3: Paleopolymorphina Cushman and Ozawa; 4–6:
Palaeomiliolina Antonova; 7–9: Pealerina Lalicker; 10–11: Pleurostomella Reuss; 12–14: Protoglobobulimina Hofker;
15–17: Polystomammina Yabe and Hanzawa; 18–20: Pseudotrochammina Frerichs; 21: Quinqueloculina d’Orbigny;
22: Reophax de Montfort; 23–24: Rectobolivina Cushman; 23: Microspheric; 24: Megalospheric; 25–26: Recurvoides
Earland; 27: Siphotextularia Finlay; 28: Spiroplectammina Cushman; 30: Spirillina Ehrenberg; 31: Spirillinoides
Rhumbler; 32: Syringammina; 33: Stetsonia Brotzen
3 Geological History 191

Fig. 17 Some representative foraminifera species with their major characteristics. 1–3: Trochammina Parker and
Jones; 4: Valvobifarina Hofker; 5–6: Verneuilina d’Orbigny; 7–10: Vulvulina d’Orbign; 11–12: Verneuilinoides
Loeblich and Tappan; 13: Vertebralina d’Orbigny; 14: Uvigerina d’Orbigny

foraminifera (Oxford et al. 2002). Cretaceous
tropical regions witnessed a flowering of the
larger forms of miliolines and rotaliines while the
widespread chalk seas and newly opened Atlan-
tic Ocean favored a thriving planktonic
population.
Representative benthic foraminiferal species
through time are illustrated in Figs. 13, 14, 15,
16 and 17.
References
Ehrenberg, C. G. (1843). Verbreitung und Finfluss des
Mikniskopischen Lebens in Süd– und Nord-Amerika.
Physikalische Abhandlungen der Königlichen Akade-
mie der Wissenschaften zu Berlin, 1841. Theil, I, 291–
446.
Haynes, J. R. (1981). Foraminifera (p. 433). London:
Macmillan.
Hottinger, L. (2006). Illustrated glossary of terms used in
foraminiferal research. Carnets de Géologie (Note-
books on Geology), (Mémoires). https://doi.org/10.
4267/2042/5832.
Jain, S., & Farouk, S. (2017). Shallow Water Aggluti-
nated Foraminiferal response to Late Cretaceous—
early Paleocene sea-level changes in the Dakhla Oasis,
Western Desert. Egypt. Cretaceous Research, 78, 1–
18.
Jones, R. W. (1984). A revised classification of the
unilocular Nodosariida and Buliminida (Foramini-
fera). Revista Espanola de Micropaleontologia, 16,
91–160.
192
Jones, R. W. (2014). Foraminifera and their Applications
(391 p.). Cambridge: Cambridge University Press.
Kender, S., Kaminski, M. A., & Jones, R. W. (2008).
Early to middle Miocene foraminifera from the
deep-sea Congo Fan, offshore Angola. Micropaleon-
tology, 54(6), 477–568.
Loeblich, A. R., & Tappan, H. (1964). Sarcodina-chiefly
“thecamoebians” and foraminiferida. In R. C. Moore
(Ed.), Treatise on invertebrate paleontology (Part C,
Protista 2, Vol. 2, pp. 1–900). University of Kansas
Press.
Mikhalevich, V., & Debenay, J.-P. (2001). The main
morphological trends in the development of the
foraminiferal aperture and their taxonomic signifi-
cance. Journal of Micropalaeontology, 20, 13–28.
9 Benthic Foraminifera
Murray, J. (2006). Ecology and Applications of Benthic
Foraminifera (426 pp). Cambridge: Cambridge
University Press.
Orbigny, A. D. (1826). Tableau methodique de la classe
des Cephalopodes. Annates des Sciences Nalurelles, 7,
245–314.
Oxford, M. J., Gregory, F. J., Hart, M. B., Henderson, A.
S., Simmons, M. D., & Watkinson, M. P. (2002).
Jurassic planktonic foraminifera from the United
Kingdom. Terra Nova, 14(3), 205–209.
Revets, S. A. (2005). A key to the unilocular hyaline
Foraminifera. Journal of Micropalaeontology, 24,
145–158.
 Calcareous Nannofossils
10

seafloor. These skeletal remains along with

1 Introduction
nannoliths become part of the sedimentary record
and thus provide a high-resolution biostratigra-
The calcareous nannofossils (nannoplankton
phy for many pelagic deposits of Jurassic to
with calcareous tests) have traditionally been
Holocene age (Young et al. 1994; Perch-Nielsen
defined as marine organisms that are less than 30
in Bolli et al. 1985a, b; Bown 1998).
microns (µm) in size (Lohmann 1909) (see Fig. 1
The heterococcoliths that form the bulk of the
for finer size categorization). The coccolith size
microfossil record are well represented at least
is maximum in plan view, i.e., the length
since the Triassic (Crucirhabdus minutus being
observed in the light microscope (Fig. 1). Based
one of the earliest representatives from the Nor-
on appearance, major terms are: Minuscule
ian–Rhaetian boundary; Gardin et al. 2012). The
(<1 µm); Very small, 1–3 µm; Small 3–5 µm;
calcareous nannoplankton, through time, have
Medium 5–8 µm; Large 8–12 µm; and Very
proved to be excellent biostratigraphic markers
large >12 µm (Fig. 1). These, mostly disk-
for regional and global correlation (Tables 1, 2,
shaped forms, are produced by marine
3, 4, 5, 6, 7, 8, 9, and 10; the age model used is
golden-brown unicellular photosynthetic algae (a
after Gradstein et al. 2102). Their usefulness as a
living haptophyte, the coccolith-bearing sub-
biostratigraphic tool is largely due to four rea-
group) called coccolithophore (Fig. 2(1)). The
sons: (a) they are abundant in marine sediments
external surface of a coccolithophore cell, called
and thus preserve a continuous paleontological
a Coccosphere, is a composite exoskeleton that is
record (Bown et al. 2004), (b) they are cos-
made up of skeletal plates set in an interlocking
mopolitan in their distribution and thus are
arrangement (Figs. 2(2–8)). Closely related are
excellent tool for regional and global correlation,
another significant and variously shaped cal-
(c) their rapidly evolution through time (appear-
careous nannofossils called the nannoliths
ances and disappearances) also makes them an
(Fig. 3) that are formed by dinoflagellates. Their
excellent tool for subdividing or dating a rock
biological affinity is uncertain due to lack of
strata, and lastly, (d) their small size (<30 µm)
living analogs but they do possess a comparable
means that even a small sample size is enough
distribution similar to the coccoliths.
for analyses, contrary to the need for larger
The present-day coccolithophores live in the
sample size for other fossil forms.
top 200 m of the marine water column (photic
The nannoliths are an assorted artificial group
zone) and play an important role in the food
with a wide range of shapes (Fig. 3). Most of
chain. After death, the individual coccoliths
them show some heterococcolith features, such
separate from the coccolithosphere (Fig. 2(1)) or
as rotational symmetry, complex crystal unit
form part of a fecal pellet, and sink down to the

© Springer Nature India Private Limited 2020 193

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_10
194
Fig. 1 Size of coccoliths. The maximum dimension (in
µm; micron) of the coccosphere in plan view (length). The
terms are based on the appearance in light microscope.
Modified after Young et al. (1997) and with permission
from Dr. Jeremy Young (UK). See text for further
explanation
shape, and chirality; they are, however, distinct
enough. Their relationship to coccolithophores is
uncertain, and considered tentatively either as
modified heterococcoliths evolved from a range
of ancestors (Nannotetrina, Discoaster, Flori-
sphaera) or are modified holococcoliths (Cera-
tolithus) (Bown 1998), but some others may have
10 Calcareous Nannofossils
been formed by haptophytes (unicellular alga
belonging to the division Haptophyta, and
includes all coccolithophores) with calcification
mechanisms unrelated to either mode (Braaru-
dosphaera, Nannoconus) or even by
non-haptophytes (Schizosphaerella) (for details
see Bown 1998).
The terminology used in this chapter is after
several authors but mainly follows Young and
Bown (1997). Prominent among them whose
terms have been incorporated are: Bukry (1969),
Black (1972), Roth and Thierstein (1972), Hay
(1977), Okada and McIntyre (1977), Brasier
(1980), Roth (1973), Norris (1985),
Perch-Nielsen (1985a, b, 1986), Young and
Bown (1991, 1997), Kleijne (1991), Young
et al. (1992, 1997), Young and Westbroek
(1991), Siesser (1993), Young et al. (1994),
Jordan et al. (1995), Bown and Young (1997),
Bown (1998), Young et al. (2003), Bown et al.
(2004), Young and Bown (2014), and Bown
et al. (2017). The classification is dynamic;
hence, the basic framework of Perch-Nielsen in
Bolli et al. (1985a, b) is followed (supplemented
by the illustrations of families). Wherever pos-
sible, the species and genera are updated to a
newer classification of Bown (1998) and this list
is given at the end of the book; updated from the
website www.mikrotax.org/Nannotax3/; cour-
tesy Dr. Jeremy R. Young).
2 Morphology of Coccoliths
On, the basis of their (a) morphology, (b) mode of
formation (calcite element construction), and the
(c) life-cycle stage during which they are
produced, the coccolithophore cells produce
two types of coccoliths—Heterococcoliths and
Holococcoliths. In heterococcoliths, the calcite
elements vary in both size and shape, whereas in
holococcoliths, the calcite elements are of
identical size and shape (*0.1 lm sized euhedral
crystallites). But few structures do not conform
to either of this pattern and are called nannoliths
(Fig. 3). Thus, the coccolithophores are classi-
fied into three groups—heterococcoliths,
2 Morphology of Coccoliths 195

Fig. 2 Coccolith terminology (1–8) as discussed in the text. 1: Cyclococcolithina (a recent coccolithophore) showing
coccoliths. 2: a coccolith showing two differentiated parts; the outer part is higher than the inner part of the coccolith. 3.
Proximal view of a coccolith showing various elements. 4–6: coccosphere showing the placement of shield elements. 7–
8: side view of a coccolith showing the placement of shield elements. 9–10: orientation of a coccolith. 11–13: types
based on the rim having different angles relative to central area and are designated as Murolith (11), Placolith (12), and
Planolith (13). Figures 10–13. Modified after Young et al. (1997) and with permission from Dr. Jeremy Young (UK).
See text for further explanation
196
Fig. 3 Nannoliths. See text for further explanation
holococcoliths, and nannoliths (as originally done
by Braarud et al. 1955; see also Young and Bown
1997).
2.1 Heterococcoliths
A coccolith is built from circular to elliptical and
rarely polygonal disks or rings, constructed from
one or more radial array (cycles) of crystal units
of different sizes and shapes. In most hetero-
coccoliths, the outer part is higher than the inner
part of the coccolith (Fig. 2(2)). This difference
in the outer and inner parts of a coccolith (Fig. 1
(2)) enables the differentiation of a coccolith into
several elements (Fig. 2(3–8)). But before, we
delve into detail, it is important to understand the
10 Calcareous Nannofossils
orientation of a coccolith under study (Fig. 2(9
and 10)).
2.2 Holococcoliths
The fossil record of holococcoliths is sporadic
and sparse, reflecting their low preservation
potential despite their diversity in modern
assemblages. Holococcoliths are disk- or
dome-shaped, formed by numerous, minute
(*0.1 µm), equidimensional calcite crystallites
(Fig. 8(1–5)) of simple shape (as in Coccolithus
pelagicus and Calyptrosphaera sphaeroidea; see
Parke and Adams 1960; Manton and Leedale
1969; Rowson et al. 1986; Klaveness 1973;
Pienaar 1994) and held together by an organic
matrix (Halldal and Markali 1955). Compared to
heterococcoliths, the holococcolith morphologies
are more conservative through time, with less
well-defined divisions between rim and central
area structures (as they are constructed using
identical building blocks; see Fig. 8(1–5)).
The holococcolith surfaces often show
hexagonal arrays of crystals with rhombohedral
faces developed on the crystallites (see Fig. 8(1–
5)); light microscope (LM) observations suggest
that this is because their c-axes are orientated
perpendicular to the surface. Other surfaces,
however, are covered by crystallites lying with a
rhombohedral face parallel to the surface (Fig. 8
(5)), their c-axes oblique to the surface (see also
Gartner and Bukry 1969a, b; Kleijne 1991; Bown
1993; Farhan et al. 1994). Further details of both
their structural and morphological types and
terms used for light microscope (LM) observa-
tions are discussed below (see Fig. 8(8–30)).
3 Terminology for Heterococcoliths
and Nannoliths
3.1 Coccolith Orientation
3.1.1 Proximal
This is directed toward the center of the cocco-
sphere (Fig. 2(9)).
3 Terminology for Heterococcoliths and Nannoliths 197

Table 1 Stratigraphic occurrences of important marker taxa (202–236 Ma)

198 10 Calcareous Nannofossils

Table 2 Stratigraphic occurrences of important marker taxa (175–201 Ma)

3 Terminology for Heterococcoliths and Nannoliths 199

Table 3 Stratigraphic occurrences of important marker taxa (146–157 Ma)

Table 4 Stratigraphic occurrences of important marker taxa (164–173 Ma)

200 10 Calcareous Nannofossils

Table 5 Stratigraphic occurrences of important marker taxa (145–101 Ma)

3 Terminology for Heterococcoliths and Nannoliths 201

Table 6 Stratigraphic occurrences of important marker taxa (100–67 Ma)

202 10 Calcareous Nannofossils

Table 7 Stratigraphic occurrences of important marker taxa (65–56 Ma)

3.1.2 Distal 3.1.8 Horizontal

This is directed toward the outer surface of the This is perpendicular to the proximo-distal
coccosphere (Fig. 2(9)). direction (Fig. 2(10)).

3.1.3 Longitudinal 3.1.9 Internal/Inner/Inward

This direction is parallel to the long axis of an This is toward the center of coccolith (Fig. 2
elliptical/elongated coccolith (Fig. 2(9)). (10)).

3.1.4 Transverse 3.1.10 External/Outer/Outward

This direction is parallel to the short axis of an This is away from center of the coccolith (Fig. 2
elliptical/elongated coccolith (Fig. 2(9)). (10)).

3.1.5 End 3.1.11 Length/Width/Height

This is the edge of the coccolith and is parallel to These are maximum dimensions of a coccolith in
the short axis (Fig. 2(9)). the longitudinal, transverse, and vertical direc-
tions, respectively (Fig. 2(10)).
3.1.6 Side
This is the edge of a coccolith and is parallel to
the long axis (Fig. 2(9)). 3.2 Central Area and Rim

3.1.7 Vertical The coccolith disk or ring is called the rim which
This is the proximo-distal direction (Fig. 2(9)). encloses a central area (Fig. 2(11–13)).
3 Terminology for Heterococcoliths and Nannoliths 203

Table 8 Stratigraphic occurrences of important marker taxa (55–34 Ma)

204 10 Calcareous Nannofossils

Table 9 Stratigraphic occurrences of important marker taxa (33–6 Ma)

3 Terminology for Heterococcoliths and Nannoliths
Table 10 Stratigraphic occurrences of important marker taxa (5Ma–Present)
3.2.1 Central Area
This is the inner part of the coccolith and is
enclosed by the rim (see Fig. 2(10)). In com-
parison with the rim, it possesses less regularly
placed cyclical elements and has inward element
growth. The central area is either entirely closed
or may have a central opening (see Fig. 2(10)).
3.2.2 Rim
This is the outer part of the coccolith and is
marked by regular cycles, with some vertically
directed structures and outward element growth
(Fig. 2(10)).
In heterococcoliths, both central area and rim
are important morphologically differentiated
parts. The central area may be open, virtually
closed, or spanned by different structures,
resulting in great variations in the form of the rim
and designated as Murolith, Placolith, and
Planolith (based on the rim having different
angles relative to the central area) (Fig. 2(11–
13), respectively); the former two are most
common (Fig. 2(11 and 12), respectively). The
Muroliths are characterized by a wall-like, sub-
vertical rim (Fig. 2(11)) and Placoliths have two
subhorizontal shields (distal and proximal), sep-
arated by a tube (Fig. 2(12)).
Murolith (from the Latin Word Murus = Wall)
The rim has a subvertical wall, and hence, the
shape of the coccolith is bowl-shaped (Fig. 2
(11)). Although the rim is elevated, it is without
205
well-developed shields (distal and proximal).
Example: Zeugrhabdotus Reinhardt.
Placolith
The rim has two shields with a tube that separates
them (Fig. 2(12)). The rim has two (distal and
proximal), or more, well-developed shields. Ex:
Coccolithus, Emiliania and Watznaueria.
Planolith (from the Latin Word Planus = Flat)
The rim is low, and hence, the shape of the
coccolith is disk-shaped (Fig. 2(13)). The rim is
not significantly elevated. Ex: Rhabdosphaera
Haeckel and Discoaster Tan.
3.3 Coccolith Outline and Axial Ratio
The coccoliths, in plan view (i.e., outline), occur
in varied shapes, also (Fig. 3(1–16)) such as:
3.3.1 Asymmetrical
These coccoliths lack bilateral symmetry, due to
the presence of a wing or a similar structure
(Fig. 4(1 and 2)). Example: Helicopontosphaera
Hay and Mohler and Helicosphaera Kamptner.
3.3.2 Wing
These are characterized by the presence of the
local extension of rim (Fig. 4(1 and 2)). Exam-
ple: Helicosphaera Kamptner and Kamptnerius
Deflandre.
206 10 Calcareous Nannofossils

Fig. 4 Coccolith shapes in plan view with examples, and their characteristics with axial ration. Modified after Young
et al. (1997) and with permission from Dr. Jeremy Young (UK). See text for further explanation
3 Terminology for Heterococcoliths and Nannoliths
3.3.3 Irregularly Elliptical
These coccoliths possess an almost elliptical shape
(Fig. 4(3 and 4)). Example: Reticulofenestra
3.3.4 Elliptical
These coccoliths are curved with two axes of
symmetry (the alternative terms used in the lit-
erature are oval and ovoid) (Figs. 4(5 and 6)).
Example: Coccolithus pelagicus (Wallich 1877).
3.3.5 Oblong
They have a symmetrical form that is interme-
diate between a rectangle and an ellipse, i.e., the
ends are curved with subparallel sides (Fig. 4(7
and 8)). Example: Pseudoemiliania.
3.3.6 Lenticular
They have a symmetrical form that is interme-
diate between a rhomb and an ellipse; they have
pointed ends (Fig. 4(9 and 10)). Example:
Stradnerlithus bifurcatus Noël.
3.3.7 Polygonal
This form has straight sides (with sharp edges as
triangular, pentagonal, etc.) (Figure 4(11 and
12)). Example: Rhombolithion duodecostatum
(Goy).
Fig. 5 Types of ultrastructural component showing element (2), crystal units (3) and segment (4). Modified after
Young et al. (1997) and with permission from Dr. Jeremy Young (UK). See text for further explanation
207
3.3.8 Reniform
This form is kidney-shaped (Fig. 4(13 and 14)).
Example: Nephrolithus frequens Górka)
3.3.9 Ring-Shaped
These are circular or elliptical forms character-
ized by a narrow rim and a central area that is
open (Fig. 4(15 and 16)). Example: Manivitella
pemmatoidea (Deflandre).
The ratio of coccolith length to width is called
the Axial Ratio and based on this ratio, various
terms of shape are coined for coccoliths, such as
Narrowly elliptical, Normally elliptical, Broadly
elliptical, Subcircular, and Circular (see Fig. 4
(17 and 21)), respectively).
3.4 Ultrastructure
The Element, Crystal unit, and Segment form a
hierarchy of structural components (Fig. 5). The
structures (Fig. 5) provide a key for the catego-
rization at the family and order level. Within a
family, both coccolith structure and crystallog-
raphy are generally constant. Various terms used
to describe the ultrastructural components of a
coccolith are given below.
208 10 Calcareous Nannofossils

3.4.1 Element 3.5.2 Tile

This is a discrete component of a coccolith where
several elements unite to form a crystal unit
(Fig. 5(1 and 2)).
3.4.2 Crystal Unit
This is a fundamental component of a coccolith
and is composed of a group of elements from
different cycles but in crystallographic continuity
(Fig. 5(3)).
3.4.3 Segment
This is a symmetrically repeated part of the
coccolith and includes elements from each cycle,
consisting of one or more crystal units (Fig. 5
(4)).
3.4.4 Contact-Surface
This is the plane of contact between two ele-
ments (Fig. 5(4)). An alternative term for this is
attachment surface.
3.4.5 Cycle
This is the ring of elements or of crystal units
(Fig. 5(4)).
3.4.6 Suture
This is the trace of contact-surface on the surface
of a coccolith (Fig. 5(4)).
This element is broad and thin and appears like a
“tile” (a  b > c) (Fig. 6(2)).
3.5.3 Lath
This element is elongate and wide (longish
rectangle; a > b > c) (Fig. 6(2)).
3.5.4 Rod
This element is elongate and narrow (a > b  c);
it is narrower than the lath (Fig. 6(2)).
3.5.5 Wedge
This element tapers at one end but overall, all three
elements are nearly of the same size (Fig. 6(3)).
3.5.6 Petal/Petaloid Element
This element also tapers on one end (like a petal
of a flower) but is much broader, bigger, and
thinner than the Wedge (Fig. 6(3)).
3.5.7 Ray
This element also tapers on one end but is much
elongated and wide (Fig. 6(3)).
3.5.8 Spine
This element also tapers on one end
(“spine-like”) and elongates, but is much nar-
rower than a ray (Fig. 6(3)).

3.4.7 Lamina 3.5.9 Granule

This is a platy substructure within a crystal unit.
Example: Braarudosphaera.
3.4.8 Tier
This is a set of vertically superposed cycles.
Example: Arkhangelskiella and Lapideacassis.
This element is small and variable-shaped ele-
ment. Example: the blanket elements of Heli-
cosphaera and the spine-forming elements of
Cretarhabdus.
3.6 Element Modifications

3.5 Element Shapes 3.6.1 Kink

The names of the element shapes are a function
of the orientation of the three orthogonal axes, a,
b, and c (Fig. 6(1)).
3.5.1 Block
This element is nearly of the same size
(a  b  c) (Fig. 6(2)).
It is characterized by an angular bend in the
element (Fig. 6(4)). Example: Discorhabdus
criotus Bown.
3.6.2 Offset
It is the displacement of an element from the radial
growth caused by a double kink (Fig. 6(4)). Exam-
ple: Grantarhabdus coronadventis (Reinhardt).
3 Terminology for Heterococcoliths and Nannoliths 209

Fig. 6 Element terminology. Orientation of the three orthogonal axes (1). The element shapes (2–3) are a function of
the orientation of these three axes. Element modifications (4–5), types of openings (6–8), and description of the parts of
rimz (9–10). Modified after Young et al. (1997) and with permission from Dr. Jeremy Young (UK). See text for further
explanation
210
3.6.3 Node
It is a block-shaped projection from the element
(Fig. 6(5)). Example: Discoaster mediosus
Bramlette and Sullivan.
3.6.4 Keel
It is a lath-shaped projection from the element
(Fig. 6(5)).
3.6.5 Ridge
It is a rod-shaped projection that runs all along
the element (Fig. 6(5)). Example: Pontosphaera
multipora (Kamptner).
3.6.6 Tooth
It is a rod- or wedge-shaped projection from the
element (Fig. 6(5)). Example: Eprolithus floralis
(Stradner).
3.7 Types of Openings
3.7.1 Hole
This is an opening that runs in one of the elements
(Fig. 6(6)). Example: Pemma basquense (Martini).
3.7.2 Slit
It is an elongate perforation (Fig. 6(6)). Example:
Gephyrocapsa ericsonii (McIntyre and Bé).
3.7.3 Perforation
It is a small opening present between two or
more elements (Fig. 6(6)). Example: Anthos-
phaera periperforata Kleijne.
3.7.4 Canal
This is a narrow elongated opening within a
coccolith or nannolith. Example: Nannoconus
kamptneri Brönnimann.
3.7.5 Cavity
This is a broad opening within a coccolith or a
nannolith. Example: Polycostella beckmannii
Thierstein.
3.7.6 Depression
This is a pit on the surface of a coccolith or a
nannolith. Example: Micula concava (Stradner).
10 Calcareous Nannofossils
3.8 Element Curvature
3.8.1 Dextrogyre
The elements curve to the right when traced
radially outward (Fig. 6(7)). Example: Cyc1a-
gelosphaera reinhardtii (Perch-Nielsen) Romein.
3.8.2 Laevogyre
The elements curve to the left when traced
radially outward (Fig. 6(8)). Example: Fasci-
culithus involutus Bramlette and Sullivan.
3.8.3 Straight
The elements are not curved.
3.8.4 Rims
This Is Largely Formed of Cyclic Structures
Possessing High Rotational Symmetry (Fig. 6(9
and 10)).
3.9 Parts of Rims
Each of these parts may be formed of a single
cycle of elements, part of a cycle, or several
cycles illustrated in Fig. 6(9 and 10).
3.9.1 Wall
This is a subvertical structure that is not associ-
ated with shields (Muroliths) (Fig. 6(9)). The
wall together with the shield is composed of
several elements that vary from species to spe-
cies. The Chiasmolithus eograndis Perch-Nielsen
may have 60–100 elements each, whereas in
Chiasmolithus bidens (Bramlette and Sullivan),
the range is between 51 and 75 elements.
3.9.2 Flange
It is the subhorizontal protrusion (in the
outward-lateral direction) from the rim (Fig. 6
(9)). Example: Lophodolithus nascens Bramlette
and Sullivan.
3.9.3 Collar
It is the subvertical protrusion from rim (may
occur on proximal or distal surfaces) (Fig. 6(9
and 10)). Example: Rhabdosphaera perlonga
(Deflandre) Bramlette and Sullivan.
3 Terminology for Heterococcoliths and Nannoliths 211

3.9.4 Shield 3.10.2 Tangential

It is a broad (sub-) horizontal structure (Fig. 6 This is the direction on the surface of the base-
(10)). Example: Biscutum castrorum Black. plate parallel to its margin (Fig. 6(1)). It is (as
seen in the distal view) clockwise/dextral/
3.9.5 Tube right or anticlockwise/sinistral/left senses of
It is the subvertical structure between two shields direction.
(Fig. 6(10)).
3.10.3 Vertical
3.9.6 Crown This is the direction perpendicular to the base-
It is the discontinuous/beaded collar. Example: plate—Up/down distal-proximal directions
Prinsius dimorphosus (Perch-Nielsen). (Fig. 6(1)).

3.10.4 Flare and Taper

3.10 Directions on the Rim The divergence of orientation from
horizontal/vertical in the radial direction (Fig. 6
3.10.1 Radial (1); see also Fig. 7(1) and (3)).
This is the direction on the surface of the base-
plate perpendicular to its margin—inward, out- 3.10.5 Taper Surfaces
ward, toward from center, and away from center The convergence upward, producing conical
(Fig. 6(1)). bodies (Fig. 7(1)).

Fig. 7 Element arrangements as seen in side view (1–9). 1–3: element arrangements based on the divergence of
orientation from horizontal/vertical in the radial direction; 4–6: element arrangements based on the divergence from the
horizontal in the tangential direction; 7–9: element arrangements based on the divergence from the horizontal from the
radial direction. 10–12: types of sutures. 13–17: element arrangement in plan view showing different types of coccolith
placement (13–17). Modified after Young et al. (1997) and with permission from Dr. Jeremy Young (UK). See text for
further explanation
212
3.10.6 Flare Surfaces
The divergence upward, producing obconical/
funnel-shaped bodies (Fig. 7(3)).
3.10.7 Element Arrangement as Seen
in Side View
. lowii (Gran and Braarud).
3.10.8 Imbrication
This is the divergence from the horizontal in the
tangential direction (Fig. 7(4) and (6)).
3.10.9 Anticlockwise/Clockwise
Imbrication
This is the offset of the upper part of element
from the lower. Example: anticlockwise (Fig. 7
(9)): Coronosphaera Gaarder; clockwise (Fig. 7
(11)): Pontosphaera Lohmann.
3.10.10 Imbrication Angle
This is the angle of contact-surface from the
horizontal. There are two types, high angle with
subvertical contact-surfaces and low angle with
subhorizontal contact-surfaces. Example:
Gephyrocapsa Kamptner.
3.10.11 Zeugoid Rim
This is characterized by a rim that has
high-angled imbrication but is without distinct
shields. Example: Bownia Varol and Girgis.
3.11 Element Arrangement as Seen
in Plan View
3.11.1 Obliquity
This is the horizontal divergence from the radial
direction (see Fig. 7(7–9)). Example: Cal-
ciopappus caudatus Gaarder and Ramsfjell.
3.11.2 Dextral/Sinistral Obliquity
This is the deflection from the radial of the outer
part of element relative to the inner part, as seen
in distal view. A dextrally oblique cycle displays
clockwise obliquity in distal view but anticlock-
wise obliquity in proximal view (Fig. 7(9)).
Examples: anticlockwise obliquity (chirality; see
Fig. 7(7))—Syracosphaera corolla Lecal;
10 Calcareous Nannofossils
clockwise obliquity (see Fig. 7(9))—Syra-
cosphaera rotula Okada and McIntyre.
3.11.3 Butting
These are elements with simple subradial sutures
(Fig. 7(10)). Example: Braarudosphaera bige-
3.11.4 Interlocking
These are elements with complex sutures
(Fig. 7(11)).
3.11.5 Overlapping
These are elements with low angle oblique su-
tures (Fig. 7(12)).
The orientation of these crystal units is mostly
radial and vertical but some are orthogonal (the
latter arrangement gives chirality to the coccolith
structure). The chirality is expressed in terms of
imbrication and obliquity direction of individual
elements (Fig. 5) which remains the same at the
species level (Didymus and Young 1994). Details
of the structures/terms mentioned above are fur-
ther elaborated and explained below in Sect. 3,
Terminology for Coccoliths and Nannoliths.
3.12 Coccolith Arrangement
3.12.1 Non-Overlapping
The adjacent coccoliths are arranged with edges
directly butting with each other, rather than
overlapping. These types are either non-over-
lapping interlocking, where the adjacent coccol-
iths interlock or are non-overlapping non-
interlocking where the adjacent coccoliths do
not interlock (Figs. 7(13–14)).
3.12.2 Interlocking
In this condition, the adjacent coccoliths inter-
lock (Fig. 7(15)).
3.12.3 Overlapping
In this condition, the adjacent coccoliths overlap
(Fig. 7(16 and 17)).
Both overlapping and interlocking patterns
can occur in varied combinations.
4 Terms Used in Describing the Parts of a Holococcolith
4 Terms Used in Describing
the Parts of a Holococcolith
4.1 Holococcolith
A coccolith that is built by many small-sized
(<0.1 µm) calcite crystallites of similar shape
and size (Fig. 8(1)).
4.1.1 Crystallite
These are individual small-sized (<0.1 µm) cal-
cite crystals forming the coccolith (Fig. 8(2)).
4.1.2 Perforation
These are the opening in the wall (Fig. 8(3)); the
openings are the size of a single calcite crystallite.
4.2 Crystallite Arrangement
This pattern/arrangement of calcite crystallites
noted on the outer surface (Fig. 8(3–5)). Three
types of arrangement are noted: Hexagonal
meshwork, Hexagonal, and Rhomboid.
4.2.1 Hexagonal Meshwork
The crystallites are of the same size with a reg-
ular pattern of perforations. The perforations are
a result of the absence of single crystals (Fig. 8
(3)). Example: Syracosphaera pulchra Lohmann.
4.2.2 Hexagonal
The crystallites are arranged in a hexagonal
pattern, with their c-axes directed radially (Fig. 8
(4)). Example: Syracolithus quadriperforatus
(Kamptner).
4.2.3 Rhomboid
The crystallites are arranged in a rhombohedral
pattern, with their c-axes oblique to the surface
(Fig. 8(5)).
4.3 Plug
This is the distally placed block (Fig. 8(6)).
A spine extends distally and may be closed by a
distal plug. Example: Owenia hillii Crux.
213
4.4 Rim
This is the marginal part best observed in
cross-polarized light that typically has elements
with radial c-axes (Fig. 8(6 and 7).
4.5 Block
A zone of holococcolith that behaves as one unit
in cross-polarized light (Fig. 8(7)). Examples:
Anfractus, Calculites, and Lucianorhabdus.
4.6 Cavity
This is formed when the crystallites do not fill the
open central part of the holococcolith (Fig. 8(8)).
Example: Calyptrosphaera and Zygosphaera.
4.7 Pore
This is the opening within the wall that is larger than
a crystallite (Fig. 8(8)). Example: Gliscolithus.
4.8 Proximal Flange
This is the subhorizontal protrusion from base of
the rim (Fig. 8(8)).
4.9 Proximal Ring
This is the proximal layer(s) of the crystallites that
are confined to the edge of the coccolith (Fig. 8
(8)). Example: Lacunolithus Bown (Nannolith).
4.10 Wall
This is the layer(s) of crystallites that forms the
subvertical structure (Fig. 8(8)).
4.11 Distal Cover
This is the distal layer(s) of crystallites, and it
covers the cavity (Fig. 8(9)). Example: Floscu-
losphaera Jordan and Kleijne.
214 10 Calcareous Nannofossils

Fig. 8 Holococcolith terminology for describing parts of it (1–7), structural (8–10) and morphological types (11–30)
describing the entire holococcolith. Modified after Young et al. (1997) and with permission from Dr. Jeremy Young
(UK). See text for further explanation
4 Terms Used in Describing the Parts of a Holococcolith
4.12 Proximal Plate
This is the proximal layer(s) of crystallites that at
times, almost covers the base of a coccolith
(Fig. 8(9)). Example: Acuturris Wind and Wise.
4.13 Septum
This is the layer(s) of crystallites that forms the
internal wall (Fig. 8(9)).
4.14 Depression
This is a pit on the surface (Fig. 8(10)). It is not
an opening into the cavity. Example: Daktylethra
Gartner.
5 Terms Used in Describing
the Entire Holococcolith
5.1 Cavate
This is characterized by a large cavity within the
rim (Fig. 8(8)). Example: Calyptrosphaera.
5.2 Septate
The space inside the rim is subdivided by a septa
(Fig. 8(9)). Example: Anfractus harrisonii Medd.
5.3 Solid
This is characterized by a coccolith that has a
single mass of crystallites and is without a dis-
tinct cavity or septa. It may or may not possess
depressions, perforations, or pores (Fig. 8(10)).
Example: Helicosphaera carteri (Wallich).
5.4 Morphological Types
5.4.1 Laminolith
These are solid holococcoliths formed by several
(usually more than two) horizontal layers of
215
crystallites (Fig. 8(11–13)). They may be with or
without perforations/pores. Example: Zygo-
sphaera Kamptner.
5.4.2 Crystallolith
These are disk-like solid holococcoliths formed
by one or two layers of crystallites, with a low
rim (Fig. 8(14 and 15)). Example: Coccolithus
pelagicus (Wallich).
5.4.3 Zygolith
These holococcoliths possess a bridge-shaped
distal-cover (Fig. 8(16–18)). Example: Cori-
sphaera Kamptner.
5.4.4 Calicalith
These holococcoliths are open cavate, but with-
out a distal cover (Fig. 8(19 and 20)). Example:
Calicasphaera Kleijne.
5.4.5 Flosculolith
These holococcoliths are cavate with a distal
opening that is partially closed by a vaulted
distal-cover (Fig. 8(21 and 22)). Example:
Flosculosphaera Jordan and Kleijne.
5.4.6 Gliscolith
These holococcoliths are also cavate but with a
bulbous distal part (Figs. 8(23 and 24)). Exam-
ple: Gliscolithus Norris.
5.4.7 Fragariolith
These holococcoliths possess a proximal plate
which has a blade-like process on top of it
(Figs. 8(25 and 26)). Example: Syracosphaera
molischii Schiller.
5.4.8 Calyptrolith
These are domal cavate, with a nearly continuous
distal-cover (Fig. 8(27 and 28)). Example:
Calyptrosphaera Lohmann and Anthosphaera
Kamptner.
5.4.9 Helladolith
These holococcoliths are similar to the zygoliths
but possess a bridge that is expanded distally into
a double-layered leaf-like process (Fig. 8(29 and
30)). Example: Helladosphaera Kamptner.
216 10 Calcareous Nannofossils

5.5 Central Areas Narrow Central Area

The width of the central area is less than one time
The central area structures are very variable in the width of the rim (Fig. 9(7)). Example:
both form and structure (Fig. 9); the individual Reticulofenetra bisecta (Hay, Mohler, and
species often have unique structures. Wade).

Normal Central Area

5.6 Structural Types
5.6.1 Disjunct Bar
This is formed from crystal units that are discrete
from the rim structure (Fig. 9(1)). Example:
Eiffellithus angustus. Examples for plate:
Arkhangelskiella; bar: Coccolithus pelagicus,
Helicosphaera seminulum, and Watznaueria
britannica.
5.6.2 Conjunct Bar
This is formed from crystal units of the rim
structure (Fig. 9(2)). Example: Helicosphaera
walbersdorfensis Müller. These can be in the
shape of bridge and grill (Gephyrocapsa), bar
(Helicosphaera sellii and Watznaueria biporta),
or plate (Kamptnerius).
5.7 Orientation in Profile
5.7.1 Basal
They occur on the proximal surface (Fig. 9(3)).
5.7.2 Elevated
They occur above the proximal
(Fig. 9(4)).
5.7.3 Vaulted
They are cone-shaped, arising from the rim
toward the center (Fig. 9(5)).
5.7.4 Planar
They are flat, but not vaulted (Fig. 9(6)).
The width of the central area is one or two times
the width of the rim (Fig. 9(8)).
Wide Central Area
The width of the central area is more than two
times the width of the rim (Fig. 9(9)). Example:
Crepidolithus perforatus (Medd).
5.8.2 Transverse
The central area is parallel with short axis of
(elliptical) of the coccolith (Fig. 9(10)). Exam-
ple: Biscutum finchii Crux.
5.8.3 Longitudinal
.
5.8.4 Diagonal
The central area appears inclined relative to axes
9 Fig. 9(12–15). The angle is measured from the
traverse direction. Based on this angle, there are
two major types—Low and High angle (Fig. 9
(12 and 13), respectively).
Low Angle
This is near to the transverse direction
surface (Fig. 9(12)).
High Angle
This is near to the longitudinal direction
(Fig. 9(12)).
5.8.5 Sinistral/Dextral
The central area is inclined to the left or right of
the long axis as seen in distal view (Fig. 9(14 and
15), respectively).

5.8 Orientation in Plan View

5.9 Structures Spanning Central Area
5.8.1 Relative Width
The width of central area relative to the width of 5.9.1 Arm
the rim is categorized into three types: See Fig. 9(16).
5 Terms Used in Describing the Entire Holococcolith 217

Fig. 9 Central area structures. Modified after Young et al. (1997) and with permission from Dr. Jeremy Young (UK).
See text for further explanation
218 10 Calcareous Nannofossils

5.9.2 Cross-Bar Closed Central Area

A bar spans the central area (Fig. 9(16)). The central area is without a central opening.

5.9.3 Cross Open Central Area

Part of cross-bar meets in the center (Fig. 9(17)). The central area is without any structures.

Axial Cross (Abb. +) 5.10.2 Grill

The cross-bar is longitudinal and transverse
(Fig. 9(18)). Example: Coccolithus hulliae Bown
and Newsam.
Diagonal Cross (Abb. X)
The cross-bar is diagonal; it may be symmetrical
or asymmetrical with respect to the axes (Fig. 9
(19 and 20), respectively).
A system of bars closing the central area (Fig. 9
(24)). Example: Emiliania.
5.10.3 Net
A mesh-like structure closing the central area
(Fig. 9(25)). An alternative term is Cribrate
central area. Example: Reticulofenestra and
Cribrosphaerella.

Offset Cross 5.10.4 Perforated Plate

The cross with an offset between the arms of one,
or both, of the cross-bars (Chiasmolithus) (Fig. 9
(21)).
5.9.4 Foot
The broadening of the bar as it meets the rim
(Fig. 9(22)). Example: Cruciplacolithus tenius.
5.9.5 Lateral Bar
The bars run from rim to the cross- bar (Fig. 9
(23)). Example: Retecapsa.
5.9.6 Blanket
The covering of small elements on distal side of
the central area. Example: Helicosphaera and
Coccolithus.
5.9.7 Bridge
An elevated bar spanning the central area.
Example: Gephyrocapsa.
It is characterized by a plate with perforations
(Fig. 9(26)). Example: Arkhangelskiella.
5.10.5 Plate
A continuous or nearly continuous structure
closing the central area (Fig. 9(27)).
5.10.6. Processes
Cavity
Wide opening within process (Fig. 9(28)).
Example: Podorhabdus grassei and Algiro-
sphaera robusta.
Protrusion
Broad low process, with height similar to width,
and width near that of entire central area.
Conical
Cone-shaped protrusion (Fig. 9(29)). Example:
Acanthoica.

5.10 Structures Closing Central Area Sacculiform Sac-like

5.10.1 Central Opening
The opening at the center of a coccolith may be
spanned by bars or other central area structures
(Figs. 9(24–27)), but not by a continuous structure
such as a grill (Fig. 9(24)) or plate (Fig. 9(27)).
Protrusion with more or less rounded upper part
(Fig. 8(28)). Example: Algirosphaera.
Labiatiform
The term is for the elongate double-lipped sac-
culiform protrusions.
5 Terms Used in Describing the Entire Holococcolith 219

Spine 6.1 Crystallographic Orientation

Elongated process, height greater than width
(Figs. 9(30–32)).
Styliform
Spine tapers toward the distal end (Fig. 9(30)).
Claviform
Spine has blunt end, without calyx (Fig. 9(31)).
Calicate
Spine is surmounted by a calyx (Fig. 9(32)).
Salpingiform
Spine (or protrusion) trumpet-shaped (Fig. 9
(33)). Example: Discosphaera.
6.1.1 V-Unit
A crystal unit with a subvertical orientation of
Calcite c-axis (Fig. 10).
6.1.2 R-Unit
A crystal unit with subradial orientation of Cal-
cite c-axis, relative to its point of origin (nucle-
ation) on the proto-coccolith ring.
6.1.3 T-Unit
A crystal unit with subtangential orientation of
Calcite c-axis. Example: Braarudosphaeraceae
and Polycyclolithaceae.

Calyx 7 Nannolith Shapes

A flaring structure at the tip of a process (Fig. 9
(32)). Example: Podorhabdus and Papposphaera
Boss
A low process whose height is similar to or less
than the width (an alternative term is Knob).
Process
A general term for any structure that is rising
from the central area.
Stem
A part of a process below the calyx (Fig. 9(32)).
Nannoliths display a wide range of shapes that
are independent of structure. Some of the major
shapes are illustrated in Fig. 11.
7.1 Dibrachiate
These consist of two subparallel arms joined at
one end (such shapes as horseshoe, arrow-head,
and arcuate) (Fig. 11(1–6)). Example: Amau-
rolithus, Ceratolithus, Ceratolithina, and Cera-
tolithoides (except in C. verbeekii).

Canal
A narrow opening that runs along the length of 7.2 Compact
the process (Fig. 9(33)).
These are more or less equidimensional nanno-
Proximal Pore liths and include shapes such as conical (Exam-
An opening of canal, on the proximal side of the ple: Sphenolithus), obconical (i.e., inverted
central area (Fig. 9(33)). cone-shaped; cylindrical (Example: Fasci-
culithus); Example: Conusphaera), and cubic
(Example: Micula) (Fig. 11(7–14)).
6 Crystallography
7.3 Radiate
The crystallographic orientation provides a key
character for determining phylogenetic relation- These shapes have radial symmetry (Fig. 11
ships (Fig. 9). (15–35)).
220 10 Calcareous Nannofossils

Fig. 10 Crystallographic orientation of elements and V-, R- and T-units. 2–4: distal view of Coccolithus pelagicus
showing the intergrowth of V- and R-units, with one pair of units outlined. See text for further explanation. Modified
after Young et al. (1997) and with permission from Dr. Jeremy Young (UK). See text for further explanation
7 Nannolith Shapes 221

Fig. 11 Nannolith shapes. See text for further explanation

7.3.1 Triradiate 7.4 Rod-Shaped

Threefold radial symmetry. Example: Marthas-
terites and Trochasterites (Fig. 11(28 and 32)). These are elongated with no basal disk. Included
shapes are: bladed (Example: Lithraphidites quadra-
7.3.2 Tetraradiate tus and Triquetrorhabdulus carinatus) and (sub)
Fourfold radial symmetry. Example: Micula, \(Example: Microrhabdulus) (Fig. 11(36–39)).
Quadrum, and Nannotetrina (Fig. 11(29 and
33)).
7.5 Central Body
7.3.3 Pentaradiate
Core part of radiate nannolith where elements are
Fivefold radial symmetry. Example: Goniolithus
in contact (Fig. 12).
and Braarudosphaera (Fig. 11(30 and 34)).

7.3.4 Multiradiate 7.6 Free Rays

More than fivefold radial symmetry. Example:
As in many Discoaster species (Fig. 11(31 Parts of radiate nannolith extending beyond
and 35)). central body.
222
Fig. 12 Terminology used in describing species of family Arkhangelskiellaceae (Bukry 1969a, b)
7.6.1 Short Free Rays
Shapes in which the length of the free ray is less
than the radius of the central body; this results in a
rosette-shaped outline. Example: Hayesites irreg-
ularis (Thierstein in Roth and Thierstein, 1972).
7.6.2 Long Free Rays
Shapes in which the length of free ray is greater
than the radius of central body; this results in a
star-shaped outline.
7.7 Convex Outline
Shapes without a free ray (as in Braaru-
dosphaera). Included shapes are: triangular,
square, and pentagonal shapes.
7.8 Stellate
Shapes with free rays (including rosette and star-
shaped). Example: Micrantholithus and Discoaster.
10 Calcareous Nannofossils
8 Additional Taxon–Based
Terminology
8.1 Braarudosphaeraceae
The nannolith Family Braarudosphaeraceae
Deflandre ranges from Early Cretaceous to pre-
sent day (Table 11). They are of minimal bios-
tratigraphic relevance but often occur in large
numbers, as in the Early Danian just after the
Cretaceous–Tertiary boundary (K/T boundary)
and in the Oligocene of the South Atlantic. Some
of the terms specifically used in describing their
morphology are briefly enumerated below (major
terms only) and illustrated in Fig. 13(1–5).
8.1.1 Pentalith
A nannolith formed by the Braarudosphaeraceae
(this does not include other nannoliths formed of
five elements, such as in Discoaster pentara-
diatus) (Fig. 13(1–5)).
8 Additional Taxon–Based Terminology 223

Fig. 13 Terminology used in describing species of Braarudosphaeraceae Deflandre (1947), Ceratolithaceae Norris
(1965), and Discoasteraceae Tan (1927). Modified from Bolli et al. (1989)
224
8.1.2 Segment
It is one of the five component parts of a pentalith
(Fig. 13(1 and 3)). These appear to be single
crystal units, with a tangential c-axis orientation.
8.1.3 Lamina
Plate-like subelement of a segment (Fig. 13(2)).
8.2 Ceratolithaceae
The Ceratolithaceae is a biostratigraphically
useful nannolith family (see Tables 1, 2, 3, 4, 5,
6, 7, 8, 9, and 10). The group is extant and ranges
from the Late Miocene to Recent (Table 11).
Major terms are enumerated below, whereas
others are illustrated in Fig. 13(6–12).
8.2.1 Orientation
Ceratolith
These are dibrachiate nannoliths (Fig. 13(6 and
7)) including Amaurolithus (Fig. 13(10)).
Anterior/Posterior
Closed end is designated as anterior (Fig. 13(6)).
Left/Right
Based on upper view, and looking toward the
anterior end.
8.2.2 Parts
Apical Region
Anterior end of a ceratolith (also called apical
node) (Fig. 13(8)).
Arch
Part of apical region connecting the two arms
(Fig. 13(7)).
Arm
Rod-like extension back from apical region
(Fig. 13(11 and 12)).
10 Calcareous Nannofossils
Rod
Rod-shaped structure attached to the nannolith
(Fig. 13(10)). Example: Amaurolithus bizarrus.
Spur
Projection from apical region (Fig. 13(8)).
Keel
Lath-like structure running along an arm (Fig. 13
(9)). Keel can be dentate or smooth.
Dentate Keel
keel formed of subparallel teeth.
Smooth Keel
keel without teeth.
Tooth
Rod-like subelement of a keel.
Wing
Plate-like extension from main body of nanno-
lith. Example: Amaurolithus ninae.
8.3 Discoasters
Dicoasters are the most important single group of
Cenozoic nannoliths, ranging from the Paleocene
to the Pliocene and are of great biostratigraphic
use (see Tables 1, 2, 3, 4, 5, 6, 7, 8, 9, and 10).
Major terms are enumerated below, whereas
others are illustrated in Fig. 13(13–19).
8.3.1 Ray-Related Terms
See Fig. 13(13–17).
Ray
This is a disk element (Fig. 13(13 and 14)).
Free Ray
Part of ray that protrudes beyond the central area
(Fig. 13(13)).
8 Additional Taxon–Based Terminology 225

Ray Tip Sutural Ridge

Outermost part of ray (Fig. 13(14)). Ridge running along suture (Fig. 13(16)).

Bifurcate Tip Proximal

Ray tip divides into two bifurcations (Fig. 13
(16)). Example: Discoaster variabilis.
Simple Tip
Ray tip without bifurcation or proximal exten-
sion (Fig. 13(14)).
Proximal Extension
Continuation of the ray downward from the tip
(Fig. 13(17)). Example: Discoaster brouweri.
8.3.2 Other Useful Terms
Concave side of eudiscoaster (Fig. 13(17)).
Distal
Convex side of eudiscoaster (Fig. 13(16 and 17)).
Laevogyral Face
Side of heliodiscoaster showing left-handed
curvature of rays (Fig. 13(18)).
Dextrogyral Face
Side of heliodiscoaster showing right-handed
curvature of rays (Fig. 13(19)).

Central Area
Portion of Discoaster with rays in contact 8.4 Fasciculiths
(Fig. 13(13)).
Fasciculiths (Family Fasciculithceae) (Fig. 14(1–
Contact-Surface 5)) are an extinct Paleogene group of nannoliths,
Surface between adjacent elements (alternative ranging from the Paleocene to the Early Eocene
term attachment surface). (Table 11). Major terms are enumerated below,
whereas others are illustrated in Fig. 14(1–5).
Disk
Main part of Discoaster, excluding bosses or 8.4.1 Orientation
stems (Fig. 13(15)). The concave end of the nannolith is assumed to
be proximal (Fig. 14(1)).
Boss
Low distal or proximal protrusion from center of Apical Cycle
disk (alternative term knob) (Fig. 13(15)). Distal cycle of fasciculith. (An alternative term is
Cone) (Fig. 14(1–3)).
Rosette-Shaped
Discoaster with short free rays (Fig. 13(18 and Central Body
19)). Optically distinct body occurring in the center of
fasciculith (Fig. 14(2 and 5)).
Segment
Ray and associated boss or stem elements. Column Cycle
Proximal cycle of fasciculith, usually forms most
Stem of the fasciculith (Fig. 14(4)).
High distal or proximal protrusion from center of
disk (Fig. 13(15)). Example: Discoaster Fenestra
kuepperi. Regular depression on fasciculith (Fig. 14(3)).

Star-Shaped Longitudinal Ridge

Discoaster with long free rays (Fig. 13(13)). Ridge parallel to length of fasciculith.
226 10 Calcareous Nannofossils

Fig. 14 Terminology used in describing species of Fasciculithaceae Hay and Mohler (1967), Helicopontosphaeraceae
Black (1971a), Nannoconaceae Deflandre (1959), and Prediscosphaeraceae Rood, Hay, and Barnard (1971). Modified
from Bolli et al. (1989)
8 Additional Taxon–Based Terminology
Proximal Surface
Lower surface of fasciculith (Fig. 14(2)).
8.5 Helicosphaera
The Helicosphaeraceae is an important hetero-
coccolith family ranging from the Early Eocene
to the present day; the extant species include
Helicosphaera carteri and H. pavimentum. There
is only one genus but many fossil species, many
of them have biostratigraphic relevance, but only
as secondary markers (Tables 1, 2, 3, 4, 5, 6, 7,
8, 9 and 10). Major terms are enumerated below,
whereas others are illustrated in Fig. 14(6–10).
8.5.1 Orientation
The asymmetry of helicoliths (nannolith formed
by the Helicosphaeraceae) allows more-precise
description of orientation than for most other
coccoliths (Fig. 14(7–10)).
Anterior
End with origin of flange and usually with
broader flange on distal side often with distinct
wing or spur. An alternative term is Pterygal
(Fig. 14(10)).
Posterior
Opposite end to anterior. An alternative term is
Antipterygal (Fig. 14(10)).
Dextral/Sinistral
Right/Left sides of helicolith as seen in distal
view with anterior end upwards. The wing, when
present is on the sinistral side (Fig. 14(7 and 9),
respectively).
8.5.2 Parts
Bar
A structure crossing the central opening (Fig. 14
(9 and 10)).
Conjunct Bar
Developed from rim elements (Fig. 14(6)).
Alternative terms include optically continuous
bar, bar, bridge. Example: Helicosphaera carteri.
227
Disjunct Bar
Formed from elements that are discrete from the
rim. Alternative terms are optically discontinuous
bar and bridge. Example: H. intermedia.
Normally Oriented Bar
Diagonal bar with dextral orientation, i.e., rotated
to the right of the long axis in distal view/anterior
end on opposite side to the wing (Fig. 14(9)).
Inversely Oriented Bar
Diagonal bar with sinistral orientation, i.e.,
rotated to the left of the long axis in distal view/
anterior end on same side as wing (Fig. 14(10))
Blanket
Mass of elements forming distal cover (Fig. 14(8
and 9)).
Flange
Rim structure of helicolith (shield is also used by
some workers) (Fig. 14(7–9)).
Origin
Location of first/shortest elements of flange on
the proximal side (Fig. 14(7)).
Proximal Plate
Inward radiate elements on proximal side of
central area (Fig. 14(7 and 8)).
Spur
Spike-like expansion of flange near its termina-
tion (Helicosphaera recta) (Fig. 148–11)).
Termination
Location of last elements of flange on the distal
side (Fig. 14(9)).
Wing
Broad expansion of flange near its termination
(Fig. 14(9)). Example: H. carteri.
8.6 Nannoconids
The Nannoconaceae is a monogeneric group
spanning Late Jurassic to Late Cretaceous of
228 10 Calcareous Nannofossils

uncertain affinities (Fig. 14(12)). Major terms are
enumerated below, whereas others illustrated in
Fig. 14(12).
8.6.1 Central Opening/Central Cavity
This is an opening that runs longitudinally all
through the nannoconid (Fig. 14(12)). Types
include: Canal narrow, <1 µm, and Cavity—
wide, >1 µm.
8.7.2 Apical Cycle
Upper part of sphenolith, formed of most steeply
inclined cycle of elements (Fig. 15(3)). Three
types are noted:
Monocrystalline
These are formed by a single crystal unit.
Example: Sphenolithus heteromorphosus and S.
belemnos.

8.6.2 Aperture Duocrystalline

It is an expression of the central opening at the These are formed by two crystal units. Example:
ends of the specimen (Fig. 14(12)). S. distentus and S. furcatulithoides.

8.6.3 Bulb Compound

It is a distinct swelling of the outline. Example: These are formed by several crystal units.
single: N. borealis; double: Nannoconus Example: S. radians and S. abies.
paskentiensis; triple: N. multicadus.
8.7.3 Apical Spine
8.6.4 Constriction This is the elongate extension of the apical cycle
It is an external indentation of the wall, between (Figs. 15(2 and 3)).
bulbs.
8.7.4 Blade
8.6.5 Flange This is one of three subparts of an element and is only
It is a horizontal projection around the end of a noted in well-preserved specimens (Fig. 15(1)).
nannoconid.
8.7.5 Core
8.6.6 Wall This is the center of radiation of elements
A structure that encloses the central opening (Fig. 15(6)).
(Fig. 14(12)).
8.7.6 Element
This is the basic component of a sphenolith; each
8.7 Sphenoliths element is a single crystal unit.

Sphenoliths (formed by the Sphenolithaceae) 8.7.7 Lateral Cycles

(Fig. 15(1–6)) are an extinct group of nannoliths
that are classified under a single genus Spheno-
lithus ranging from the Paleocene to the Plio-
cene) (Table 11). They are of great
biostratigraphic value during Late Oligocene and
Early Miocene (Table 9).
It is the cycles between apical and proximal cy-
cles, but is not always present (Fig. 15(1)).
8.7.8 Proximal Cycle
It is the lowermost cycle of elements (Fig. 15
(1)).

8.7.1 Orientation 8.7.9 Upper/Lower Quadrant

The concave end of the nannolith is assumed to It is the part above or below the core (Fig. 15
be proximal/basal (Fig. 15(1)). (6)).
9 Brief Note on Classification and Illustration of Major Families 229

Fig. 15 Terminology used in describing species of Sphenolithaceae Deflandre (1952), Stephanolithiaceae Black
(1968), Triquetrorhabdulaceae Lipps (1969), and Zygodiscaceae Hay and Mohler (1967). Modified from Bolli et al.
(1989)

nannoliths generally prevents any standardized

9 Brief Note on Classification
and Illustration of Major Families
The bipartite differentiation in nannofossil clas-
sification (coccoliths and nannoliths) is funda-
mental, although the significance of the
heterococcolith/holococcolith division is equally
indispensable (see Young and Bown 1997; Bown
1998). The morphological variability of
taxonomic approach, and they are best treated on
a group-by-group basis. Coccolith rim structure
is the most significant and conserved morpho-
logical feature and most useful for familial and
higher-level classification. Fine details of rim
construction, central area size, and central area
structure enable generic- and specific-level clas-
sification. At family level, coccolith morphology
is the major criterion for classification (Bown and
230
Young 1997; Young and Bown 1997; Hay 1977;
Perch-Nielsen 1985a, b). At order level, how-
ever, there is no consensus and there is a sig-
nificant gulf between the ways in which they are
applied in fossil and living material (see also the
classification after Perch-Nielsen in Bolli et al.
1985a, b; Hay 1977; Tappan 1980).
10 Geological History
About 250 coccolithophore species are extant
(Young et al. 2003), and off them, about 50 are
known from the deep-sea sediments (Bown et al.
2004). Calcareous nannofossils first appeared in
the latest Triassic (Norian–Rhaetian boundary;
Gardin et al. 2012) as abundant but low-diversity
assemblages, restricted to low latitudes. All but
one species of coccolith disappeared during the
Triassic–Jurassic boundary extinction event.
Fig. 16 Calcareous nannofossil diversity through time (after Bown et al. (2004) and Bown and Young (1997))
10 Calcareous Nannofossils
Thereafter, they were rapidly re-established in
the earliest Jurassic and subsequently colonized
all marine environments. The Early Jurassic
radiation saw the establishment of most major
coccolith families (Fig. 16). Thereafter,
throughout the Jurassic and Cretaceous, diversity
increased steadily with a diversity peak in the
Late Cretaceous. The Cretaceous/Tertiary
boundary mass-extinction event saw the demise
of around 90% of species. Subsequently, they
recovered rapidly in the Early Paleocene, reach-
ing a second diversity peak in the Middle
Eocene. The Paleocene radiation was more rapid
and diverse than the Mesozoic radiations and
established significantly different coccolith fam-
ilies accompanied by numerous new nannolith
groups. There followed a significant decline into
the Oligocene, increased diversity in the Miocene
and a general decrease into the Pleistocene
(Fig. 16).
10 Geological History 231

The distribution of coccolithophore families families discussed in the text is provided in

and nannolith groups is given in Fig. 17 (after Figs. 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28,
Bown et al. 2004) and illustration of represen- 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41,
tative calcareous nannofossil species of various 42, and 43.

Fig. 17 Distribution of coccolithophore families and nannolith groups (after Bown et al. 2004)
232 10 Calcareous Nannofossils

Fig. 18 1–16: Representative calcareous nannofossil species of Arkhangelskiellaceae Bukry (1969); 17–18:
Kamptneriaceae Bown and Hampton (1997) in Bown and Young (1997). 1–3: Arkhangelskiella cymbiformis
Vekshina; 4–5: Arkhangelskiella specillata Vekshina; 6: Arkhangelskiella cribrata (Gazdzicka); 7–9: Broinsonia parca
parca (Stradner); 10: Broinsonia parca constricta Hattner et al.; 11: Broinsonia parca constricta Hattner et al.; 12:
Broinsonia parca expansa Wise and Watkins; 13: Broinsonia parca expansa Wise and Watkins; 14: Thiersteinia
ecclesiastica Wise and Watkins; 15–16: Thiersteinia Wise and Watkins; 17–18: Gartnerago Bukry; 18: Proximal view.
A. = Arkhangelskiella; B. = Broinsonia. 1: Light microscope image; 2–18. Figures adapted from scanning electron
microscope (SEM) images. For systematic details refer to Nannotaxa (www.ina.tmsoc.org/Nannotax3)
10 Geological History 233

Fig. 19 1–17: Representative calcareous nannofossil species of Axopodorhabdaceae Bown and Young (1997). 1:
Cribrosphaerella Deflandre in Piveteau; 2: Cribrosphaerella daniae Perch-Nielsen; 3: Nephrolithus Górka; 4:
Cribrocorona Perch-Nielsen; 5: Ethmorhabdus hauterivianus (Black); 6–9: Axopodorhabdus albianus (Black); 10:
Axopodorhabdus atavus (Grün et al.); 11–12: Axopodorhabdus cylindratus (Noël); 13–14: Axopodorhabdus
dietzmannii (Reinhardt); 15–16: Axopodorhabdus rahla (Noël); 17: Boletuvelum Wind and Wise; A. = Axopodorhab-
dus; 8: Light microscope image. All other figures are adapted from scanning electron microscope (SEM) images. For
systematic details refer to Nannotaxa (www.ina.tmsoc.org/Nannotax3)
234 10 Calcareous Nannofossils

Fig. 20 Representative calcareous nannofossil species of Biscutaceae Black, 1971a. 1–3: Biscutum dubium (Noël); 4:
Discorhabdus exilitus (conical) Noël; 5: Discorhabdus patulus (Deflandre); 6–8: Podorhabdus grassei Noël; 9–11:
Discorhabdus corollatus Noel; 12: Discorhabdus Noël; 13: Sollasites pristinus Noël; 14: Sollasites bipolaris Rood,
Hay and Barnard; 15: Sollasites concentricus Rood, Hay and Barnard; 16: Sollasites lowei (Bukry) Rood, Hay and
Barnard; 17: Sollasites arcuatus Black; 18: Sollasites falklandensis Filewicz et al.; 19–20: Sollasites horticus Stradner
and Adamiker; 21: Sollasites thiersteinii Filewicz et al.; 22: Sollasites granulifera Black. 23: Crucibiscutum pinnatus
(Black); 24: Crucibiscutum hayi (Black). All figures are adapted from scanning electron microscope (SEM) images.
For systematic details refer to Nannotaxa (www.ina.tmsoc.org/Nannotax3)
10 Geological History 235

Fig. 21 1–22: Representative calcareous nannofossil species of Braarudosphaeraceae Deflandre (1947). 1: Braaru-
dosphaera Deflandre; 2: Braarudosphaera batilliformis Troelsen and Quadros; 3–4: Braarudosphaera hayi (Bukry); 5:
Braarudosphaera quinquecostata Hill; 6: Braarudosphaera stenorhetha Hill; 7: Braarudosphaera primula Black; 8–
10: Braarudosphaera africana Stradner; 11: Braarudosphaera hockwoldensis Black; 12: Braarudosphaera bigelowii
(Gran and Braarud); 13: Braarudosphaera regularis Black; 14: Braarudosphaera turbine Stradner; 15: Micrantholithus
Deflandre; 16: Micrantholithus hoschulzii (Reinhardt); 17: Micrantholithus obtusus Stradner; 18: Micrantholithus
speetonensis Perch-Nielsen; 19: Micrantholithus attenuates Bramlette and Sullivan; 20: Micrantholithus belgicus Hay
and Towe; 21: Microrhabdulus decoratus Deflandre; 22: Trapezopentus sarmatus Wind and Cepek; B: Braaru-
dosphaeraceae; M. = Micrantholithus. All figures are adapted from scanning electron microscope (SEM) images. For
systematic details refer to Nannotaxa (www.ina.tmsoc.org/Nannotax3)

Fig. 22 1–23: Representative calcareous nannofossil species of Calciosoleniaceae Kamptner (1927). 1–3: Calciosole- c
nia brasiliensis (Lohmann); 4: Rhombolithion duodecostatum (Goy); 5: Rhombolithion horrellii (Rood and Barnard); 6:
Rhombolithion rotatum (Rood, Hay and Barnard); 7: Rhombolithion octocostatum (Rood and Barnard); 8:
Rhombolithion rhombicum (Stradner and Adamiker); 9: Rhombolithion speetonensis (Stradner and Adamiker); 10:
Stradnerlithus bifurcatus Noël; 11: Corollithion impontus Grün; 12: Corollithion kennedyi Crux; 13: Corollithion
signum Stradner; 14: Corollithion? madagaskarensis Perch-Nielsen; 15: Corollithion exiguum Stradner; 16:
Corollithion completum Perch-Nielsen; 17: Crepidolithus crecifer Prins; 18: Crepidolithus crassus (Deflandre); 19–
20: Cylindralithus serratus Bramlette and Martini; 21: Cylindralithus oweinae Perch-Nielsen; 22: Cylindralithus nudus
Bukry; 23: Cylindralithus duplex Perch-Nielsen. 24–52: Representative calcareous nannofossil species of Calyptro-
sphaeraceae Boudreaux and Hay. 24: general morphology; 25: Anfractus Medd; 26: Semihololithus Perch-Nielsen; 27:
Petrobrasiella Troelsen and Quadros; 28: Athenagalea Hattner and Wise; 29: Octolithus Romein; 30: Metadoga Wind
and Cepek; 31: Acuturris Wind and Wise; 32: Lucianorhabdus Deflandre; 33: Isocrystallithus Verbeek; 34: Munarinus
Risatti; 35: Multipartis Risatti; 36: Pharus Wind and Wise; 37: Calculites Prins and Sissingh; 38: Russellia Risatti; 39:
Okkolithus Wind and Wise; 40: Ramsaya Risatti; 41: Orastrum Wind and Wise; 42: Ottavianus Risatti; 43–45:
Zygrhablithus Deflandre; 46–47: Lucianorhabdus Deflandre; 48: Calculites obscurus (Deflandre); 49–50: Lucianorhab-
dus maleformis Reinhardt; 51–52: Lucianorhabdus cayeuxii Deflandre
236 10 Calcareous Nannofossils
10 Geological History 237

Fig. 23 1–25: Representative calcareous nannofossil species of Calyptrosphaeraceae Boudreaux and Hay (1969). 1:
Calciosolenia Gran; 2–3: Calculites obscurus (Deflandre); 4–5: Calculites ovalis (0 and 45º); 6: Orastrum campanensis
(Cepek); 7: Orastrum asarotum Wind and Wise; 8: Petrobrasiella venata Troelsen and Quadros; 9: Semihololithus
priscus Perch-Nielsen; 10: Acuturris scotus (Risatti, 1973) Wind and Wise; 11: Semihololithus bicornis Perch-Nielsen;
12: Lucianorhabdus inflatus Perch-Nielsen; 13: Lucianorhabdus windii Hattner and Wise; 14: Lucianorhabdus
arborius Wind and Wise; 15–16: Lucianorhabdus maleformis Reinhardt; 17–18: Lucianorhabdus arcuatus
Forchheimer; 19–20: Lucianorhabdus cayeuxii Deflandre; 21–22: Lucianorhabdus compactus (0 and 45º) (Verbeek);
23–24: Lucianorhabdus quadrifidus (0 and 45º) (Verbeek); 25: Lucianorhabdus quadrifidus Forchheimer
238 10 Calcareous Nannofossils

Fig. 24 1–23: Representative calcareous nannofossil species of Chiastozygaceae Rood, Hay and Barnard (1973). 1–2:
Crucirhabdus Rood, Hay and Barnard; 3: Staurorhabdus elongatum (Medd); 4: Staurolithites stradneri (Rood, Hay and
Barnard); 5: Staurolithites acutifera (Vekshina); 6–8: Staurolithites Caratini; 9: Staurolithites angustus (Stover); 10:
Turbirhabdus Prins; 11: Bukrylithus Black; 12–13: Vacheruvillius Goy; 14: Ahmuellerella Reinhardt; 14: Chiastozygus
platyrhethus Hill; 15–16: Heteromarginatus Burky; 17: Diadorhombus Worsley; 18: Diadorhombus rectus Worsley;
19: Rhombolithion minutum Rood, Hay and Barnard; 20: Chiastozygus Gartner; 21: Chiastozygus platyrhethus Hill; 22:
Stradnerlithus primitus Rood, Hay and Barnard; D. = Diadorhombus
10 Geological History 239

Fig. 25 1–17: Representative calcareous nannofossil species of Coccolithaceae Poche (1913). 1–3: Calcidiscus
Kamptner; 1: Distal view; 2: Profile view; 3: Oblique profile view; 4–5: Cruciplacolithus tenuis (Stradner); 6:
Coccolithus formosus (Kamptner); 7: Coccolithus gigas Bramlette and Sullivan; 8: Chiasmolithus titus Gartner; 9:
Chiasmolithus californicus (Bramlette and Sullivan); 10: Chiasmolithus consuetus (Bramlette and Sullivan); 11–13:
Chiasmolithus oamaruensis (Deflandre); 14: Chiasmolithus bidens (Bramlette and Sullivan); 15: Chiasmolithus
expansus (Bramlette and Sullivan); 16–17: Chiasmolithus solitus (Bramlette and Sullivan); 18: Ceratolithoides
Bramlette and Martini; 18–34: Ceratolithaceae Norris; 19–20: Ceratolithoides arcuatus Prins and Sissingh; 21–22:
Ceratolithoides aculeus (Stradner); 23–34: Ceratolithus Kamptner; 25: Ceratolithoides aculeus (<90º); 26:
Ceratolithoides aculeus (>90º); 27–28: Ceratolithoides verbeekii Perch–Nielsen; 29: Ceratolithoides kamptneri
Bramlette and Martini; 30: Ceratolithina hamata Martini; 31: Ceratolithus cristatus (Kamptner)
240 10 Calcareous Nannofossils

Fig. 26 1–16: Representative calcareous nannofossil species of Cretarhabdaceae Thierstein (1973). 1: Cretarhabdus
Bramlette and Martini; 2: Miravetesina Grün; 3: Retecapsa Black; 4: Retecapsa crenulata (Bramlette and Martini); 5–6:
Retecapsa escaigii (Noël); 7: Flabellites Thierstein; 8: Helenea Worsley; 9: Grantarhabdus Black; 10: Cretarhabdus
Bramlette and Martini; 11: Cruciellipsis Thierstein; 12: Speetonia Black; 13–14: Cruciellipsis cuvillieri (Manivit); 15–
16: Speetonia colligata Black
10 Geological History 241

Fig. 27 1–39: Representative calcareous nannofossil species of Discoasteraceae Tan (1927). 1–6: Discoaster Tan; 7:
Discoaster pentaradiatus Tan; 8–10: Discoaster surculus Martini and Bramlette; 11–12: Discoaster deflandrei
Bramlette and Riedel; 13: Discoaster barbadiensis (Tan); 14–15: Discoaster lodoensis Bramlette and Riedel; 16–17:
Discoaster sublodoensis Bramlette and Sullivan; 18–19: Discoaster variabilis Martini and Bramlette; 20–21:
Discoaster challengeri Bramlette and Riedel; 23–25: Discoaster brouweri Tan; 26–27: Discoaster pentaradiatus Tan;
28: Discoaster tani Bramlette and Riedel; 29: Discoaster nobilis Martini; 30: Discoaster mohleri Bramlette and
Percival; 31: Discoaster saipanensis Bramlette and Riedel; 32: Discoaster kugleri Martini and Bramlette; 33–34:
Discoaster multiradiatus Bramlette and Riedel; 35: Discoaster hamatus Martini and Bramlette; 36: Discoaster
quinqueramus Gartner; 37: Discoaster asymmetricus Gartner; 38: Discoaster druggii Bramlette and Wilcoxon
242 10 Calcareous Nannofossils

Fig. 28 1–11: Representative calcareous nannofossil species of Eiffellithaceae Reinhardt (1965). 1–3: Eiffellithus
Reinhardt, 1965; 1: Profile view; 2: Proximal view; 3: Distal view. 4: Eiffellithus collis Hoffmann; 5: Eiffellithus eximius
(Stover); 6: Eiffellithus parallelus Perch-Nielsen; 7: Eiffellithus turriseiffelii (Deflandre); 8: Eiffellithus gorkae
Reinhardt; 9: Manivitella inversus Black; 10: Tegumentum stradneri Thierstein. 11–14: Representative calcareous
nannofossil species of Fasciculithaceae Hay and Mohler (1967). 11–13: Fasciculithus Bramlette and Sullivan; 12:
Lateral view; 12–13: Sectional views; 14: Fasciculithus tympaniformis Hay and Mohler. 15–20: Representative
calcareous nannofossil species of Heliolithaceae Hay and Mohler (1967). 15–17: Heliolithicus Bramlette and Sullivan
1961; 15: Frontal view; 16: Lateral view; 17: Sectional view; 18–19: Heliolithus kleinpelli Sullivan; 20: Heliolithus
riedelli Bramlette and Sullivan
10 Geological History 243

Fig. 29 1–40: Representative calcareous nannofossil species of Helicosphaeraceae Black 1971b (size is fundamental
for species identification) . 1–2: Helicosphaera Kamptner, 1954. 3–6: Helicosphaera carteri (Wallich); 7:
Helicopontosphaera? subantarctica Edwards and Perch–Nielsen; 8: Helicosphaera pacifica Müller and Brönnimann;
9: Helicosphaera papillata (Bukry and Bramlette); 10: Helicosphaera ampliaperta Bramlette and Wilcoxon; 11:
Helicosphaera scissura Miller; 12: Helicopontosphaera minima Martini; 13–15: Helicosphaera walbersdorfensis
Müller; 16: Helicosphaera gartae Bukry; 17: Helicosphaera orientalis Black; 18: Helicosphaera hyaline Gaarder; 19:
Helicosphaera wallichii (Lohmann); 20: Helicosphaera granulata (Bukry and Percival); 21: Helicosphaera kampteri;
22: Helicosphaera inversa (Gartner); 23: Helicosphaera obliqua Bramlette and Wilcoxon; 24: Helicosphaera orientalis
Black; 25: Helicosphaera perch-nielseniae (Haq); 26: Helicosphaera truempyi Biolzi and Perch-Nielsen; 27:
Helicosphaera willcoxonii; 28: Helicosphaera seminulum Bramlette and Sulliva; 29: Helicosphaera mediterranea
Muller; 30: Helicosphaera recta (Haq); 31: Helicosphaera compacta Bramlette and Wilcoxon; 32: Helicosphaera
rhomba (Bukry); 33: Helicosphaera intermedia; 34: Helicosphaera euphratis Haq; 35: Helicosphaera bramlettei
(Muller); 36–39: Helicosphaera reticulata Bramlette and Wilcoxon; 40: Helicosphaera lophota (Bramlette and
Sullivan); 41: Helicosphaera vedderi Bukry; 42: Helicosphaera sellii (Bukry and Bramlette)
244 10 Calcareous Nannofossils

Fig. 30 Insertae sedis. 1: Marthasterites Deflandre; 2–6: Marthasterites furcatus (Deflandre); 7–8: Petrarhabdus
copulatus (Deflandre); 9: Annulithus arkelli Rood, Hay and Barnard; 10: Tribrachiatus orthostylus Shamrai; 11:
Tribrachiatus contortus (Stradner)
10 Geological History 245

Fig. 31 1–24: Representative calcareous nannofossil species of Microrhabdulaceae Deflandre (1963). 1: Lithraphidites
carniolensis Deflandre; 2: Lithraphidites carniolensis serratus Shumenko; 3: Lithraphidites carniolensis carniolensis
Deflandre; 4: Lithraphidites grossopectinatus Bukry; 5–7: Lithraphidites quadratus Bramlette and Martini; 8:
Lithraphidites kennethii Perch-Nielsen; 9: Lithraphidites pseudoquadratus Crux; 10–11: Lithraphidites acutus var.
acutus Verbeek and Manivit; 12: Lithraphidites alatus Thierstein; 13: Lithraphidites praequadratus Roth; 14:
Lithraphidites bollii (Thierstein); 15: Pseudolithraphidites quattuorba cillatus Keupp; 16: Pseudolithraphidites
multibacillatus Keupp; 17–19: Lithostromation Deflandre; 20: Pseudomicula quadrata Perch-Nielsen; 21: Microrhab-
dulus undosus Perch-Nielsen; 22–24: Microhabdulus; L. = Lithraphidites
246 10 Calcareous Nannofossils

Fig. 32 1–38: Representative calcareous nannofossil species of Nannoconaceae Deflandre (1959). 1: Nannoconus
truitti rectangularis Deres and Acheriteguy; 2: Nannoconus truitti frequens Deres and Acheriteguy; 3: Nannoconus
truitti truitti Bronnimann; 4: Nannoconus fragilis Deres and Acheriteguy; 5: Nannoconus donnatensis Deres and
Acheriteguy; 6: Nannoconus cornuta Deres and Acheriteguy; 7: Nannoconus quadratus (Noel); 8: Nannoconus
farinacciae Bukry; 9: Nannoconus regularis Deres and Acheriteguy; 10: Nannoconus multicadus Deflandre and
Deflandre; 11: Nannoconus elongatus Bronnimann; 12: Nannoconus colomii (de Lapparent); 13: Nannoconus
inconspicuus Deflandre and Deflandre; 14: Nannoconus abundans Stradner and Griin; 15: Nannoconus borealis
Perch-Nielsen; 16: Nannoconus bermudezii Bronnimann; 17: Nannoconus quadriangulus quadriangulus Deflandre and
Deflandre; 18: Nannoconus minutus Bronnimann; 19: Nannoconus quadriangulus apertus Deflandre and Deflandre; 20:
Nannoconus broennimannii Trejo; 21: Nannoconus carniolensis latus Deres and Achersiteguy; 22: Nannoconus
steinmannii Kamptner; 23: Nannoconus steinmannii minor Deres and Acheriteguy; 24: Nannoconus dolomiticus Cita
and Pasquare; 25: Nannoconus boletus Deflandre and Deflandre; 26: Nannoconus calpidomorphus Deflandre and
Deflandre; 27: Nannoconus circularis Deres and Acheriteguy; 28: Nannoconus globulus Bronnimann (1955); 29:
Nannoconus arniolensis carniolensis Deflandre and Deflandre; 30: Nannoconus vocontiensis Deres and Achéritéguy;
31: Nannoconus boletus curtus Deflandre and Deflandre-Rigaud; 32: Nannoconus wassallii Bronnimann; 33:
Nannoconus aquitanicus Deres and Acheriteguy; 34: Nannoconus bucheri Bronnimann; 35: Nannoconus kamptneri
Bronnimann; 36: Nannoconus boneti Trejo; 37: Nannoconus grandis Deres and Acheriteguy; 38: Nannoconus
dauvillieri Deflandre and Deflandre; N. = Nannoconus
10 Geological History 247

Fig. 33 1–35: Representative calcareous nannofossil species of Polycyclolithaceae Forchheimer (1972). 1–10:
Eprolithus floralis (Stradner): 11–14: Eprolithus Stover; 15–16: Eprolithus antiquus Perch-Nielsen; 17–18:
Tegulalithus septentrionalis (Stradner); 19–23: Hayesites albiensis Manivit; 24–25: Uniplanarius sissinghii
(Perch-Nielsen); 26: Uniplanarius trifidus (Stradner); 27–29: Lithastrinus grillii Stradner; 30–31: Lithastrinus moratus
(Stover); 32–33: Lithastrinus septenarius Forchheimer; 34–35: Radiolithus planus Stover
248 10 Calcareous Nannofossils

Fig. 34 1–30: Representative calcareous nannofossil species of Polcyclolithaceae Forchheimer (1972). 1–4: Micula
prinsii Perch-Nielsen; 5–7: Micula decussata Vekshina; 8–13: Micula concava (Stradner); 14–16: Micula murus
(Martini); 17: Micula praemurus (Bukry); 18–19: Micula quadrata Perch-Nielsen; 20–23: Micula staurophora
(Gardet); 24–26: Quadrum gartneri Prins and Perch-Nielsen; 27–28: Uniplanarius gothicus (Deflandre); 29:
Uniplanarius sissinghii (Perch-Nielsen); 30: Uniplanarius trifidus (Stradner)
10 Geological History 249

Fig. 35 1–26: Representative calcareous nannofossil species of Prediscosphaeraceae Rood, Hay and Barnard (1971).
1–5: Prediscosphaera Vekshina; 1: Profile view; 2: Distal extremity of stem; 3: Distal view; 4: Proximal view; 5: Side
view; 7–10: Prediscosphaera columnata (Stover); 11: Prediscosphaera microrhabdulina Perch-Nielsen; 12: Predis-
cosphaera spinosa (Bramlette and Martini); 13: Prediscosphaera arkhangelskyi (Reinhardti); 14: Prediscosphaera
serrata Noël; 15: Prediscosphaera bukryi Perch-Nielsen; 16–17: Prediscosphaera stoveri (Perch-Nielsen); 18:
Prediscosphaera rhombica (Perch-Nielsen); 19: Prediscosphaera mgayae Lees; 20: Prediscosphaera grandis
Perch-Nielsen; 21–22: Prediscosphaera columnata (Stover); 23: Prediscosphaera ponticular; 24–25: Prediscosphaera
cretacea (Arkhangelsky); 26: Prediscosphaera majungae Perch-Nielsen. 27–34: Representative calcareous nannofossil
species of Prinsiaceae Hay and Mohler (1967). 27: Emiliania huxleyi (Lohmann); 28–29: Pseudoemiliania Gartner; 30:
Pseudoemiliania lacunosa Kamptner; 31: Rhabdosphaera Haeckel; 32: Reticulofenestra pseudoumbilicus (Gartner); 33:
Reticulofenestra umbilicus (Levin); 34: Reticulofenestra umbilicus (Levin)
250 10 Calcareous Nannofossils

Fig. 36 1–5: Representative calcareous nannofossil species of Pontosphaeraceae Lemmermann (1908). 1–5:
Scyphosphaera Lohmann; 1: Profile view of base with rods and elliptical pores between; 2: Profile view of buccal
discolith with central stem; 3: Profile view; 4: Coccosphere; 5: Proximal view. 6–21: Representative calcareous
nannofossil species of Parhabdolithaceae Bown 1987. 6: Stoverius achylosus (Stover); 7: Stoverius helotatus; 8–9:
Stoverius achylosus (Stover); 10: Stoverius coronatus Bukry; 11–12: Cylindralithus biarcus Bukry; 13: Mitrolithus
Deflandre; 14–15: Crucirhabdus Rood, Hay and Barnard; 16: Nephrolithus frequens Górka; 17: Grantarhabdus Black;
18: Retecapsa Black; 19–20: Amphizygus Bukry; 21: Cretarhabdus Bramlette and Martini
10 Geological History 251

Fig. 37 Representative calcareous nannofossil species of Rhagodiscaceae Hay (1977). 1–3: Blackites Hay and Towe;
4: Blackites gladius (Locker); 5: Calcicalathina oblongata (Worsley)
252 10 Calcareous Nannofossils

Fig. 38 1–22: Representative calcareous nannofossil species of Stephanolithiaceae Black (1968). 1–7: Stephanolithion
Deflandre; 1: Distal view; 2: profile view; 3: section; 4: specimens with digenetic crystalline growth; 6: oblique
proximal view; 7: distal view. 8–11: Stephanolithion hexum Rood and Barnard; 12–15: Stephanolithion speciosum
Deflandre; 16: Stephanolithion speciosum var. speciosum Deflandre; 17 Stephanolithion speciosum var. octum (Rood
and Barnard); 18: Stephanolithion bigoti Deflandre; 19: Stephanolithion bigotii var. bigotii Deflandre; 20:
Stephanolithion bigotii var. maximum (Deflandre); 21: Stephanolithion bigotii Deflandre; 22: Stephanolithion
carinatum Medd
10 Geological History 253

Fig. 39 1–36: Representative calcareous nannofossil species of Stephanolithiaceae Black (1968). 1–2: Truncatosca-
phus hexaporus Moshkovitz and Ehrlich; 3–4: Truncatoscaphus octoporus Moshkovitz and Ehrlich; 5–6:
Truncatoscaphus pauciramous (Black); 7: Truncatoscaphus intermedius Perch-Nielsen; 8–11: Truncatoscaphus
delftensis (Stradner and Adamiker); 12: Stradnerlithus asymmetricus Rood, Hay and Barnard; 13–15: Stradnerlithus
tortuosus Noël; 16: Stradnerlithus geometricus (Górka); 17: Stradnerlithus langii (Rood and Barnard); 18–20:
Stradnerlithus comptus Black; 21: Stradnerlithus humilis Goy, Noël, and Busson; 22: Stradnerlithus callomonii Rood,
Hay and Barnard; 23: Stradnerlithus fragilis (Rood and Barnard); 24: Stradnerlithus escovillensis (Rood and Barnard);
25: Thurmannolithion Grün and Zweili; 26: Thurmannolithion clatratum Grün; 27: Stephanolithion Deflandre; 28:
Truncatascaphus Rood, Hay and Barnard; 29: Corollithion Stradner; 30: Cylindralithus Bramlette and Martini; 31:
Rotelapillus Noël; 32–33: Rotelapillus crenulatus (Stover); 34: Rotelapillus munitus (Perch-Nielsen); 35: Rotelapillus
radians Noël; 36: Rotelapillus crenulatus (Stover)
254 10 Calcareous Nannofossils

Fig. 40 1–5: Representative calcareous nannofossil species of Schizophaerellaceae Deflandre (1959). 1–3:
Schizophaerella Deflandre; 1: section view; 2: lateral view; 3: surface view (juxtaposition of crystals). 4:
Schizosphaerella punctulata Deflandre (type figure); 5: Schizosphaerella punctulata Deflandre
10 Geological History 255

Fig. 41 1–35: Representative calcareous nannofossil species of Sphenolithaceae Deflandre (1952). 1: Sphenolithus
heteromorphus Deflandre; 2: Sphenolithus pseudoradians Bramlette and Wilcoxon; 3: Sphenolithus belemnos
Bramlette and Wilcoxon; 4: Sphenolithus paratintinnabulum Bergen and de Kaenel; 5: Sphenolithus distentus (Martini);
6–7: Sphenolithus abies Deflandre; 8: Sphenolithus dissimilis Bukry and Percival; 9: Sphenolithus orphanknollensis
Perch-Nielsen; 10: Sphenolithus editus Perch-Nielsen; 11: Sphenolithus radians Delfandre; 12: Sphenolithus
pseudoradians Bramlette and Wilcoxon; 13: Sphenolithus predistentus Bramlette and Wilcoxon; 14: Sphenolithus
distentus (Martini); 15: Sphenolithus ciperoensis Bramlette and Wilcoxon; 16: Sphenolithus spiniger Bukry; 17:
Sphenolithus conicus Bukry; 18: Sphenolithus anarrhopus Bukry and Bramlette; 19: Sphenolithus conspicuous Martini;
20: Sphenolithus belemnos Bramlette and Wilcoxon; 21: Sphenolithus heteromorphus Deflandre; 22: Sphenolithus
delphix Bukry; 23: Sphenolithus calyculus Bukry; 24: Sphenolithus tribulosus Roth; 25: Sphenolithus abies Deflandre;
26: Sphenolithus primus Perch-Nielsen; 27: Sphenolithus paratintinnabulum Bergen and de Kaenel; 28: Sphenolithus
neoabies Bukry and Bramlette; 29: Sphenolithus moriformis (Bronnimann and Stradner); 30: Sphenolithus grandis Haq
and Berggren; 31: Sphenolithus furcatolithoides Locker; 32: Sphenolithus capricornutus Bukry and Percival; 33:
Sphenolithus celsus Haq; 34: Sphenolithus obtusus Bukry; 35: Sphenolithus intercalaris Martini
256 10 Calcareous Nannofossils

Fig. 42 1–10: Representative calcareous nannofossil species of Thoracosphaeraceae Schiller (1930). 1–3: Thora-
cosphaera Kamptner; 1: section view; 2: lateral view; 3: surface view (joined crystals). 4: Triquetrorhabdulus Martini;
5: T Triquetrorhabdulus carinatus Martini; 6–7: Isthmolithus Deflandre; 6: frontal view; 7: lateral view. 8: Isthmolithus
recurvus Deflandre. 9–10: Reinhardtites levis Prins and Sissingh. 11–13: Representative calcareous nannofossil species
of Zygodiscaceae Hay and Mohler (1967). 11: Zeugrhabdotus embergeri (Noël); 12: Tranolithus orionatus (Reinhardt);
13: Zygodiscus Bramlette and Sullivan
10 Geological History 257
 258 10 Calcareous Nannofossils

b Fig. 43 1–25: Representative calcareous nannofossil species of Watznaueriaceae (Rood, Hay and Barnard, 1971). 1–2:
Ansulasphaera helvetica Grün and Zweili; 3: Lotharingius sigillatus (Stradner); 4: Lotharingius barozii Noël; 5:
Lotharingius primitivus Rood, Hay and Barnard; 6: Lotharingius crucicentralis (Medd); 7: Lotharingius hauffii Grün; 8:
Cyclagelosphaera reinhardtii (Perch-Nielsen); 9: Cyclagelosphaera margerelii Noël; 10: Cyclagelosphaera rota-
clypeata Bukry; 11. Watznaueria fossacincta Black; 12: Watznaueria barnesae (Black); 13: Watznaueria biporta
Bukry; 14–16: Watznaueria britannica (Stradner); 17–18: Watznaueria ovata Bukry; 19: Watznaueria reinhardtii
Rood, Hay and Barnard; 20: Watznaueria fasciata Wind and Cepek; 21–23: Watznaueria fossacincta Black; 24:
Watznaueria britannica (Stradner); 25: Ellipsagelosphaera plena Grün and Zweili

References Tertiary in California. Micropaleontology, 7, 129–

Black, M. (1968). Taxonomic problems in the study of
coccoliths. Palaeontology, 11, 793–813.
Black, M. (1971). Coccoliths of the Speeton Clay and
Sutterby Marl. Proceeding of the Yorkshire Geolog-
ical Society, 38, 381–424.
Black, M. (1972). British Lower Cretaceous Coccoliths.
I – Gault Clay (Part I). Palaeontological Society of
London (Monograph), 126, 1–48.
Bolli, H. M., Saunders, J. B., & Perch-Nielsen, K. (Eds.).
(1989). Plankton Stratigraphy (p. 1032). Cambridge
University Press.
Boudreaux, J. E., & Hay, W. W. (1969). Calcareous
nannoplankton and biostratigraphy of the Late
Pliocene-Pleistocene-Recent sediments in the Sub-
marex Cores. Revista Espanola de Micropaleontolo-
gia, 1, 249–292.
Bown, P. R. (1987). Taxonomy, evolution, and biostratig-
raphy of Late Triassic-Early Jurassic calcareous nan-
nofossils. Special Papers in Palaeontology, 38, 1–118.
Bown, P. R. (1993). New holococcoJiths from the
Toarcian-Aalenian (Jurassic) of northern Germany.
Senckenbergiana Lethaea, 73, 407–419.
Bown, P. R. (1998). Calcareous Nannofossil Biostratig-
raphy (p.315). British Micropalaeontological Society
Publication Series: Chapman & Hall.
Bown, P. R., Lees, J. A., & Young, J. R. (2004).
Calcareous nannoplankton evolution and diversity
through time. In H. R. Thierstein & J. R. Young
(Eds.), Coccolithophore; from molecular Progress to
Global Impact, 481–508. Berlin: Springer-Verlag.
Bown, P. R., & Young, J. R. (1997). Mesozoic calcareous
nannoplankton classification. Journal of Nannoplank-
ton Research, 19, 21–36.
Bown, P. R., Young, J. R., & Lees, J. A. (2017). On the
cretaceous origin of the order syracosphaerales and the
genus syracosphaera. Journal of Micropalaeontology,
36, 153–165.
Braarud, T., Deflandre, G., Halldal, P., & Kamptner, E.
(1955). Terminology, nomenclature, and systematics
of the Coccolithophoridae. Micropaleontology, 1(2),
157–159.
Bramlette, M. N., & Sullivan, F. R. (1961) Coccol-
ithophorids and related Nannoplankton of the early
188.
Brasier, M. (1980). Microfossils. George Alien & Unwin
(p. 193).
Bronnimann, P. (1955). Microfossils incertae sedis from
the Upper Jurassic and Lower Cretaceous of Cuba.
Micropaleontology, 1, 28–51.
Bukry, D. (1969). Upper Cretaceous coccoliths from
Texas and Europe. The University of Kansas Paleon-
tological Contributions, Article 51 (Protista 2): 1–79.
Deflandre, G. (1947). Braarudosphaera nov. gen., type
d'une famille nouvelle de Coccolithophoridés actuels á
éleéments composites. Comptes Rendus (Hebdo-
madaires des Séances) de l’ Académie des Sciences,
225, 439–441.
Deflandre, G. (1952). Cia sse des Coccolithophoridés
(Coccolithophoridae Lohmann, 1902). Traité de
zoologie, 1, 439–470.
Deflandre, G. (1959). Sur les nannofossiles calcaires et
leur systematique. Revue de Micropaléontologie, 2,
127–152.
Deflandre, G. (1963). Sur les Microrhabdulidés, famille
nouvelle de nannofossiles calcaires. Comptes Rendus
(Hehdomadaires des Sciences) de l’ Académie des
Sciences, 256, 3484–3487.
Didymus, J. M., & Young, J. R. (1994). A net for the
construction of model calcite rhombohedra. Journal of
Nannoplankton Research, 16(2), 79.
Falkowski, P. G., Schofield, O., Katz, M. E., Van de
Schootenbruggem, B., & Knoll, A. H. (2004b). Why
is land green and the ocean red? In H. R. Thierstein &
J. R. Young (Eds.), Coccolithophores: from molecular
processes to global impact (pp. 429–453). Heidelberg:
Springer-Verlag.
Farhan, A. J., Burnett, J. A., Bown, P. R., & Lord, A. R.
(1994). Holococcoliths from the Upper Cretaceous of
Alabama and Mississippi (U.S.A.). Cahiers de
Micropalionlologie, 9, 57–72.
Forchheimer, S. (1972). Scanning electron microscope
studies of Cretaceous coccoliths from the Köpingsberg
Borehole No.1, SE Sweden. Sveriges Geologiska
Undersokning, Serie., C, #668, 65, 1–141.
Gardin, S., Krystyn, L., Richoz, S., Bartolini, A., &
Galbrun, B. (2012). Where and when the earliest
coccolithophores? Lethaia. https://doi.org/10.1111/j.
1502-3931.2012.00311.x.
References
Gradstein, F. M., Ogg, J. G., Schmitz, M. D., & Ogg, G.
M. (2012). The Geologic Time Scale, 2, 801–853.
Halldal, P., & Markali, J., (1955). Electron microscope
studies on coccolithophorids from the Norwegian Sea,
the Gulf Stream and the Mediterranean. Avhandlinger
utgitt av det Norske Videnskapsakademi i Oslo. Mat.
Naturvid. Klasse, 1, 1–30.
Hay, W. W. (1977). Calcareous nannofossils. In: A. T. S.
Ramsay (Ed.), Oceanic Micropalaeontology
(pp. 1055–1200). London: Academic Press.
Hay, W. W., & Mohler, H. P. (1967) Calcareous nanno-
plankton from early Tertiary rocks at Pont Labau,
France, and Paleocene Eocene correlations. Journal of
Palaeotology, 41, 1505–1541.
Jordan, R. W., Kleijne, A., Heimdal, B. R., & Green,
J. C., (1995). A glossary of the extant Haptophyta of
the world. Journal of the Marine Biological Associ-
ation of the United Kingdom, 75, 769–814.
Kamptner, E. (1927). Beitrag zur Kenntnis adriatischer
Coccolithophoriden. Archiv für Protistenkunde, 58,
173–184.
Kamptner, E. (1954). Untersuchungen uber den Feinbau
der Coccolithen. Archiv fur Protistenkunde, 100,
1–90.
Klaveness, D. (1973). The microanatomy of Calyptro-
sphaera sphaernidea with some supplementary obser-
vations on the motile stage of Coccolithus pelagicus.
Norwegian Journal of Botany, 20, 151–162.
Kleijne, A. (1991). Holococcolithophorids from the
Indian Ocean, Red Sea, Mediterranean Sea and North
Atlantic Ocean. Marine Micropaleontology, 17, 1–76.
Lemmermann, E. (1908). Flagellatae, Chlorophyceae,
Coccosphaerales und Silicoflagellatae in Nordische.,
Plankton. Botanischer Teil. In K. Brandt & C. Apstein
(Eds.), Lipsius and Tischer, Kiel and Leipzig,
pp. 1–40.
Lohman, H., (1909). Die Gehause und Gallertblasen der
Appendicularien und ihre Bedeutung fur die
Erforschung des Lebens in Meer. Verhandlungen
Deutsche Zoologische Geselleschaft, 19, 200–239.
Manton, J., & Leeda!e, G. F. (1969). Observations on the
microanatomy of Coccolithus pelagicus and Cri-
cosphaera carterae with special reference to the origin
and nature of coccoliths and scales. Journal of the
Marine Biological Association of the United Kingdom,
49, 1–16.
Norris, R. E. (1965). Living cells of Ceratolithus cristatus
(Coccolithophorineae). Archiv für Proristenkunde,
108, 19–24.
Norris, R. E. (1985). Indian Ocean nannoplanklon II.
Holococcolithophorids (Calyptrosphaeraceae, Prym-
nesiophyceae) with a review of extant genera. Journal
of Phycology, 21, 619–641.
Okada, H., & McIntyre, A. (1977). Modern coccolil-
hophores of the Pacific and North Atlantic Oceans.
Micropaleontology, 2, 155.
Parke, M., & Adams, I. (1960). The motile (Crystallithus
hyalinus Gaarder and Markali) and non-motile phases
in the life history of Coccolithus pelagicus (Wallich)
259
Schiller. Journal of the Marine Biological Association
of the United Kingdom, 39, 263–274.
Perch-Nielsen, K. (1985a). Mesozoic calcareous nanno-
fossils. In H. M. Bolli, J. B. Saunders, K.
Perch-Nielsen (Eds.), Plankton Stratigraphy
(pp. 329–426). Cambridge: Cambridge University
Press
Perch-Nielsen, K. (1985b). Cenozoic calcareous nanno-
fossils. In H. M. Bolli, J. B. Saunders, K.
Perch-Nielsen (Eds.), Plankton stratigraphy
(pp. 427–554). Cambridge: Cambridge University
Press
Perch-Nielsen, K. (1986). Geologic events and the
distribution of calcareous nannofossils—some specu-
lations. Bull Cent Rech Elf E, 10, 421–432.
Pienaar, R. N. (1994). Ultrastructure and calcification of
coccolithophores. In: A. Winter, & W. G. Siesser
(Eds.), Coccolithophores (pp. 13–39). Cambridge:
Cambridge University Press.
Poche, F. (1913). Das System der Protozoa. Archiv für
Protistenkunde, 30, 125–321.
Reinhardt, P. (1965). Neue Familien für fossile Kalkflag-
ellalen (Coccolithophoriden, Coccolithineen). Monats-
berichte der Deutschen Akademie der Wissenschaften
za Berlin, 7, 30–40.
Risatti, J. B. (1973). Nannoplankton biostratigraphy of the
Upper Bluffport Marl-Lower Prairie Bluff Chalk
interval (upper Cretaceous) in Mississippi. In L.
A. Smith & J. Hardenbol (Eds.), In: Proceedings of
the Symposium on Calcareous Nannofossils. Gulf
Coast Section, SEPM (pp. 8–57).
Rood, A. P., Hay, W. W., & Barnard, T. (1973). Electron
microscope studies of Lower and Middle Jurassic
coccoliths. Eclogae Geologicae Helvetiae, 66(2),
365–382.
Roth, P. H. (1973). Calcareous nannofossils-Leg 17, Deep
Sea Drilling Project. Initial Reports of the DSDP, 17,
695–795.
Roth, P. H., & Thierstein, H. (1972). Calcareous nanno-
plankton: Leg 14 of the Deep Sea Drilling Project.
initial Reports of the DSDP, 14, 421–485.
Rowson, J. D., Leadbeater, B. S. C. & Green, J. C.
(1986). Calcium carbonate deposition in the motile
(Crystallithus) phase of Coccolithus pelagicus (Prym-
nesiophyceae). British Phycological Journal, 21,
359–370.
Schiller, J. (1913). Vorläufige Ergebnisse der Phytoplank-
ton–Untersuchungen auf den Fahrten S.M.S. Najade
in der Adria 1911/12. I. Die Coccolitophoriden.
Sitzungberichte der Kaiserlichen Akademie der Wis-
senschaften. Mathematisch–Naturwissenschaftliche
Klasse, 122(1), 597–617.
Schiller, J. (1930). Coccolithineae, in Kryptogamen-Flora
von Deutschland. Osterreich und der Schweiz. 10.
Band, 2 Aht. In: L. Rabenhorst (Ed.), Akademische
Verlagsgesellschaft (pp. 89–267). Leipzig.
Siesser, W. G. (1993). Calcareous nannoplankton. in:
J. H. Lipps (Ed.), Fossil Prokaryotes and Protists
(pp. 169–201). Blackwell Scientific Publications.
260
Tan, S. H. (1927). Over de samenstelling en het onstaan
van krijt en mergelgesteenten van de Molukken.
Jaarboek van het Mijnwezen in Nederlandsch-lndie,
55, 111–122.
Tappan, H. (1980). Haptophyta, coccolithophores, and
other calcareous nannoplankton. In: H. Tappan (Ed.),
The Paleobiology of plant protists (pp. 678–803).
W. H. Freeman and Company, San Francisco.
Thierstein, H. R. (1973). Lower Cretaceous calcareous
nannoplankton biostratigraphy. Ahlwndlunllen der
Geologischen Bundesanstalt, 29, 1–52.
Wallich, G. C. (1877). Observations on the coccosphere.
Annals and Magazine of Natural History, 19, 342–350.
Young, J. R., & Bown, P. R. (2014). Some emendments
to calcareous nannoplankton taxonomy. Journal of
Nannoplankton Research, 33, 139–46.
Young, J. R., Geisen, M., Cros, L., Kleijne, A., Probert, I.,
& Ostergaard, J. B. (2003). A guide to extant
coccolithophore taxonomy. Journal of Nannoplankton
Research, Special Issue, 1, 1–132.
10 Calcareous Nannofossils
Young, J. R., Bergen, J. A., Bown, P. R., Burnett, J. A.,
Fiorentino, A., Jordan, R. W., et al. (1997). Guidelines
for coccolith and calcareous nannofossil terminology.
Palaeontology, 40, 875–912.
Young, J. R., & Bown, P. R. (1997). Cenozoic calcareous
nannoplankton classification. Journal of Nannoplank-
ton Research, 19, 36–47.
Young, J. R., Bown P. R., Burnett, J. A. (1994). The
haptophytes: palaeontological perspectives.
In J. C. Green, B. S. C. Leadbeater (Eds.), The
Haptophyte Algae (Vol. 51, pp. 379–392). The
Systematics Association.
Young, J. R., Didymus, J. M., Bown, P. R., Prins, B., &
Mann, S. (1992). Crystal assembly and phylogenetic
evolution in heterococcoliths. Nature, 356, 516–518.
Young, J. R., & Westbroek, P. (1991). Genotypic
variation in the coccolithophorid species Emiliania
huxleyi. Marine Micropaleontology, 18, 5–23.
 Calpionellids
1 Introduction
The calpionellids are urn-shaped (Fig. 1) plank-
tic Protozoans that were first described (Calpi-
onella; Fig. 1(1)) from the Late Jurassic
(Tithonian) limestones of Alps (Lorenz 1902).
Calpionellids are typically found in deep-sea
deposits of the Late Jurassic and, less frequently,
in the earliest Cretaceous. They range in age
from the Late Jurassic (Late Tithonian) to Late
Cretaceous (Early Valanginian) (Tappan 1993;
Tappan and Loeblich 1968; Remane 1985) (see
Tables 1 and 2; Fig. 2) and are characterized by
rapid evolution (Figs. 2 and 3) with a vast geo-
graphic distribution throughout the Tethys Ocean
(from Mexico to Tibet; see Remane 1989; Pop
1994; Kowal-Kasprzyk 2018). Hence, they are
considered ideal for biostratigraphic studies and
have been used for fine subdivisions, precise
dating and reliable long-distance correlation of
pelagic carbonate deposits of late Lower Titho-
nian to Valanginian age (see Tables 1 and 2)
(Alleman et al. 1971; Remane et al. 1986; Pop
1994; Reháková and Michalík 1997a; Remane
1998; Kowal-Kasprzyk 2018) and have also been
used to define the Jurassic/Cretaceous (J/K)
boundary (Lukeneder et al. 2010; Michalík and
Reháková 2011; López–Martínez et al. 2015).
© Springer Nature India Private Limited 2020
S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8_11
11
2 General Morphology
The axially symmetrical calpionellids possess a
calcite–hyaline, microgranular or mixed test
called lorica (see Fig. 1(1–12)). The oral opening
(=aperture) is always large and surrounded by a
collar of variable form (see Fig. 1(13 and 14)). In
the thin section, the lorica appears generally
transparent (hyaline) or sometimes
micro-granular and brownish opaque. Various
families have different wall ultrastructure which
is best observed under the scanning electron
microscope (SEM). Family Chitnoidellidae Trejo
has a microgranular wall, consisting of rhombo-
hedral crystals and cemented with organic mat-
ter; family Calpionellidae Bonet has a hyaline
wall (Reháková and Michalík 1993). The lorica
is generally oblong and rarely rounded with two
ends—Aboral and Adoral (Fig. 1(13 and 14)).
The adoral end has a round wide mouth, topped
with a collar (Fig. 1(13 and 14)). The collar may
be straight, curved, to varying degrees (and bent
at a different angle), thin or thickened, or bifur-
cated (Makarieva 1979) (Fig. 1(7–12)). This
collar morphology (Fig. 1(13 and 14)) is most
useful for generic identifications (Remane 1989).
The aboral end may be rounded or sharpened
with or without the caudal horn (also referred to
261
262 11 Calpionellids

Fig. 1 Major calpionellid genera (1–6), lorica (test wall) (7–12), morphology (13–16) and sectional details (modified
after Remane 1962). 1: Calpionella Lorenz; 2: Calpionellites Colom; 3: Calpionellopsis Colom; 4: Crassicollaria
Remane; 5: Tintinnopsella Colom; 6: Lorenziella Knauer and Nagy; 7: Calpionella alpina Lorenz; 8: Calpionellites
darderi (Colom); 9: Remaniella cadischiana (Colom); 10: Tintinnopsella carpathica (Murgeanu and Filipescu); 11:
Lorenziella hungarica Knauer and Nagy; 12–13: Calpionellid morphology; 14–15: Tintinnopsella longa (Colom);
16-18: Calpionellopsis simplex (Colom); 19–25: Tintinnopsella carpathica (Murgeanu and Filipescu). Figure 1((6) and
(17–26)): Modified after Remane (1964); 7–12: Modified after Remane (1985); 13–16: Modified after Remane (1962)
2 General Morphology 263

Table 1 Calpionellid bioevent and integrated zonation of the Tithonian, Barriasian and Valanginian of Bulgaria (after
Lakova et al. 1999). 1: Remane (1971a, b); 2: Alleman et al. (1971); 3: Pop (1971, 1997); 4: Reháková and Michalík
(1997a, b); 5: Grün and Blau (1997); 6: Andreini et al. (2007); 7: Bakalova (1977); Bakalova-Ivanova (1986); 8:
Lakova and Stoyanova (1997); Lakova et al. (1997, 1999); Lakova and Petrova (2013)

as caudal appendage) (Fig. 1(13 and 14),
respectively).
The calpionellid test length varies from 40–
150 lm and is made up of a single calcite layer.
The calpionellids can only be studied under thin
sections. But, there are several problems in
studying calpionellids. These are associated with
(a) mechanical damage, (b) recrystallization, and
(c) differing orientation of test in thin sections.
Mechanical damage impacts test wall (Remane
1964, 1974), with the least durable part (generally
the Lorica), is pressed inwards (Fig. 1(15 and 16)).
In test recrystallization, the loss may occur in
various minor elements of the lorica, due to which
the bowl (inner cavity) may take a very different
contour (Fig. 1(17–19)). The orientation of test in
thin sections greatly influences species identifica-
tion as the correct diagnosis of a calpionellid
species depends on the orientation of the test sec-
tion under observation (Fig. 1(20–26)). Due to the
small size of calpionellids and their random dis-
tribution within the sediment, it is almost
impossible to determine the required cross-sec-
tional plane of the section (Fig. 1(20–26)). Hence,
based on the angle under observation, the calpi-
onellids change their shape and parameters for
different test sections (Fig. 1(20–26)). In the event
that the cross section extends perpendicular to the
major axis of the test, the shape will take the form
of thin rings (see Reháková et al. 2009, pl. 1). The
different sections are changing not only its size and
shape, but also in its outline of the collar and the
caudal horn (compare Fig. 1(21–26)). All this,
significantly affects the description and definition
of the calpionellid, under observation; damaged or
bad section of the tests often lead to erroneous
determination or the erection of a new species. It
must be noted that only the axial section (dashed
line in Fig. 1(20)) gives an exact image of lorica
size and proportions, but in most cases the exact
orientation of a given section cannot be deduced
from its outline (Remane 1989). Hence, caution
must be observed when assigning species to a
particular calpionellid.
264 11 Calpionellids

Table 2 Tithonian–Valanginian calpionellid zonation and FO (first occurrence) events for the Mediterranean Realm

Bulgria after Lakova and Petrova (2013)

3 Geological History and Distribution
Fig. 2 Evolution of the most important species of Jurassic–Cretaceous calpionellids. Spain after Reháková and
Michalík (1997b)
3 Geological History
and Distribution
There are no recent calpionellids, and hence, their
systematic position is uncertain (see Tappan
1993). Together with other simple microfossils,
some organic and some calcareous, calpionellids
have been assigned to the organic-walled Tintin-
nids (Tappan 1993). Additionally, because of their
cup-shaped loricae (Fig. 1), they have commonly
been assumed to be Tintinnids. With this view,
some taxonomists have interspersed calpionellids
into modern families based on the shape of the
lorica (Tappan and Loeblich 1968; Lynn 2008).
Others have separated the calpionellids into their
own Tintinnid subgroup (see Bonet 1956). How-
ever, the calcareous tests of calpionellids readily
distinguish them from Tintinnids. Additionally, in
all cases where mineralized tests occur in recent
Protozoa, they are in direct contact with the living
cell over most of their surface, quite unlike the
organic lorica of Tintinnids. For these reasons,
none of the calcareous forms, including calpi-
onellids, can be assigned to the order Tintinnida
nor to class Ciliatea. There are only a few fossil
265
species with an organic lorica which may well be
fossil Tintinnids (see Rüst 1885; Eicher 1965).
4 Classification
The calpionellids have a controversial taxonomic
position and are classified either as Ciliata–
Tintinnida (Colom 1948; Bonet 1956; Trejo
1975; Borza 1984; Tappan 1993) or as Incertae
sedis (Remane 1969, 1971a, b, 1989; Lipps et al.
2013). In general, the calpionellid classification
(Pop 1994; Blau and Grün 1997; Reháková and
Michalík Reháková and Michalík 1997a or b;
Reháková 2002) is in a state of flux. However,
currently, the calpionellids are classified into
three extinct families of Chitinoidellidae Trejo
1975 (test wall or lorica of calcite with an or-
ganic cement; in transmitted light it appears dark;
Lower–Upper Tithonian boundary), Calpionelli-
dae Bonet 1956 (lorica is microgranular, hyaline,
in transmitted light it appears bright; Late
Tithonian–Valanginian) and Colomiellidae Toso
(lorica is hyaline; Aptian–Albian).
Representative calpionellid species are illus-
trated in Figs. 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, and 12.
266 11 Calpionellids

Fig. 3 Stratigraphic distribution and lineages in the evolution of chitinoidellids (after Reháková 2002); the biozonation
(Western Carpathian; Tethyan) is after Reháková and Michalík (1997b; Spain)
4 Classification 267
 268 11 Calpionellids

b Fig. 4 Representative calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–2: Chitinoidella boneti Doben; 3: Chitinoidella carthagensis Sallouhi, Boughdiri and
Cordey; 4: Chitinoidella elongata Pop; 5: Chitinoidella elongata Pop; 6: Chitinoidella sp. indet.; 7–8: Chitinoidella
hegarati Sallouhi, Boughdiri and Cordey; 9–10: Chitinoidella popi Sallouhi, Boughdiri and Cordey; 11: Chitinoidella
sp. indet.; 12–21: Chitinoidella boneti Doben; 22–25: Chitinoidella elongata Pop; 26: Chitinoidella bermudezi
(Furrazola–Bermudezj); 27: Chitinoidella cristobalensis (Furrazola–Bermudez). See Tables 1 and 2 for biozonation. 1–
11: Boneti subzone, Chitinoidella Zone, modified after Kowal-Kasprzyk (2018), courtesy and with permission from
Justyna Kowal-Kasprzyk; 13–18: Modified after Platonov et al. (2014); 19–27: Modified Andreini et al. (2007)
 4 Classification 269

b Fig. 5 Representative calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1: Calpionella alpina Lorenz—large form, close to the Crassicollaria/Calpionella boundary;
2: Calpionella alpina Lorenz—medium form, close to the Crassicollaria/Calpionella boundary; 3: Calpionella alpina
Lorenz—large form (C. “grandalpina”), ?intermedia subzone, Crassicollaria Zone; 4: Calpionella sp. (C. “elliptalpina”),
?intermedia subzone, Crassicollaria Zone; 5: Calpionella alpina Lorenz—medium spherical form, Alpina Subzone,
Calpionella Zone; 6: Calpionella alpina Lorenz—small slightly elliptical form, Elliptica Subzone, Calpionella Zone; 7–
8: Calpionella alpina Lorenz; 9-14: Calpionella alpina Lorenz; 15: Calpionella elliptica Cadisch; 16–17: Calpionella
elliptica Cadisch, Elliptica subzone, Calpionella Zone; 18–19: Calpionella minuta Houša, Elliptica Subzone,
Calpionella Zone; See Tables 1 and 2 for biozonation. 1–6 and 16–19: Courtesy and with permission from Justyna
Kowal-Kasprzyk (see also Kowal–Kasprzyk 2018); 9–14: Modified after Brönnimann (1954)
 270 11 Calpionellids

b Fig. 6 Representative Calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–2: Crassicollaria brevis Remane; 3–4: Crassicollaria colomi Doben; 5–8: Crassicollaria
intermedia (Durand Delga); 9: Crassicollaria massutiniana (Colom); 10–14: Crassicollaria parvula Remane; 15:
Praetintinnopsella andrusovi Borza. See Tables 1 and 2 for biozonation. 1–7 and 9-11: Courtesy and with permission
from Justyna Kowal-Kasprzyk (see also Kowal-Kasprzyk 2018). 1–4, 9 and 11: ?Intermedia subzone, Crassicollaria
Zone. 5–6: Remanei Subzone, Crassicollaria Zone. 8: close to the Crassicollaria/Calpionella boundary. 1–7 and 9–11:
Courtesy and with permission from Justyna Kowal-Kasprzyk (see also Kowal-Kasprzyk 2018); 8, 12, and 13: Modified
after Remane (1964); 14–15: Modified after Platonov et al. (2014)

Fig. 7 Representative Calpionellid species. 1–3: Remaniella ferasini (Catalano); 4–5: Remaniella catalanoi Pop; 6:
Remaniella borzai Pop; 7–10: Remaniella duranddelgai Pop; 11: Remaniella filipescui Pop; 12–13: Remaniella
cadischiana (Colom); 14–15: Remaniella colomi Pop; 16: Sturiella oblonga Borza. 1–9 and 11–16: Modified after
Andreini et al. (2007). 10: Courtesy and with permission from Justyna Kowal-Kasprzyk (see also Kowal-Kasprzyk
2018)
4 Classification 271

Fig. 8 Representative Calpionellid species. 1: Chitinoidella sp. indet. (=Tintinnopsella carpathica (Murgeanu and
Filipescu)—small form of Kowal-Kasprzyk 2018), ?intermedia Subzone, Crassicollaria Zone; 2: Dobeniella colomi
(Borza) (=Tintinnopsella remanei Borza of Kowal-Kasprzyk 2018), Remanei Subzone, Crassicollaria Zone. 3–4:
Daciella danubica Pop (3: Tintinnopsella carpathica (Murgeanu and Filipescu)—large form, Elliptica Subzone,
Calpionella Zone; 4: Tintinnopsella carpathica (Murgeanu and Filipescu)—large form, Elliptica Subzone, Calpionella
Zone of Kowal-Kasprzyk 2018); see Tables 1 and 2 for biozonation. 1–4: Courtesy and with permission from Justyna
Kowal-Kasprzyk (see also Kowal-Kasprzyk 2018). The biozonation (Western Carpathian; Tethyan) is after Reháková
and Michalík (1997b; Spain)
272 11 Calpionellids

Fig. 9 Representative Calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–12: Tintinnopsella carpathica (Murgeanu and Filipescu); 13–17: Tintinnopsella longa
(Colom); 18: Lorenziella gr. hungarica Knauer and Nagy; 19–20: Lorenziella hungarica Knauer and Nagy; 21:
Lorenzlella ruggierii (Catalano); 22 and 25: Praecalpionellites dadayi (Knauer); 23–24 and 28: Praecalpionellites
murgeanui (Pop); 26: Praecalpionellites siriniaensis Pop; 27: Praecalpionellites filipescui (Pop). See Tables 1 and 2 for
biozonation. 1–10 and 14–17: Modified after Platonov et al. (2014); 19–28: Modified after Andreini et al. (2007); 11–13
and 18: Modified after Remane (1964)
4 Classification 273

Fig. 10 Representative Calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–2: Calpionellites major (Colom); 3–4: Calpionellites coronatus (Trejo); 5–8: Calpionellites
darderi (Colom); 9: Calpionellites caravacensis Allemann; 10: Calpionellopsis oblonga (Cadisch); 11: Calpionellopsis
simplex (Colom). See Tables 1 and 2 for biozonation. 1–6 and 9: Modified after Andreini et al. (2007); 7–8 and 10–12:
Modified after Remane (1964)
274 11 Calpionellids

Fig. 11 Representative Calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–2: Dobeniella bermudezi (Furrazola–Bermudez); 3: Dobeniella cubensis (Furrazola–
Bermudez); 4–5: Borzaiella atava Grün and Blau; 6: Borzaiella slovenica (Borza) (=could well be a chitinoidellid
species; personal communication, Mohamed Benzaggagh); 7: Longicollaria dobeni (Borza); 8: Colomiella mexicana
Bonet; 9–10: Colomiella recta Bonet. See Tables 1 and 2 for biozonation. 1–7: Modified after Andreini et al. (2007); 8–
10: Modified after Remane (1964)
4 Classification 275

Fig. 12 Representative calpionellid species. The biozonation (Western Carpathian; Tethyan) is after Reháková and
Michalík (1997b; Spain). 1–2: Daciella danubica Pop; 3: Dobeniella colomi (Borza); 4: Longicollaria dobeni (Borza);
5: Dobeniella cf. tithonica (Borza); 6: Borziella slovenica (Borza); 7: Longicollaria cf. insueta (Řehánek); 8: Popiella
oblongata Reháková; 9: Praetintinnopsella andrusovi Borza. See Tables 1 and 2 for biozonation. 1–9: Courtesy
Justyna Kowal-Kasprzyk (see also Kowal-Kasprzyk, 2018). 1–6: Dobeni Subzone, Chitinoidella Zone; 7–8: Boneti
Subzone, Chitinoidella Zone; 9: Remanei Subzone, Crassicollaria Zone

References Andreini, G., Caracuel, J. E., & Parisi, G. (2007).

Alleman, F., Catalano, R., Fares, F., Remane, J. (1971).
Standard calpionellid zonation (Upper Tithonian–
Valanginian) of the Western Mediterranean province.
In A. Farinacci (Ed.), Proceedings of the II Planktonic
Conference. Roma 1970, 1337–1340.
Calpionellid biostratigraphy of the Upper Tithonian–
Upper Valanginian interval in Western Sicily (Italy).
Swiss Journal of Geoscience, 100, 179–198.
Bakalova, D. (1977). La succession a calpionelles de la
coupe pres du village de Ginci, Bulgarie du
Nord-Ouest. Comptes rendus de l’Academie bulgare
des Sciences, 30, 423–426.
276
Bakalova-Ivanova, D. (1986). Peculiarities of the calpi-
onella zone in Bulgaria. Acta Geologica Hungarica,
29, 89–92.
Blau, J., & Grün, B. (1997). Late Jurassic/Early Creta-
ceous revised Calpionellid zonal and subzonal divi-
sion and correlation with ammonite and absolute time
scales. Mineralia Slovaca, 29, 297–300.
Bonet, F. (1956). Zonificación microfaunística de las
calizas cretácicas del este de México. Boletín de la
Asociación Mexicana de Geólogos Petroleros, 8, 389–
488.
Borza, K. (1984). The Upper Jurassic—Lower Cretaceous
parabiostratigraphic scale on the basis of Tintinninae,
Cadosinidae, Stomiosphaeridae, Calcisphaerulidae
and other microfossils from the West Carpathians.
Geologický Zborník—Geologica Carpathica, 35, 539–
550.
Brönnimann, P. (1954). On the occurrence of Calpionel-
lids in Cuba. Schweizerische Palaeontologische
Gesellschaft, 1953, 263–268.
Colom, G. (1948). Fossil tintinnids: Loricated infusoria of
the order of the oligotricha. Journal of Paleontology,
22(2), 233–263.
Eicher, D. L. (1965). Cretaceous tintinnids from the
western interior of the United States. Micropaleontol-
ogy, 4, 449–456.
Grün, B., & Blau, J. (1997). New aspects of calpionellid
biochronology: Proposal for a revised calpionellids
zonal and subzonal division. Revue de Paléobiologie,
16, 197–214.
Kowal-Kasprzyk, J. (2018). Calpionellid zones of the
Tithonian–Berriasian exotic limestone clasts from the
Outer Carpathians, Poland. Cretaceous Research, 81,
36–50.
Lakova, I., & Petrova, S. (2013). Towards a standard
Tithonian to Valanginian calpionellid zonation of the
Tethyan Realm. Acta Geologica Polonica, 63, 201–
222.
Lakova, I., & Stoyanova, D. (1997). Calpionellid zonation
of Tithonian-Valanginian carbonate successions in the
West Balkan Mts. (Montana Region). Proceedings of
Conference, 50 years of Geology, Sofia University, 7–
10.
Lakova, I., Stoykova, K., & Ivanova, D. (1997). Tithonian
to Valanginian bioevents and integrated zonation on
calpionellids, calcareous nannofossils and calcareous
dinocycts from the Western Balkanides. Bulgaria,
Mineralia Slovaca, 29, 301–303.
Lakova, I., Stoykova, K., & Ivanova, D. (1999). Calpi-
onellid, nannofossil and calcareous dinocyst bioevents
and integrated biochronolgy of the Tithonian to
Valanginian in the Western Balkanides, Bulgaria.
Geologica Carpathica, 50, 151–168.
Lipps, J. H., Stoeck, T., & Dunthorn, M. (2013). Fossil
Tintinnids. In J. R. Dolan, D. J. Montagnes, S. Agatha,
D. W. Coats, & D. K. Stoecker (Eds.), The biology
and ecology of tintinnid ciliates: Models for marine
plankton (pp. 186–197). Oxford: Wiley-Backwell.
López–Martínez, R., Barragán, R., Reháková, D. (2015).
Calpionellid biostratigraphy across the
11 Calpionellids
Jurassic/Cretaceous boundary in San José de Iturbide,
Nuevo León, northeastern Mexico. Geological Quar-
terly, 59, 581–592.
Lorenz, Th. (1902). Geologische Studien im Grenzgebiet
zwischen helvetischer und ostalpiner Fazies. II Der
südliche Rhätikon, Berichte der Naturforschenden
Gesellschaft zu Freiburg, 12, 35–95.
Lukeneder, A., Halásová, E., Kroh, A., Mayrhofer, S.,
Pruner, P., Reháková, D., et al. (2010). High resolu-
tion stratigraphy of the Jurassic-Cretaceous boundary
interval in the Gresten Klippen belt (Austria). Geo-
logica Carpathica, 61, 365–381.
Lynn, D. H. (2008). The ciliated protozoa: Characteri-
zation, classification, and guide to the literature (3rd
ed.). Dordrecht: Springer.
Makarieva, S. F. (1979). Mesozoic Tintinnids from the
North Caucasus, and the Jurassic-Cretaceous bound-
ary. In: Upper Jurassic and the boundary to the
Cretaceous system. Academy Nauk, USSR, Siberian
branch, Institute for Geology and Geophysics, Novosi-
birsk, 168–171.
Michalík, J., & Reháková, D. (2011). Possible markers of
the Jurassic/Cretaceous boundary in the Mediterranean
Tethys: A review and a state of art. Geoscience
Frontiers, 2, 475–490.
Platonov, E. S., Lakova, I., & Arkadiev, V. V. (2014).
Tithonian—Berriasian tintinnids (Ciliophora) of the
Eastern Crimea. Bulletin of Saint-Petersburg State
University, 7(2), 57–81.
Pop, G. (1994). Calpionellid evolutive events and their
use in biostratigraphy. Romanian Journal of Stratig-
raphy, 76, 7–24.
Pop, G. (1997). Révision systématique des chitinoïdelles
tithoniennes des Carpathes méridionales. Comptes
Rendus de l’Académie des Sciences Paris, 324, 931–
938.
Reháková, D. (2002). Chitinoidella Trejo, 1975 in Middle
Tithonian carbonate pelagic sequences of the West
Carpathian Tethyan area. Geologica Carpathica, 53
(6), 369–380.
Reháková, D., Halásová, E., & Lukeneder, A. (2009). The
Jurassic-Cretaceous boundary in the Gresten Klippen-
belt (Nutzhof, Lower Austria): Implications for Micro-
and Nannofacies analysis. Annalen des Naturhis-
torischen Museums in Wien, 110A, 345–381.
Reháková, D., & Michalík, J. (1993). Observations of
Ultrastructure of the Upper Jurassic and Lower
Cretaceous Calpionellid Tests. Geologica Carpathica,
44(2), 75–79.
Reháková, D., & Michalík, J. (1997a). Calpionellid
associations versus Late Jurassic and Early Cretaceous
sea-level fluctuations. Mineralia Slovaca, 29, 306–
307.
Reháková, D., & Michalík, J. (1997b). Evolution and
distribution of calpionellids—the most characteristic
constituents of Lower Cretaceous Tethyan
microplankton. Cretaceous Research, 18, 493–504.
Remane, J. (1962). Zur Calpionellen-Systematik, Neus
Jahbuch für Geologie und Paläontologie. Monatshefte,
1, 8–24.
References
Remane, J. (1964). Untersuchungen zur Systematik und
stratigraphie der Calpionellen in den
Jura-Kreide-Grenzschichten des Vocontischen Troges.
Palaeontographica, 123, 1–57.
Remane, J. (1969). Nouvelles données sur la position
taxonomique des Calpionellidea Bonet (1956) et leurs
rapports avec les Tintinnina actuels et les autres
groups de “Tintinnoidienns” fossils. In P. Brönnimann,
H. H. Renz (Eds.), Proceedings of the first interna-
tional conference on planktonic microfossils. Geneva
1967. Brill, 778 Leiden, 574–587.
Remane, J. (1971a). Les Calpionelles, Protozoaires
planctoniques des mers mésogéennes de l’époque
secondaire. Annales Guébhard, 47, 369–432.
Remane, J. (1971b). Les Calpionells. Protozoaires planc-
toniques des mers mésogéennes de l’époque sec-
ondaire, Annales Guébhard, 47, 369–432.
Remane, J. (1974). Les Calpionelles, Cours de 3 éme
cycle en Sciences de la Terre, partie II, Paléontologie
(p. 58). Genéve: Univ.
Remane, J. (1985). Calpionellids. In H. M. Bolli,
J. B. Saunders, K. Perch-Nielsen (Eds.), Plankton
stratigraphy (pp. 555–572). Cambridge: Cambridge
University Press.
Remane, J. (1989). Calpionellids. In H. M. Bolli,
J. B. Saunders, L. Perch-Nielsen (Eds.), Plankton
stratigraphy, Planktic Foraminifera, Calcareous
277
Nannofossils and Calpionellids (Vol. 1, 555–572
pp.). Cambridge: Cambrigde University Press.
Remane, J. (1998). Calpionellids: possibilities of bios-
tratigraphic correlations versus palaeobiogeographic
limitations. Bulletin de la Société géologique de
France, 168, 829–839.
Remane, J., Bakalova-Ivanova, D., Borza, K., Knauer, J.,
Nagy, I., Pop, G., et al. (1986). Agreement on the
subdivision of the standard calpionellid zones defined
at the II planktonic conference, Roma 1970. Acta
Geologica Hungarica, 29, 5–14.
Rüst, D. (1885). Beiträge zur Kenntniss der fossilen
Radiolarien aus Gesteinen des Jura. Palaeontograph-
ica, 36, 269–322.
Tappan, H. (1993). In W. Tintinnids, J. H. Lipps (Eds.),
Fossil prokaryotes and protists (285–303 pp.). Bos-
ton: Blackwell Scientific Publications.
Tappan, H., & Loeblich, A. R. (1968). Lorica composi-
tion of modern and fossils Tintinnids (Ciliate Proto-
zoa), systematics, Geologic distribution, and some
new Tertiary taxa. Journal of Paleontology, 42, 1378–
1394.
Trejo, M. (1975). Tintinidos mesozoicos de Mexico
(taxonomía y datos paleobiológicos). Boletín de la
Asociación Mexicana de Geólogos Petroleros, 27,
329–449.
 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
Chitinozoan
1 Chitinozoan Eisenackitina? sp. Cambrian Stage 5 1(2–10)
1 Chitinozoan Spinachitina Schallreuter Upper Silurian–Lower Devonian
1 Chitinozoan Eisenackitina cupellata Wrona Bostovian Lower Devonian 2(5)
1 Chitinozoan Sphaerochitina sphaerocephala Eisenack Upper Silurian–Lower Devonian 2(6)
1 Chitinozoan Urnochitina Paris Late Silurian 3(3)
1 Chitinozoan Conochitina Eisenack Late Ordovician 3(4)
1 Chitinozoan Margachitina banwyensis Mullins Late Ordovician–Early Silurian 4(8)
1 Chitinozoan Margachitina margaritana Eisenack Late Ordovician–Early Silurian 4(9)
1 Chitinozoan Anthochitina superba Eisenack Silurian 5(6–13)
1 Chitinozoan Anthochitina radiata Wrona Late Silurian–Early Devonian 5(14 and
15)
1 Chitinozoan Pseudoclathrochitina Cramer Late Ordovician 9(1)
1 Chitinozoan Armoricochitina Paris Late Ordovician–Early Silurian 9(2)
1 Chitinozoan Pterochitina Eisenack Ordovician 9(3)
1 Chitinozoan Hoegisphaera Staplin Middle Precambrian–Early 9(4)
Mississippian
1 Chitinozoan Bulbochitina Paris Early Devonian 9(5)
1 Chitinozoan Urnochitina Paris Late Silurian 9(6)
1 Chitinozoan Cingulochitina Paris Late Silurian 9(7)
1 Chitinozoan Calpichitina Wilson and Hedlund Late Ordovician 9(8)
1 Chitinozoan Bursachitina Taugourdeau Late Ordovician–Early Silurian 9(9)
1 Chitinozoan Desmochitina Eisenack Late Ordovician 9(10)
1 Chitinozoan Margachitina Eisenack Late Ordovician–Early Silurian 9(11)
1 Chitinozoan Linochitina Eisenack Early Silurian 9(12)
1 Chitinozoan Eisenackitina Jansonius Late Ordovician–Early Silurian 9(13)
1 Chitinozoan Ordochitina Achab Asselin and Soufiane Late Ordovician 9(15)
1 Chitinozoan Orbichitina Achab Asselin and Soufiane Late Ordovician 9(16)
1 Chitinozoan Salopochitina Swire Silurian 9(17)
1 Chitinozoan Conochitina Eisenack Late Ordovician 10(1)
1 Chitinozoan Euconochitina Taugourdeau Early Silurian 10(2)
1 Chitinozoan Pistillachitina Taugourdeau Ordovician 10(3)
1 Chitinozoan Rhabdochitina Eisenack Late Ordovician–Early Silurian 10(4)
1 Chitinozoan Clavachitina Taugourdeau Early Ordovician 10(5)
(continued)

© Springer Nature India Private Limited 2020 279

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8
280 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
1 Chitinozoan Velatachitina Poumot Late Ordovician 10(6)
1 Chitinozoan Siphonochitina Jenkins Late Ordovician 10(7)
1 Chitinozoan Eremochitina Taugourdeau and de Jekhowsky Early Ordovician 10(8)
1 Chitinozoan Acanthochitina Eisenack Ordovician 10(9 and
10)
1 Chitinozoan Hercochitina Jansonius Ordovician 10(11)
1 Chitinozoan Belonechitina Jansonius Late Ordovician 10(12)
1 Chitinozoan Tanuchitina Jansonius Middle Ordovician 10(13)
1 Chitinozoan Hyalochitina Paris Middle Ordovician 10(14)
1 Chitinozoan Laufeldochitina Paris Middle Ordovician 10(15)
1 Chitinozoan Pogonocihitina Early Silurian 10(16)
1 Chitinozoan Lagenochitina Eisenack Late Ordovician 11(1)
1 Chitinozoan Saharochitina Paris et Grahn Ordovician 11(2)
1 Chitinozoan Sphaerochitina Eisenack Early Silurian 11(3)
1 Chitinozoan Urochitina Early Devonian 11(4)
1 Chitinozoan Pellichitina Achab Asselin and Soufiane Late Ordovician 11(5)
1 Chitinozoan Sagenachitina Jenkins Early–Middle Ordovician 11(6)
1 Chitinozoan Cyathochitina Eisenack Late Ordovician–Early Silurian 11(7)
1 Chitinozoan Acanthochitina Eisenack Ordovician 11(8)
1 Chitinozoan Parisochitina Boumendjel Silurian 11(9)
1 Chitinozoan Fungochitina Taugourdeau Late Devonian 11(10)
1 Chitinozoan Angochitina Eisenack Middle to Late Ordovician 11(11)
1 Chitinozoan Ramochitina Sommer et van Boekel Late Ordovician–Early Silurian 11(12)
1 Chitinozoan Muscochitina Early–Middle Ordovician 11(13)
1 Chitinozoan Ancyrochitina Eisenack Late Ordovician–Early Silurian 11(14)
1 Chitinozoan Plectochitina Cramer Late Ordovician–Early Silurian 11(15)
1 Chitinozoan Clathrochitina Late Ordovician–Early Silurian 11(16)
1 Chitinozoan Alpenachitina Dunn and Miller Middle Devonian 11(17)
Acritarch
2 Acritarch Dictyosphaera macroreticulata Precambriam 1(1)
2 Acritarch Shuiyousphaeridium macroreticulatum Late Precambrian 1(2)
2 Acritarch Pterospermopsimorpha insolita Neoproterozoic 1(3)
2 Acritarch Micrhystridium Deflandre Ordovician 3(18)
2 Acritarch Riculasphaera (Strother) Early Devonian 3(19)
2 Acritarch Polyancisrrodoms 3(20)
2 Acritarch Diexallophasis gotlandica (Cramer) Silurian 3(21)
2 Acritarch Eupoikilofusa cantabrica (Cramer) Late Silurian 3(22)
2 Acritarch Veryhachium Deunff Late Cambrian 3(23)
2 Acritarch Retisphaeridium howellii Middle Cambrian 4(1)
2 Acritarch Cristallinum cambriense (Bula and Middle Cambrian 4(2)
Jachowicz)
2 Acritarch Timofeeva lancarae (Cramer and Diez de Cambrian–Devonian 4(3)
Cramer)
2 Acritarch Vulcanisphaera turbata Martin Late Cambrian 4(4)
2 Acritarch Cristallinum randomense Martin Cambrian 4(5)
2 Acritarch Vulcanisphaera africana Deunff Late Cambrian 4(6)
2 Acritarch Veryhachium minutum Downie Ordovician 4(7)
2 Acritarch Cymatiogalea velifera (Downie) Early Ordovician 4(8)
(continued)
Species List 281

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
2 Acritarch Cristallinum dentatum Vozzhennikova Late Cambrian–Early Ordovician 4(9)
2 Acritarch Striatotheca principalis Burmann Early Silurian 4(10)
2 Acritarch Dilatisphaera wimanii (Eisenack) Late Ordovician 4(11)
2 Acritarch Tunisphaeridium eisenackii Loeblich et Late Ordovician 4(12)
Tappan
2 Acritarch Dilatisphaera williereae (Martin) Early Silurian 4(13)
2 Acritarch Psenotopus chondrocheus Martin Silurian 4(14)
2 Acritarch Neoveryhachium carminae (Cramer) Early Silurian 4(15)
2 Acritarch Cymbosphaeridium pilar (Cramer) Late Silurian 4(16)
2 Acritarch Daillydium pentaster Staplin Middle Devonian 4(17)
2 Acritarch Umbellasphaeridium saharicum Jardiné et al. Late Devonian 4(18)
2 Acritarch Ammonidium exoticum Deunff Late Silurian 4(19)
2 Acritarch Winwaloeusia ranulaeforma Martin Late Devonian 4(20)
2 Acritarch Volkovia Downie Early Cambrian 5(3)
2 Acritarch Revinotesta Vanguestaine Middle Cambrian 5(4)
2 Acritarch Cymatiogalea Deunff Early Ordovician 5(5)
2 Acritarch Corollasphaeridium Cambrian–Ordovician 5(6)
2 Acritarch Asketopalla Loeblich and Tappan Ordovician 5(7)
2 Acritarch Polyancistrodorus (Loeblich and Tappan) Middle Ordovician 5(8)
2 Acritarch Veryhachium Deunff Ordovician 5(9)
2 Acritarch Diexallophasis Loeblich Late Ordovician 5(10)
2 Acritarch Visbysphaera Lister Silurian 5(11)
2 Acritarch Onondagaella Cramer Late Silurian–Early Devonian 5(12)
2 Acritarch Pterospermopsimorpha Timofeev Tonian Neoproterozoic 8(1)
2 Acritarch Vandalosphaeridium Samuelson Early Neoproterozoic 8(2)
2 Acritarch Octoedryxium Rudavskaja Middle to Late Neoproterozoic 8(3)
2 Acritarch Podolina Timofeev Late Precambrian 8(4)
2 Acritarch Micrhystridium Deflandre Late Permian 8(5)
2 Acritarch Comasphaeridium Staplin Early Cambrian 8(6)
2 Acritarch Comasphaeridium velvetum Moczydowska Early Cambrian 8(6)
2 Acritarch Rhopalophora Tapan and Loeblich Early Cambrian 8(7)
2 Acritarch Skiagia Downie Early Cambrian 8(8)
2 Acritarch Asteridium tornatum (Volkova) Early Cambrian 8(9)
2 Acritarch Annulum squamaceum (Volková) Early Cambrian 8(10)
2 Acritarch Comasphaeridium strigosum Jankauskas Early Cambrian 8(11)
2 Acritarch Skiagia orbicularis (Volkova) Early Cambrian 8(13)
2 Acritarch Skiagia ornata (Volkova) Early Cambrian 8(14)
2 Acritarch Skiagia scottica Downie Early Cambrian 8(15)
2 Acritarch Arbusculidium ornatum (= A. ramusculosum) Early Cambrian 9(1)
Combaz
2 Acritarch Arbusculidium destombesii (= A. frondiferum) Late Cambrian 9(2)
Deunff
2 Acritarch Ladogella Golub and Volkova Late Cambrian–Early Ordovician 9(3)
2 Acritarch Actinotodissus achrasi (Martin) Late Cambrian–Early Ordovician 9(4)
2 Acritarch Actinotodissus Loeblich and Tappan Late Cambrian–Early Ordovician 9(5)
2 Acritarch Actinotodissus achrasi (Martin) Late Cambrian–Early Ordovician 9(6)
2 Acritarch Acanthodiacrodium angustum (Downie) Late Cambrian 9(7)
(continued)
282 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
2 Acritarch Vulcanisphaera africana Deunff Late Cambrian 9(8)
2 Acritarch Vulcanisphaera cf. africana Late Cambrian 9(9)
2 Acritarch Vulcanisphaera imparilis Rasul Early Ordovician 9(10)
2 Acritarch Buedingiisphaeridium tremadocum Rasul Early Ordovician 9(11–12)
2 Acritarch Visbysphaera ?occultata Martin Late Devonian 9(13)
2 Acritarch Visbysphaera ?fecunda Vanguesta Late Devonian 9(14)
2 Acritarch Baltisphaeridium crinitum Martin Early Cambrian 9(15)
2 Acritarch Dasydiacrodium glabrum Combaz Precambrian 9(16)
2 Acritarch Dasydiacrodium cf. glabrum Precambrian 9(17)
2 Acritarch Ephelopalla media (Stockmans and Williere) Late Devonian 9(18)
2 Acritarch Villosacapsula ceratioides (Stockmans and Late Devonian 9(19)
Williere)
2 Acritarch Villosacapsula globosa Vanguestaine et al. Late Devonian 9(20)
2 Acritarch Corollasphaeridium wilcoxianum Martin Late Cambrian–Early Ordovician 9(21)
2 Acritarch Cymatiogalea Deunff Late Cambrian–Early Ordovician 9(22–23)
2 Acritarch Cymatiogalea cuvillieri (Deunff) Early–Middle Ordovician 9(24)
2 Acritarch Daillydium pentaster (Staplin) Late Devonian 9(25)
2 Acritarch Multiplicisphaeridium abnormisum Yin Cambrian–Ordovician boundary 9(26)
Pollen and spores
3 Pollen and Dictyophyllidites harrisii Couper Middle–Late Triassic 1(1)
Spores
3 Pollen and Raistrickia Potonié and Kremp Devonian–Permian 13(1)
Spores
3 Pollen and Geminospora Balme Devonian–Carboniferous 13(2)
Spores
3 Pollen and Rhabdosporites Richardson Devonian–Carboniferous 13(3)
Spores
3 Pollen and Leiotriletes (Naumova) Devonian–Miocene 13(4)
Spores
3 Pollen and Illinites (Kosanke) Carbonferous–Permian 13(5)
Spores
3 Pollen and Torispora Balme Carboniferous 13(6 and
Spores 7)
3 Pollen and Schulzospora Kosanke Carboniferous 13(8–11)
Spores
3 Pollen and Alatisporites Ibrahim Carboniferous–Permian 13(12–
Spores 15)
3 Pollen and Spencerisporites = Microsporites Chaloner Carboniferous–Permian 13(16
Spores and 17)
3 Pollen and Triquitrites (Wilson and Coe) Carboniferous–Permian 13(18)
Spores
3 Pollen and Endosporites (Wilson and Coe) Carboniferous–Permian 13(19–
Spores 20)
3 Pollen and Florinites Schopf Carboniferous–Permian 13(21–
Spores 22)
3 Pollen and Reinschopora Carboniferous–Permian 13(23)
Spores
3 Pollen and Schopfipollenites Potonie and Kremp Carboniferous–Permian 13(24–
Spores 25)
(continued)
Species List 283

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
3 Pollen and Lycospora (Schopf Wilson and Bentall) Carboniferous–Jurassic 13(26)
Spores
3 Pollen and Endosporites (Schopf Wilson and Bentall) Carboniferous–Permian 14(1–2)
Spores
3 Pollen and Tuberculatisporites (Ibrahim) Carboniferous–Triassic 14(4)
Spores
3 Pollen and Striatopodocarpites Sorittch and Sedova Permian 14(5)
Spores
3 Pollen and Vesicaspora (Schemel) Permian 14(6–7)
Spores
3 Pollen and Limitisporites (Leschik) Permian 14(8)
Spores
3 Pollen and Taeniaesporites (Leschik) Permian–Lias 14(9)
Spores
3 Pollen and Nuskoisporites Potonié and Klaus Permian–Trlassic 14(10)
Spores
3 Pollen and Inaperturopollenites Pflug and Thomson Permian–Pliocene 14(11)
Spores
3 Pollen and Cycadoptites Permian–Tertiary 14(12)
Spores
3 Pollen and Classopollis Pflug Triassic–Cretaceous 14(13–
Spores 15)
3 Pollen and Ephedripites Bolkhovitina Triassic–Eocene 14(16)
Spores
3 Pollen and Zonalopollenites (Tsuga) Jurassic–Cretaceous 14(17)
Spores
3 Pollen and Parvisaccites Couper Jurassic–Cretaceous 14(18)
Spores
3 Pollen and Appendicisporites Weyland and Krieger Jurassic–Cretaceous 14(19)
Spores
3 Pollen and Atlantopollis Krutzsch Cretaceous 14(20
Spores and 21)
3 Pollen and Trudopollis Pflug Cretaceous–Eocene 14(22)
Spores
3 Pollen and Phyllocladitites Cretaceous–Tertiary 14(23
Spores and 24)
3 Pollen and Gnetaceapollenites Thiergart Triassic–Recent 15(1)
Spores
3 Pollen and Quercus (Oak) Eocene–Recent 15(2 and
Spores 3)
3 Pollen and Fagus Oligocene–Recent 15(4 and
Spores 5)
3 Pollen and Salicornia (Cuvier) Miocene–Recent 15(6)
Spores
3 Pollen and Abeis Recent 15(7–9)
Spores
3 Pollen and Ephedra Linnaeus Recent 15(10)
Spores
3 Pollen and Microstrobus Garden and Johnson Recent 15(12
Spores and 13)
3 Pollen and Corylus (Hazel) Oligocene–Recent 15(14
Spores and 16)
(continued)
284 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
3 Pollen and Ginkgo Linnaeus Recent 15(17)
Spores
3 Pollen and Alnus (Alder) Oligocene–Recent 15(18
Spores and 19)
3 Pollen and Polypodium Linnaeus Recent 15(20)
Spores
3 Pollen and Lycopodium Linnaeus Recent 15(21–
Spores 23)
Dinoflagellates
4 Dinoflagellates Peridinium leonis (Pavillard) Recent 2(1–5)
4 Dinoflagellates Wetzeliella reticulata Early Eocene 2(6 and
7)
4 Dinoflagellates Gonyaulax spinifera (Claparède and Recent 3(1–4)
Lachmann)
4 Dinoflagellates Ceratium Schrank Recent 3(5–8)
4 Dinoflagellates Hystrichosphaeridium Deflandre Jurassic, Early Kimmeridgian 4(4)
4 Dinoflagellates Gonyaulacysta Deflandre Jurassic 4(1–3)
4 Dinoflagellates Deflandrea Lentin and Williams Cretaceous–Miocene 4(5 and
6)
4 Dinoflagellates Spinifirites Mantell Cenozoic 4(7)
4 Dinoflagellates Hystrichokolpoma tubiferum (Ehrenberg) Eocene 5(1 and
2)
4 Dinoflagellates Spiniferites ramosus (Ehrenberg) Late Jurassic to Present 5(3)
4 Dinoflagellates Oligosphaeridium complex Davey and Recent 5(4)
Williams
4 Dinoflagellates Hystrichokolpoma eisenacki Williams and Eocene 5(5)
Downie
4 Dinoflagellates Diphyes colligerum (Deflandre and Cookson) Paleocene–Eocene 5(6)
4 Dinoflagellates Hystrichodinium Deflandre Jurassic–Cretaceous 11(1 and
2)
4 Dinoflagellates Cannosphaeropsis Wetzel Jurassic–Tertiary 11(3)
4 Dinoflagellates Chordosphaeridium Eisenack Cretaceous–Tertiary 11(4)
4 Dinoflagellates Achomosphaera Evitt Cretaceous–Tertiary 11(5)
4 Dinoflagellates Litosphaeridium Davey and Williams Cretaceous–Eocene 11(6)
4 Dinoflagellates Operculodinium Wall Tertiary 11(7)
4 Dinoflagellates Dapsilidinium langii (Wall) Early Jurassic 11(8)
4 Dinoflagellates Ellipsodinium rugilosum Sarjeant Late Cretaceous 11(9)
4 Dinoflagellates Dacrodinium simile Deflandre Ypresian; Eocene 11(10)
4 Dinoflagellates Chatangiella tripartita (Cookson and Late Cretaceous 11(11)
Eisenack)
4 Dinoflagellates Valensiella Eisenack Jurassic 12(1)
4 Dinoflagellates Meiourogonyaulax Sarjeant Jurassic 12(2 and
3)
4 Dinoflagellates Scriniodinium Klement Jurassic–Cretaceous 12(4)
4 Dinoflagellates Palaeohystrichophora Deflandre Cretaceous 12(5)
4 Dinoflagellates Gonyaulacysta Deflandre Jurassic–Tertiary 12(6 and
7)
4 Dinoflagellates Deflandrea Eisenack Cretaceous–Tertiary 12(8 and
9)
4 Dinoflagellates Pareodinia Eisenack Jurassic–Recent 12(10)
(continued)
Species List 285

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
4 Dinoflagellates Odontochitina Eisenack Cretaceous 12(11)
4 Dinoflagellates Nannoceratopsis Eisenack Jurassic 12(12
and 13)
4 Dinoflagellates Chiropteridium Gocht Oligocene 12(14)
4 Dinoflagellates Wetzeliella Eisenack Eocene–Oligocene 12(15)
4 Dinoflagellates Endoscrinium galeritum (Deflandre) Early to Middle Oxfordian 13(1)
Jurassic
4 Dinoflagellates Deflandrea denticulata Alberti Paleogene 13(2)
4 Dinoflagellates Diconodinium dispersum (Cookson and Albian Early Cretaceous 13(3)
Eisenack)
4 Dinoflagellates Histiophora ornata Klement Middle–Late Jurassic 13(4)
4 Dinoflagellates Oligosphaeridium prolixispinosum Davey and 13(5)
Williams
4 Dinoflagellates Sirmiodiniopsis orbis Drugg Callovian–Oxfordian Jurassic 13(6)
4 Dinoflagellates Spinidinium echinoideum (Cookson and Middle Jurassic–Early 13(7)
Eisenack) Cretaceous
4 Dinoflagellates Spiniferites pseudofurcatus (Klumpp) Oligocene–Early Miocene 13(8)
4 Dinoflagellates Comparodinium diacrorhaetium Morbey Rhaetian Late Triassic 13(9)
4 Dinoflagellates Polykrikos kofoidii Chatton Late Quaternary to Recent 14(1)
4 Dinoflagellates Polykrikos schwartzii Bütschli Pleistocene to Recent 14(2)
4 Dinoflagellates Polykrikos hartmannii (Matsuoka and Recent 14(3)
Fukuyo)
4 Dinoflagellates Balechina coerulea (Dogiel) Recent 14(4)
4 Dinoflagellates Pomatodinium impatiense Cachon and Recent 14(5)
Cachon–Enjumet
4 Dinoflagellates Gonyaulax fragilis (Schütt) Recent 14(6)
4 Dinoflagellates Goniodoma sphaericum Murray and Whitting Recent 14(7)
4 Dinoflagellates Protoperidinium brevipes (Paulsen) Recent 14(8)
4 Dinoflagellates Protoceratium spinulosum (Murray and Recent 14(9)
Whitting)
4 Dinoflagellates Amylax triacantha (Jörgensen) Recent 14(10)
4 Dinoflagellates Spiraulax kofoidii Graham Recent 14(11)
4 Dinoflagellates Protoperidinium oceanicum (VanHöffen) Recent 14(12)
4 Dinoflagellates Ceratium candelabrum (Ehrenberg) Recent 14(13)
4 Dinoflagellates Heterocapsa triquetra (Ehrenberg) Recent 14(14)
4 Dinoflagellates Crypthecodinium cohnii (Seligo) Recent 14(15)
Conodonts
5 Conodonts Spathognathodus Branson and Mehl Pennsylvanian Late 1(2)
Caboniferous
5 Conodonts Ozarkodina Branson Silurian–Devonian 1(3)
5 Conodonts Palmatolepis Ulrich and Bassler Late Devonian 1(4)
5 Conodonts Gnathodus Pander Mississippian Early 1(8 and
Caboniferous 9)
5 Conodonts Polygnathus Hinde Devonian 1(10 and
11)
5 Conodonts Westergaardodina Müller Cambrian–Ordovician 3(1)
5 Conodonts †Chirognathus Branson and Mehl Ordovician 3(2)
5 Conodonts Drepanodus Pander Ordovician 3(3 and
4)
5 Conodonts Coleodus Branson Ordovician 3(5)
(continued)
286 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
5 Conodonts Pygodus Lamont and Lindström Ordovician 3(6)
5 Conodonts Acanthodus Furnish Ordovician 3(7)
5 Conodonts Ozarkodina Branson and Mehl Silurian–Devonian 3(8)
5 Conodonts Prioniodus Pander Ordovician 3(9 and
10)
5 Conodonts Hibbardella Bassler Ordovician–Triassic 3(11–13)
5 Conodonts Ligonodina Branson and Mehl Ordovician–Triassic 3(14)
5 Conodonts Icriodus Branson and Mehl Devonian–Carboniferous 3(15–17)
5 Conodonts Amorphognathus Branson and Mehl Ordovician–Silurian 3(18–21)
5 Conodonts Polygnathus Hinde Devonian–Carboniferous 3(22–27)
5 Conodonts Cavusgnathus Harris and Hollingsworth Carboniferous 3(28)
5 Conodonts Gnathodus Pander Devonian–Permian 3(29–31)
5 Conodonts Ancyrognathus Branson and Mehl Devonian 3(32 and
33)
5 Conodonts Ancyrodella Ulrich and Bassler Devonian 3(34)
5 Conodonts Palmatolepis Müller Devonian 3(35–37)
5 Conodonts Gondolella Stauffer and Plummer Carboniferous–Permian 3(38–40)
5 Conodonts Protopanderotus Ordovician 2(1–4)
5 Conodonts Belodina Ethington Late Ordovician 2(5)
5 Conodonts Clavohamulus Furnish Ordovician 2(6)
5 Conodonts Cornuodus Fåhraeus Early Ordovician 2(7)
5 Conodonts Dapsilodus Cooper Middle Ordovician 2(8)
5 Conodonts Strachanognathus Dzik Ordovician 2(9)
5 Conodonts Parapanderodus Stouge Early Ordovician 2(10)
5 Conodonts Drepanoistodus Lindström Early Ordovician 2(11)
5 Conodonts Oneotodus Lindström Early Ordovician 2(12)
5 Conodonts Protoconodontus Miller Ordovician 2(13)
5 Conodonts Cordylodus Pander Early Ordovician 2(14)
5 Conodonts Fryxellodontus Miller Early Ordovician 2(15)
Radiolarians
6 Radiolarians Albaillella Deflandre Carboniferous 7(1)
6 Radiolarians Haplentactina Deflandre Middle Ordovician 7(2)
6 Radiolarians Paronaella Pessagno Carboniferous–Cretaceous 7(3 and
4)
6 Radiolarians Alievium Pessagno Triassic 7(5 and
6)
6 Radiolarians Capnuchosphera Low Triassic 7(7)
6 Radiolarians Cenosphera (Molina-Cruz) Eocene–Recent 7(8)
6 Radiolarians Ommatartus Haeckel Miocene–Recent 7(9)
6 Radiolarians Canartus Oligocene–Miocene 7(10)
6 Radiolarians Collosphera Müller Miocene–Recent 7(11)
6 Radiolarians Actinomma Haeckel Miocene–Recent 7(12)
6 Radiolarians Amphirhoplanum Pliocene–Recent 7(13)
6 Radiolarians Saturnalis Haeckel Pleistocene–Recent 7(14)
6 Radiolarians Dorcadospyris Haeckel Eocene–Miocene 7(15–17)
6 Radiolarians Heliosphaera clavata Hinde Devonian 8(1)
6 Radiolarians Theodiscus hastus Carter Whalen and Guex Devonian 8(2)
(continued)
Species List 287

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
6 Radiolarians Cenosphaera affinis Hinde Devonian 8(3)
6 Radiolarians Dictyomitra andersonii (Campbell and Clark) Cretaceous 8(4)
6 Radiolarians Saturnalis lateralis Campbell and Clark Cretaceous 8(5)
6 Radiolarians Hexadoridium magnificum Campbell and Cretaceous 8(6)
Clark
6 Radiolarians Lithomelissa armata Petrushevskaya Cretaceous 8(7)
6 Radiolarians Trochodiscus nicholsoni Rüst Carboniferous 8(8)
6 Radiolarians Porodiscus (Trematodiscus) charlestonensis Eocene 8(9)
(Clark and Campbell)
6 Radiolarians Calocyclas semipolita Clark and Campbell Eocene 8(10)
6 Radiolarians Tripilidium clavipes advena Clark and Eocene 8(11)
Campbell
6 Radiolarians Rhopalodictyum malagaense Clark and Miocene 8(12)
Campbell
6 Radiolarians Stylodictya aculeata Jørgensen Miocene 8(13)
6 Radiolarians Pterocorys (Pterocyrtidium) splendens Miocene 8(14)
Campbell and Clark
6 Radiolarians Mirifusus Rüst Jurassic–Cretaceous 9(1 and
2)
6 Radiolarians Saturniforma Pessagno Jurassic–Cretaceous 9(3)
6 Radiolarians Dictyomitra Zittel Cretaceous 9(4–6)
6 Radiolarians Holocryptocapsa Tan Cretaceous 9(7)
6 Radiolarians Theocampe Haeckel Cretaceous–Recent 9(8–9)
6 Radiolarians Calocyclas Ehrenberg Eocene 9(10)
6 Radiolarians Pterocanium Ehrenberg Eocene–Recent 9(11)
6 Radiolarians Lychnocanoma Haeckel Eocene–Recent 9(17)
6 Radiolarians Peripyramis Haeckel Oligocene–Recent 9(22)
6 Radiolarians Anthocyrtidium Kling Oligocene–Recent 9(12)
6 Radiolarians Phormostichoartus Campbell Oligocene–Recent 9(19)
6 Radiolarians Stichocorys (Campbell and Clark) Miocene–Pliocene 9(20)
6 Radiolarians Cyrtocapsa Haeckel Oligocene–Miocene 9(21)
6 Radiolarians Giraffospyris Haeckel Eocene–Recent 9(13–15)
6 Radiolarians Zygocircus Butschli Eocene–Recent 9(16)
6 Radiolarians Callimitra Haeckel Eocene–Recent 9(18)
6 Radiolarians Botryopyle Haeckel Eocene–Recent 9(23)
6 Radiolarians Corocalyptra Haeckel Pleistocene–Recent 9(24)
6 Radiolarians Anthocyrtis mespihis Ehrenberg Tertiary 10(1)
6 Radiolarians Lychocanium lucerna Ehrenberg Tertiary 10(2)
6 Radiolarians Dictyomitra mespilus Ehrenberg Tertiary 10(3)
6 Radiolarians Eucyrtidium elegans Ehrenberg Tertiary 10(4)
6 Radiolarians Pterocodon compana Ehrenberg Tertiary 10(5)
6 Radiolarians Podocyrtis schomburgi Ehrenberg Tertiary 10(6)
6 Radiolarians Cyrtocalpis amphora Tafel Recent 10(7)
6 Radiolarians Bothryocampe hexathalamia Haeckel Recent 10(8)
6 Radiolarians Petalospyris foreolata Ehrenberg Tertiary 10(9)
Diatoms
7 Diatoms Pinnularia Ehrenberg Recent 1(4 and
5)
(continued)
288 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
7 Diatoms Minidiscus trioculatus (Taylor) Recent 2(1)
7 Diatoms Minidiscus comicus Takano Recent 2(2)
7 Diatoms Minidiscus chilensis Rivera Recent 2(3)
7 Diatoms Planktoniella blanda (Schmidt) Recent 2(4–6)
7 Diatoms Planktoniella sol (Wallich) Recent 2(7)
7 Diatoms Asteromphalus Ehrenberg Recent 2(8 and
9)
7 Diatoms Asterolampra marylandica Ehrenberg Recent 2(10)
7 Diatoms Actinoptychus senarius (Ehrenberg) Recent 2(11)
7 Diatoms Planktoniella muriformis (Loeblich Wight Recent 2(13)
Darley)
7 Diatoms Rhizosolenia formosa Peragallo Recent 2(14 and
15)
7 Diatoms Rhizosolenia robusta Norman Recent 2(16)
7 Diatoms Dactyliosolen phuketensis (Sundström) Recent 2(18)
7 Diatoms Guinardia cylindrus (Cleve) Recent 2(17)
7 Diatoms Chaetoceros Ehrenberg Recent 3(1)
7 Diatoms Bacteriastrum furcatum Shadbolt Recent 3(2–6)
7 Diatoms Nanoneis hasleae Norris Recent 3(7 and
8)
7 Diatoms Bacteriastrum delicatulum Cleve Recent 3(9)
7 Diatoms Rhaphoneis amphiceros (Ehrenberg) Recent 3(10)
7 Diatoms Fragilariopsis oceanica (Cleve) Recent 3(11 and
12)
7 Diatoms Achnanthes taeniata Gru Recent 3(13–15)
7 Diatoms Coscinoconus Ehrenberg Cretaceous–Recent 4(1 and
2)
7 Diatoms Actinoptychus Ehrenberg Cretaceous–Recent 4(3 and
4)
7 Diatoms Melosira Agardh Cretaceous–Recent 4(5–7)
7 Diatoms Stephanopyxis (Ehrenberg) Cretaceous–Recent 4(8 and
9)
7 Diatoms Triceratium Ehrenberg Cretaceous–Recent 4(10 and
11)
7 Diatoms Hemiaulus Ehrenberg Cretaceous–Recent 4(12 and
13)
7 Diatoms Biddulphia Gray Cretaceous–Recent 4(15–17)
7 Diatoms Trinacria Heiberg Cretaceous–Oligocene 4(14)
7 Diatoms Rhizosolenia Ehrenberg Oligocene–Recent 5(1)
7 Diatoms Chaetoceros (Ehrenberg) Miocene–Recent 5(2–4)
7 Diatoms Strangulotrema Eocene–Recent 5(5)
7 Diatoms Asterolampra Ehrenberg Eocene–Recent 5(6)
7 Diatoms Cymbella Agardh Miocene–Recent 5(7)
7 Diatoms Denticula Kützing Miocene–Recent 5(8 and
9)
7 Diatoms Pleurosigma Smith Miocene–Recent 5(10 and
11)
7 Diatoms Gomphonema Ehrenberg Pliocene–Recent 5(12 and
13)
(continued)
Species List 289

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
7 Diatoms Fragilaria Lyngbye Eocene–Recent 6(1 and
2)
7 Diatoms Cocconeis Ehrenberg Oligocene–Recent 6(3 and
4)
7 Diatoms Surirella (Turpin) Oligocene–Recent 6(5 and
6)
7 Diatoms Campylodiscus Ehrenberg Miocene–Recent 6(7 and
8)
7 Diatoms Grammatophora Ehrenberg Miocene–Recent 6(9 and
10)
7 Diatoms Epithemia Kützing Pliocene–Recent 6(11 and
12)
7 Diatoms Amophiprora Ehrenberg Miocene–Recent 6(13–15)
7 Diatoms Eunotia Ehrenberg Miocene–Recent 6(16 and
17)
Ostracods
8 Ostracods Bradoria Matthew Early Cambrian 11(1 and
2)
8 Ostracods Indiana Matthew Early Cambrian 1(3–5)
8 Ostracods ?Mononotella Ulrich and Bassler Early Cambrian 1(6 and
7)
8 Ostracods Beyrichona Matthew Early Cambrian 1(8 and
9)
8 Ostracods Sellula Wiman Early Cambrian 1(10)
8 Ostracods Aluta Matthew Early–Middle Cambrian 1(11 and
12)
8 Ostracods ?Cambria Neckaja and Ivanova Early Cambrian 1(13)
8 Ostracods Walcottella Ulrich and Bassler Middle Cambrian 1(14–16)
8 Ostracods ?Eremos Moberg and Segerberg Early Ordovician 1(17)
8 Ostracods Cytherella posterospinosa Boomer Late Cretaceous–Cainozoic 6
8 Ostracods Hyocypris bradyi Sars Late Pliocene 7(1 and
2)
8 Ostracods Hemicytheria folliculosa (Reuss) Late Miocene 7(3)
8 Ostracods Amplocypris recta (Reuss) Miocene 7(5)
8 Ostracods Ctenobolbina ciliata (Emmons) Ordovician 14(1)
8 Ostracods Raymondatia goniglypta Kay Ordovician 14(2 and
3)
8 Ostracods Isochilina bulbosa Harris Ordovician 14(4)
8 Ostracods Eukloedenella richmondensis Spivey Ordovician 14(5)
8 Ostracods Dicranella bicornis Ulrich Ordovician 14(6 and
7)
8 Ostracods Euprimitia labiosa Ulrich Ordovician 14(8)
8 Ostracods Primitiella constricta Ulrich Ordovician 14(9)
8 Ostracods Primitia tumidula Ulrich Ordovician 14(10)
8 Ostracods Sacellatia arcuamuralis Kay Ordovician 14(11
and 12)
8 Ostracods Milleratia cincinnatiensis Miller Ordovician 14(13)
8 Ostracods Bellornatia tricollis Kay Ordovician 14(14)
8 Ostracods Eurychilina reticulata parvifrons Kay Ordovician 14(15
and 16)
(continued)
290 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
8 Ostracods Leperditella rex Coryell and Schenck Ordovician 14(17)
8 Ostracods Schmidtella crassimarginata Ulrich Ordovician 14(18
and 19)
8 Ostracods Bairdia M’Coy ?Ordovician; Carboniferous– 14(20
Permian and 21)
8 Ostracods Bollia suaequata Ulrich Ordovician 14(22)
8 Ostracods Drepanella Ulrich Ordovician–Silurian 15(2 and
3)
8 Ostracods Eurychilina Ulrich Ordovician–Silurian 15(1)
8 Ostracods Aparchites Jones Ordovician–Devonian 15(4)
8 Ostracods Drepanellina clarki Ulrich and Bassler Silurian 15(5 and
6)
8 Ostracods Zygobolbina conradi Ulrich and Bassler Silurian 15(7)
8 Ostracods Zygobeyrichia regina Ulrich and Bassler Silurian 15(8 and
9)
8 Ostracods Zygobolba decora Billings Silurian 15(10)
8 Ostracods Aparchites obliquatus Ulrich and Bassler Silurian 15(11)
8 Ostracods Plethobolbina ornata Ulrich and Bassler Silurian 15(12)
8 Ostracods Mastigobolbina intermedia Ulrich and Bassler Silurian 15(13
and 14)
8 Ostracods Bonnemaia transita Ulrich and Bassler Silurian 15(15
and 16)
8 Ostracods Zygosella mimica Ulrich and Bassler Silurian 15(17)
8 Ostracods Welleria obliqua Ulrich and Bassler Silurian 15(18
and 19)
8 Ostracods Kloedenella rectangularis Ulrich and Bassler Silurian 15(20)
8 Ostracods Paraechmina postica Ulrich and Bassler Silurian 15(21)
8 Ostracods Beyrichia M’Coy Silurian 15(22–
24)
8 Ostracods Entomozoe Pribyl Silurian–Permian 15(25)
8 Ostracods Condracypris binoda Roth Devonian 16(1 and
2)
8 Ostracods Halliella pulchra Bassler Devonian 16(3)
8 Ostracods Ranapeltis typicalis Bassler Devonian 16(4)
8 Ostracods Tubulibairdia windomensis Swartz and Oriel Devonian 16(5 and
6)
8 Ostracods Aechmina cuspidata Jones and Holl Devonian 16(7)
8 Ostracods Parabolbina parvinoda Swartz and Swain Devonian 16(8)
8 Ostracods Favulella favulosa Swartz and Swain Devonian 16(9 and
10)
8 Ostracods Euglyphella numismoides Swartz and Oriel Devonian 16(11)
8 Ostracods Graphiodactylus catenulatus Van Pelt Devonian 16(12)
8 Ostracods Bicornella tricornis Coryell and Cuskley Devonian 16(13)
8 Ostracods Stibus kothornostibus Swartz and Swain Devonian 16(14)
8 Ostracods Beecherella bloomfieldensis Swartz and Swain Devonian 16(15
and 16)
8 Ostracods Euglyphella Warthin Devonian 16(17)
8 Ostracods Dizygopleura recta Roth Devonian 17(1 and
2)
(continued)
Species List 291

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
8 Ostracods Tetradella cicatricosa Warthin Devonian 17(3 and
4)
8 Ostracods Parahealdia pecorella Coryell and Cuskley Devonian 17(5)
8 Ostracods Thlipsurella putea Coryell and Cuskley Devonian 17(6)
8 Ostracods Hyphasmaphora textiligera Van Pelt Devonian 17(7)
8 Ostracods Thlipsura furcoides Bassler Devonian 17(8)
8 Ostracods Octonaria loculosa Ulrich Devonian 17(9)
8 Ostracods Ctenoloculina acanthophora Swartz and Oriel Devonian 17(10)
8 Ostracods Quasillites lobatus Swartz and Oriel Devonian 17(11)
8 Ostracods Octonariella bifurcata Bassler Devonian 17(12)
8 Ostracods Eustephanus catastephanes Swartz and Swain Devonian 17(13)
8 Ostracods Kirkbyella verticolis Coryell and Cuskley Devonian 17(14)
8 Ostracods Entomis rugatulus Van Pelt Devonian 17(15)
8 Ostracods Cytherellina punctulifera Hall Devonian 17(16)
8 Ostracods Hollina devonica Van Pelt Devonian 17(17)
8 Ostracods Monoceratina Devonian–Recent Devonian–Recent 17(18–
20)
8 Ostracods Quasillites Coryell and Malkin Devonian–Carboniferous 17(21
and 22)
8 Ostracods Leperditia Rouault Silurian–Devonian 17(23)
8 Ostracods Cornigella tuberculopsina Jones and Kirkby Pennsylvanian and Permian 18(1 and
2)
8 Ostracods Paraparchites magnus Kellett Pennsylvanian and Permian 18(3)
8 Ostracods Hollinella digitata Kellett Pennsylvanian and Permian 18(4)
8 Ostracods Sulcella sulcata Coryell and Sample Pennsylvanian and Permian 18(5)
8 Ostracods Amphissites centronotus Urich and Bassler Pennsylvanian and Permian 18(6)
8 Ostracods Ectodemites sullivanensis Payne Pennsylvanian and Permian 18(7 and
8)
8 Ostracods Coryellites lowelli Cooper Pennsylvanian and Permian 18(9–10)
8 Ostracods Fabalicypris wetumkaensis Cooper Pennsylvanian and Permian 18(11
and 12)
8 Ostracods Geisina gallowayi Bradfield Pennsylvanian and Permian 18(13
and 14)
8 Ostracods Bairdia beedei Ulrich and Bassler Pennsylvanian and Permian 19(1)
8 Ostracods Cavellina nebrascensis Geinitz Pennsylvanian and Permian 19(2 and
3)
8 Ostracods Sansabella laevis Warthin Pennsylvanian and Permian 19(3 and
4)
8 Ostracods Hastifaba robusta Cooper Pennsylvanian and Permian 19(6 and
7)
8 Ostracods Moorites minutus Warthin Pennsylvanian and Permian 19(8 and
9)
8 Ostracods Discoidella convexa Scott and Borger Pennsylvanian and Permian 19(10)
8 Ostracods Seminolites truncatus Coryell Pennsylvanian and Permian 19(11)
8 Ostracods Roundyella simiplicissima Knight Pennsylvanian and Permian 19(12
and 13)
8 Ostracods Kellettina montosa Knight Pennsylvanian and Permian 19(14)
8 Ostracods Healdia colonyi Coryell and Booth Pennsylvanian and Permian 19(15
and 16)
8 Ostracods Darwinulina Brady and Robertson Carboniferous–Recent 19(17–
19)
(continued)
292 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
8 Ostracods Glyptopleura Girty Carboniferous–Permian 19(20)
8 Ostracods Beyrichiopsis brynhildae Coryell and Jhonson Mississippian 20(1 and
2)
8 Ostracods Beyrichia contracta Cooper Mississippian 20(3 and
4)
8 Ostracods Acratia mucronata Cooper Mississippian 20(5 and
6)
8 Ostracods Incisurella prima Cooper Mississippian 20(7 and
8)
8 Ostracods Carboprimitia depressa Croneis and Mississippian 20(9 and
Funkhouser 10)
8 Ostracods Microparaparchites spinosus Corneis and Mississippian 20(11
Gale and 12)
8 Ostracods Tetratylus ellipticus Cooper Mississippian 20(13
and 14)
8 Ostracods Perprimitia turrita Croneis and Gutke Mississippian 20(15
and 16)
8 Ostracods Glyptopleura alata Corneis and Funkhouser Mississippian 20(17)
8 Ostracods Sargentina allani Coryell and Jhonson Mississippian 20(18
and 19)
8 Ostracods Jonesina insculpta Croneis and Funkhouser Mississippian 21(1 and
2)
8 Ostracods Bairdiolites procerus Cooper Mississippian 21(3 and
4)
8 Ostracods Paracavellina elliptica Cooper Mississippian 21(5 and
6)
8 Ostracods Kirkbyella quadratus Croneis and Gutke Mississippian 21(7 and
8)
8 Ostracods Verrucosella golcondensis Croneis and Gutke Mississippian 21(9)
8 Ostracods Glyptopleurina vetula Cooper Mississippian 21(10
and 11)
8 Ostracods Venula striata Croneis and Funkhouser Mississippian 21(12
and 13)
8 Ostracods Geffenina marmerae Coryell and Sohn Mississippian 21(14
and 15)
8 Ostracods Balantoides quadrilobatus Morey Mississippian 21(6)
8 Ostracods Deloia sulcata Croneis and Funkhouser Mississippian 21(17
and 18)
Foraminifera
9 Foraminifera Alabamina Toulmin Late Cretaceous (Santonian) to 13(1–3)
Holocene
9 Foraminifera Ammonia Brünnich Early Miocene to Holocene 13(4 and
5)
9 Foraminifera Amphistegina d’Orbigny Eocene to Holocene 13(6)
9 Foraminifera Archaeosepta Wernli Middle Jurassic (Late Bajocian 13(7–10)
to Bathonian)
9 Foraminifera Angulogavelinella Hofker Late Cretaceous (Maastrichtian) 13(11–
13)
9 Foraminifera Ammodicus Reuss Silurian to Holocene 13(14)
9 Foraminifera Biloculinella Wiesner Oligocene to Holocene 13(15)
9 Foraminifera Bigenerina d’Orbigny Eocene to Holocene 13(16–
18)
(continued)
Species List 293

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
9 Foraminifera Bolivina d’Orbigny Late Cretaceous (Maastrichtian) 13(19)
to Holocene
9 Foraminifera Bolivinopsis Yakovlev Late Cretaceous 13(20
and 21)
9 Foraminifera Brizalina Costa Late Cretaceous (Campanian) to 13(22
Holocene and 23)
9 Foraminifera Buliminoides Cushman Oligocene to Holocene 13(24)
9 Foraminifera Cassidulina d’Orbigny Late Eocene to Holocene 13(25–
28)
9 Foraminifera Cassidella Hofker Late Cretaceous to M. Oligocene 13(29)
9 Foraminifera Cerobertina Finlay Early Eocene (Ypresian) to 13(3)
Holocene
9 Foraminifera Cancris de Montfort Eocene to Holocene 14(1)
9 Foraminifera Citharina d’Orbigny Early Jurassic to Paleocene 14(2)
9 Foraminifera Chilostomella Reuss in Cžjžek Late Cretaceous to Holocene 14(3)
9 Foraminifera Cyclammina Brady Paleocene to Holocene 14(4 and
5)
9 Foraminifera Clavulina d’Orbigny Paleocene to Holocene 14(6 and
7)
9 Foraminifera Choffatella Schlumberger Late Jurassic (Oxfordian) to Late 14(8)
Cretaceous (Cenomanian)
9 Foraminifera Cyclogyra Holocene 14(9)
9 Foraminifera Cruciloculina d’Orbigny Pliocene to Holocene 14(10
and 11)
9 Foraminifera Eggerella Cushman Eocene to Holocene 14(12
and 13)
9 Foraminifera Ehrenberginia Reuss Eocene to Holocene 14(14–
16)
9 Foraminifera Fissurina Reuss Cretaceous to Holocene 14(17)
9 Foraminifera Frondicularia Defrance Middle Jurassic to Holocene 14(18)
9 Foraminifera Fursenkonina Loeblich and Tappan Late Cretaceous to Holocene. 14(19–
26)
9 Foraminifera Flabellammina Cushman Cretaceous 14(27)
9 Foraminifera Gyroidella Saidova Holocene 14(28
and 29)
9 Foraminifera Gyroidina d’Orbigny Holocene 14(30
and 31)
9 Foraminifera Geosella Holocene 15(1)
9 Foraminifera Gyroidinoides Brotzen Late Cretaceous (Cenomanian) 15(2)
to Holocene
9 Foraminifera Haplophragmoides Cushman Cretaceous to Holocene 15(3)
9 Foraminifera Haplophragmium Reuss Middle Jurassic to Early 15(4–6)
Cretaceous (Hauterivian)
9 Foraminifera Hopkinsina Howe and Wallace Eocene to Holocene 15(7)
9 Foraminifera Hoeglundina Brotzen Paleocene to Holocene 15(8 and
9)
9 Foraminifera Jadammina Bartenstein and Brand Middle Eocene to Holocene 15(1)
9 Foraminifera Jullienella Schlumberger Holocene 15(11)
9 Foraminifera Kurnubia Henson Jurassic 15(12
and 13)
(continued)
294 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
9 Foraminifera Lagena Popescu Middle Miocene to Holocene 15(14
and 15)
9 Foraminifera Trochulina d'Orbigny Middle Eocene to Holocene 15(16)
9 Foraminifera Lenticulina Lamarck Triassic to Holocene 15(17
and 18)
9 Foraminifera Laticarinina Galloway and Wissler Miocene to Holocene 15(19–
22)
9 Foraminifera Loxostomum Ehrenberg Late Cretaceous (Senonian) to 15(23)
Paleocene
9 Foraminifera Neocorbina Holocene 15(24)
9 Foraminifera Nonionella Voloshinova Miocene to Holocene 15(25
and 26)
9 Foraminifera Pyrgo Defrance Late Eocene (Priabonian) to 15(27
Holocene and 28)
9 Foraminifera Paleopolymorphina Cushman and Ozawa Early Cretaceous (Albian) to 16(1–3)
Late Cretaceous
9 Foraminifera Palaeomiliolina Antonova Middle Jurassic (Bajocian) to 16(4–6)
Late Jurassic (Kimmeridgian)
9 Foraminifera Pealerina Lalicker Middle Jurassic (Bathonian) 16(7–9)
9 Foraminifera Pleurostomella Reuss Early Cretaceous (Aptian) to 16(10–
Holocene 11)
9 Foraminifera Protoglobobulimina Hofker Middle Miocene (Tortonian) to 16(12–
Holocene 14)
9 Foraminifera Polystomammina Yabe and Hanzawa Pliocene 16(15–
17)
9 Foraminifera Pseudotrochammina Frerichs Holocene 16(18–
20)
9 Foraminifera Quinqueloculina d’Orbigny Cretaceous to Holocene 16(21)
9 Foraminifera Reophax de Montfort Middle Ordovician to Holocene 16(22)
9 Foraminifera Rectobolivina Cushman Middle Eocene to Holocene 16(23
and 24)
9 Foraminifera Recurvoides Earland Middle Jurassic (Callovian) to 16(25
Holocene and 26)
9 Foraminifera Siphotextularia Finlay Middle Jurassic (Callovian) to 16(27)
Holocene
9 Foraminifera Spiroplectammina Cushman Carboniferous to Holocene 16(28)
9 Foraminifera Spirillina Ehrenberg Late Triassic (Rhaetian) to 16(3)
Holocene
9 Foraminifera Spirillinoides Rhumbler Holocene 16(31)
9 Foraminifera Syringammina Holocene 16(32)
9 Foraminifera Stetsonia Brotzen Late Cretaceous (Turonian to 16(33)
Maastrichtian)
9 Foraminifera Trochammina Parker and Jones Carboniferous to Holocene 17(1–3)
9 Foraminifera Valvobifarina Hofker Holocene 17(4)
9 Foraminifera Verneuilina d’Orbigny Late Jurassic (Kimmeridgian) to 17(5 and
Late Cretaceous (Maastrichtian) 6)
9 Foraminifera Vulvulina d’Orbigny ?Late Cretaceous (Campanian); 17(7–10)
Paleocene to Holocene
9 Foraminifera Verneuilinoides Loeblich and Tappan Jurassic to Cretaceous 17(11
and 12)
9 Foraminifera Vertebralina d’Orbigny Holocene 17(13)
(continued)
Species List 295

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
9 Foraminifera Uvigerina d’Orbigny Early Eocene (Ypresian) to 17(14)
Holocene
Calcareous nannofossils
10 Calcareous Fasciculithus richardii Perch–Nielsen Ypresian–Thanetian 14(1 and
nannofossils 2)
10 Calcareous Fasciculithus Bramlette and Sullivan Ypresian–Selandian 14(3–5)
nannofossils
10 Calcareous Helicosphaeraceae Black Ypresian–Present 14(6)
nannofossils
10 Calcareous Helicosphaera carteri (Wallich) Oligocene–Present 14(7–9)
nannofossils
10 Calcareous Helicosphaera perch–nielseniae (Haq) Burdigalian–Rupelian 14(10)
nannofossils
10 Calcareous Helicosphaera recta (Haq) Serravallian–Rupelian 14(11)
nannofossils
10 Calcareous Nannoconaceae Deflandre Campanian–Oxfordian 14(12)
nannofossils
10 Calcareous Prediscosphaera Vekshina Maastrichtian–Aptian 14(13–
nannofossils 15)
10 Calcareous Sphenolithus moriformis (Bronnimann and Zanclean–Selandian 15(1)
nannofossils Stradner)
10 Calcareous Sphenolithus Deflandre Piacenzian–Danian 15(2–5)
nannofossils
10 Calcareous Sphenolithus heteromorphus Deflandre Serravallian–Burdigalian 15(6)
nannofossils
10 Calcareous Stephanolithiaceae Deflandre Maastrichtian–Pliensbachian 15(7)
nannofossils
10 Calcareous Triquetrorhabdulaceae Lipps Serravallian–Rupelian 15(8)
nannofossils
10 Calcareous Zygodiscaceae Hay and Mohler Chattian–Danian 15(9)
nannofossils
10 Calcareous Arkhangelskiella cymbiformis Vekshina Maastrichtian–Campanian 18(1–3)
nannofossils
10 Calcareous Arkhangelskiella specillata Vekshina Maastrichtian–Campanian 18(4 and
nannofossils 5)
10 Calcareous Arkhangelskiella cribrata (Gazdzicka) Maastrichtian–Campanian 18(6)
nannofossils
10 Calcareous Broinsonia parca parca (Stradner) Campanian 18(7–9)
nannofossils
10 Calcareous Broinsonia parca cf. constricta Hattner Maastrichtian–Campanian 18(10)
nannofossils
10 Calcareous Broinsonia parca constricta Hattner Maastrichtian–Campanian 18(11)
nannofossils
10 Calcareous Broinsonia parca expansa Wise and Watkins Campanian–Turonian 18(12)
nannofossils
10 Calcareous Broinsonia parca cf. expansa Wise and Campanian–Turonian 18(13)
nannofossils Watkins
10 Calcareous Thiersteinia ecclesiastica Wise and Watkins Coniacian–Turonian 18(14)
nannofossils
10 Calcareous Thiersteinia Wise and Watkins Coniacian–Turonian 18(15
nannofossils and 16)
10 Calcareous Kamptneriaceae Bown and Hampton Maastrichtian–Aptian 18(17
nannofossils and 18)
(continued)
296 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Cribrosphaerella Deflandre Maastrichtian–Aptian 19(1)
nannofossils
10 Calcareous Cribrosphaerella daniae Perch–Nielsen Late Maastrichtian–Campanian 19(2)
nannofossils
10 Calcareous Nephrolithus Górka Maastrichtian–Campanian 19(3)
nannofossils
10 Calcareous Cribrocorona Perch–Nielsen Maastrichtian–Santonian 19(4)
nannofossils
10 Calcareous Ethmorhabdus hauterivianus (Black) Barremian–Berriasian 19(5)
nannofossils
10 Calcareous Axopodorhabdus albianus (Black) Cenomanian–Albian 19(6–9)
nannofossils
10 Calcareous Axopodorhabdus atavus (Grün, Prins and Early Tithonian–Late 19(10)
nannofossils Zweili) Pliensbachian
10 Calcareous Axopodorhabdus cylindratus (Noël) Tithonian–Bajocian 19(11
nannofossils and 12)
10 Calcareous Axopodorhabdus dietzmannii (Reinhardt) Cenomanian–Berriasian 19(13
nannofossils and 14)
10 Calcareous Axopodorhabdus rahla (Noël) Oxfordian–Bathonian 19(15
nannofossils and 16)
10 Calcareous Boletuvelum Wind and Wise Maastrichtian–Albian 19(17)
nannofossils
10 Calcareous Biscutum dubium (Noël) Berriasian–Early Toarcian 20(1–3)
nannofossils
10 Calcareous Discorhabdus exilitus Noël Oxfordian 20(4)
nannofossils
10 Calcareous Discorhabdus patulus (Deflandre) (short Oxfordian 20(5)
nannofossils diverging)
10 Calcareous Podorhabdus grassei Noël Tithonian–Aalenian 20(6–8)
nannofossils
10 Calcareous Discorhabdus corollatus Noël Late Tithonian–Early Callovian 20(9–11)
nannofossils
10 Calcareous Discorhabdus Noël Maastrichtian–Toarcian 20(12)
nannofossils
10 Calcareous Sollasites pristinus Noël Early Toarcian 20(13)
nannofossils
10 Calcareous Sollasites bipolaris Rood, Hay and Barnard Early Oxfordian 20(14)
nannofossils
10 Calcareous Sollasites concentricus Rood, Hay and Early Oxfordian 20(15)
nannofossils Barnard
10 Calcareous Sollasites lowei (Bukry) Campanian–Early Pliensbachian 20(16)
nannofossils
10 Calcareous Sollasites arcuatus Black Valanginian 20(17)
nannofossils
10 Calcareous Sollasites falklandensis Filewicz et al. Middle Albian–Early Albian 20(18)
nannofossils
10 Calcareous Sollasites horticus (Stradner and Adamiker) Maastrichtian–Early Oxfordian 20(19
nannofossils and 20)
10 Calcareous Sollasites thiersteinii Filewicz et al. Aptian 20(21)
nannofossils
10 Calcareous Gaarderella granulifera Black Albian 20(22)
nannofossils
(continued)
Species List 297

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Crucibiscutum pinnatus (Black) Early Barremian–Hauterivian 20(23)
nannofossils
10 Calcareous Crucibiscutum hayi (Black) Cenomanian–Albian 20(24)
nannofossils
10 Calcareous Braarudosphaera Deflandre Extant–Cenomanian 21(1)
nannofossils
10 Calcareous Braarudosphaera batilliformis Troelsen and Aptian 21(2)
nannofossils Quadros
10 Calcareous Bukryaster hayi (Bukry) Campanian 21(3 and
nannofossils 4)
10 Calcareous Braarudosphaera quinquecostata Hill Albian 21(5)
nannofossils
10 Calcareous Braarudosphaera stenorhetha Hill Late Albian 21(6)
nannofossils
10 Calcareous Braarudosphaera primula Black Late Albian 21(7)
nannofossils
10 Calcareous Braarudosphaera africana Stradner Cenomanian–Early Albian 21(8–10)
nannofossils
10 Calcareous Braarudosphaera hockwoldensis Black Cenomanian–Late Aptian 21(11)
nannofossils
10 Calcareous Braarudosphaera bigelowii (Gran and Extant–Cenomanian 21(12)
nannofossils Braarud)
10 Calcareous Braarudosphaera regularis Black Late Albian–Early Aptian 21(13)
nannofossils
10 Calcareous Braarudosphaera turbinea Stradner Danian–Maastrichtian 21(14)
nannofossils
10 Calcareous Micrantholithus Deflandre Burdigalian–Danian 21(15)
nannofossils
10 Calcareous Micrantholithus hoschulzii (Reinhardt) Late Aptian–Berriasian 21(16)
nannofossils
10 Calcareous Micrantholithus obtusus Stradner Aptian–Berriasian 21(17)
nannofossils
10 Calcareous Micrantholithus speetonensis Perch–Nielsen Valanginian 21(18)
nannofossils
10 Calcareous Micrantholithus altenuatus Bramlette and Priabonian–Thanetian 21(19)
nannofossils Sullivan
10 Calcareous Micrantholithus belgicus Hay and Towe Coniiacian 21(20)
nannofossils
10 Calcareous Micrantholithus decoratus Deflandre Coniiacian 21(21)
nannofossils
10 Calcareous Trapezopentus sarmatus Wind and Cepek Late Valanginian 21(22)
nannofossils
10 Calcareous Calciosolenia brasiliensis (Lohmann) Extant–Quaternary 22(1–3)
nannofossils
10 Calcareous Rhombolithion duodecostatum (Goy) Early Toarcian 22(4)
nannofossils
10 Calcareous Rhombolithion horrellii (Rood and Barnard) Late Callovian 22(5)
nannofossils
10 Calcareous Rhombolithion rotatum (Rood Hay and Oxfordian 22(6)
nannofossils Barnard)
10 Calcareous Rhombolithion octocostatum (Rood and Late Callovian 22(7)
nannofossils Barnard)
(continued)
298 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Rhombolithion rhombicum (Stradner and Maastrichtian–Hauterivian 22(8)
nannofossils Adamiker)
10 Calcareous Rhombolithion speetonensis Rood and Late Maastrichtian 22(9)
nannofossils Barnard
10 Calcareous Stradnerlithus bifurcatus Noël Kimmeridgian 22(10)
nannofossils
10 Calcareous Corollithion impontus Griin Bajocian–Pliensbachian 22(11)
nannofossils
10 Calcareous Corollithion kennedyi Crux Cenomanian 22(12)
nannofossils
10 Calcareous Corollithion signum Stradner Late Maastrichtian–Late Albian 22(13)
nannofossils
10 Calcareous Corollithion madagaskarensis Perch–Nielsen Maastrichtian–Cenomanian 22(14)
nannofossils
10 Calcareous Corollithion exiguum Stradner Late Maastrichtian–Turonian 22(15)
nannofossils
10 Calcareous Corollithion completum Perch–Nielsen Maastrichtian 22(16)
nannofossils
10 Calcareous Crepidolithus crecifer Prins–Chiastozygaceae Early Pliensbachian–Sinemurian 22(17)
nannofossils
10 Calcareous Crepidolithus crassus (Deflandre)– Tithonian–Sinemurian 22(18)
nannofossils Chiastozygaceae
10 Calcareous Cylindralithus serratus Bramlette and Martini Maastrichtian–Albian 22(19
nannofossils and 20)
10 Calcareous Cylindralithus oweinae Perch–Nielsen Maastrichtian 22(21)
nannofossils
10 Calcareous Cylindralithus nudus Bukry Maastrichtian–Albian 22(22)
nannofossils
10 Calcareous Cylindralithus duplex Perch–Nielsen Maastrichtian 22(23)
nannofossils
10 Calcareous General morphology 22(24)
nannofossils
10 Calcareous Anfractus Medd Tithonian–Toarcian 22(25)
nannofossils
10 Calcareous Semihololithus Perch-Nielsen Priabonian–Selandian 22(26)
nannofossils
10 Calcareous Petrobrasiella Troelsen and Quadros Maastrichtian–Santonian 22(27)
nannofossils
10 Calcareous Athenagalea Hattner and Wise Late Cretaceous 22(28)
nannofossils
10 Calcareous Octolithus Romein Selandian–Campanian 22(29)
nannofossils
10 Calcareous Metadoga Wind and Cepek Campanian–Valanginian 22(3)
nannofossils
10 Calcareous Acuturris Wind and Wise Maastrichtian–Cenomanian 22(31)
nannofossils
10 Calcareous Lucianorhabdus Deflandre Maastrichtian–Valanginian 22(32)
nannofossils
10 Calcareous Lucianorhabdus Deflandre Maastrichtian–Valanginian 22(33)
nannofossils
10 Calcareous Munarinus Risatti Maastrichtian–Cenomanian 22(34)
nannofossils
(continued)
Species List 299

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Multipartis Risatti Lutetian–Danian 22(35)
nannofossils
10 Calcareous Pharus Wind and Wise Maastrichtian 22(36)
nannofossils
10 Calcareous Calculites Prins and Sissingh Maastrichtian–Hauterivian 22(37)
nannofossils
10 Calcareous Russellia Risatti Maastrichtian 22(38)
nannofossils
10 Calcareous Okkolithus Wind and Wise Maastrichtian–Campanian 22(39)
nannofossils
10 Calcareous Ramsaya Risatti Maastrichtian 22(4)
nannofossils
10 Calcareous Orastrum Wind and Wise Maastrichtian–Aptian 22(41)
nannofossils
10 Calcareous Ottavianus Risatti Maastrichtian–Campanian 42
nannofossils
10 Calcareous Zygrhablithus Deflandre Aquitanian–Thanetian 22(43–
nannofossils 45)
10 Calcareous Lucianorhabdus Deflandre Maastrichtian–Valanginian 22(46
nannofossils and 47)
10 Calcareous Calculites obscurus (Deflandre) Maastrichtian–Turonian 22(48)
nannofossils
10 Calcareous Lucianorhabdus maleformis Reinhardt Maastrichtian–Turonian 22(49
nannofossils and 50)
10 Calcareous Lucianorhabdus cayeuxii Deflandre Maastrichtian–Coniacian 22(51
nannofossils and 52)
10 Calcareous Calculites obscurus (Deflandre) Maastrichtian–Turonian 23(2 and
nannofossils 3)
10 Calcareous Calculites ovalis (Stradner) Maastrichtian–Coniacian 23(4–and
nannofossils 5)
10 Calcareous Orastrum campanensis (Cepek) Campanian–Santonian 23(6)
nannofossils
10 Calcareous Orastrum asarotum Wind and Wise Maastrichtian 23(7)
nannofossils
10 Calcareous Petrobrasiella venata Troelsen and Quadros Campanian–Santonian 23(8)
nannofossils
10 Calcareous Semihololithus priscus Perch–Nielsen Thanetian–Maastrichtian 23(9)
nannofossils
10 Calcareous Acuturris scotus (Risatti) Late Maastrichtian–Cenomanian 23(10)
nannofossils
10 Calcareous Semihololithus bicornis Perch–Nielsen Maastrichtian 23(11)
nannofossils
10 Calcareous Lucianorhabdus inflatus Perch–Nielsen and Campanian 23(12)
nannofossils Feinberg
10 Calcareous Lucianorhabdus windii Hattner and Wise Maastrichtian–Campanian 23(13)
nannofossils
10 Calcareous Lucianorhabdus arborius Wind and Wise Maastrichtian 23(14)
nannofossils
10 Calcareous Lucianorhabdus maleformis Reinhardt Maastrichtian–Turonian 23(15
nannofossils and 16)
10 Calcareous Lucianorhabdus cayeuxii Deflandre Late Maastrichtian–Coniacian 23(19
nannofossils and 20)
(continued)
300 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Lucianorhabdus compactus (Verbeek) Turonian–Albian 23(21
nannofossils and 22)
10 Calcareous Lucianorhabdus quadrifidus Forchheimer Maastrichtian–Turonian 23(23–
nannofossils 25)
10 Calcareous Crucirhabdus Rood, Hay and Barnard Pliensbachian–Norian 24(1 and
nannofossils 2)
10 Calcareous Staurorhabdus elongatum (Medd) 24(3)
nannofossils
10 Calcareous Staurolithites stradneri (Rood Hay and Maastrichtian–Early 24(4)
nannofossils Barnard) Pliensbachian
10 Calcareous Staurolithites acutiferrus (Vekshina) = Late Cretaceous 24(5)
nannofossils Ephippium acutiferrus Vekshina
10 Calcareous Staurolithites Caratini Ypresian–Selandian 24(6–8)
nannofossils
10 Calcareous Turbirhabdus Prins Ypresian–Selandian 24(9)
nannofossils
10 Calcareous Bukrylithus Black Maastrichtian–Berriasian 24(10)
nannofossils
10 Calcareous Vacheruvillius Goy Toarcian 24(11
nannofossils and 12)
10 Calcareous Ahmuellerella Reinhardt Maastrichtian–Cenomanian 24(13)
nannofossils
10 Calcareous Chiastozygus platyrhethus Hill Coniacian–Aptian 24(14)
nannofossils
10 Calcareous Heteromarginatus Bukry Campanian 24(15
nannofossils and 16)
10 Calcareous Diadorhombus Worsley Aptian–Berriasian 24(17)
nannofossils
10 Calcareous Diadorhombus rectus Worsley Aptian–Late Berriasian 24(18)
nannofossils
10 Calcareous Rhombolithion minutum (Rood Hay and Oxfordian 24(19)
nannofossils Barnard)
10 Calcareous Chiastozygus Gartner Maastrichtian–Berriasian 24(20)
nannofossils
10 Calcareous Chiastozygus platyrhethus Hill Coniacian–Aptian 24(21)
nannofossils
10 Calcareous Stradnerlithus primitus (Rood Hay and Aalenian–Pliensbachian 24(22)
nannofossils Barnard)
10 Calcareous Calcidiscus Kamptner Extant–Thanetian 25(1–3)
nannofossils
10 Calcareous Cruciplacolithus tenuis (Stradner) Ypresian–Danian 25(4 and
nannofossils 5)
10 Calcareous Ericsonia formosa (Kamptner) = Coccolithus Rupelian–Ypresian 25(6)
nannofossils formosus (Kamptner)
10 Calcareous Chiasmolithus gigas (Bramlette and Sullivan) Lutetian 25(7)
nannofossils = Coccolithus gigas Bramlette and Sullivan
10 Calcareous Chiasmolithus titus Gartner Rupelian–Lutetian 25(8)
nannofossils
10 Calcareous Chiasmolithus californicus (Bramlette and Ypresian–Selandian 25(9)
nannofossils Sullivan)
10 Calcareous Chiasmolithus consuetus (Bramlette and Bartonian–Selandian 25(10)
nannofossils Sullivan)
(continued)
Species List 301

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Chiasmolithus oamaruensis (Deflandre) Rupelian–Priabonian 25(11–
nannofossils 13)
10 Calcareous Chiasmolithus bidens (Bramlette and Sullivan) Ypresian–Selandian 25(14)
nannofossils
10 Calcareous Chiasmolithus expansus (Bramlette and Lutetian 25(15)
nannofossils Sullivan)
10 Calcareous Chiasmolithus solitus (Bramlette and Sullivan) Bartonian–Thanetian 25(16
nannofossils and 17)
10 Calcareous Ceratolithoides Bramlette and Martini Maastrichtian–Campanian 25(18)
nannofossils
10 Calcareous Ceratolithoides arcuatus Prins and Sissingh Maastrichtian–Campanian 25(19
nannofossils and 20)
10 Calcareous Ceratolithoides aculeus (Stradner) Maastrichtian–Campanian 25(21–
nannofossils 24)
10 Calcareous Ceratolithoides aculeus (Stradner) < 90° Maastrichtian–Campanian 25(25)
nannofossils
10 Calcareous Ceratolithoides aculeus (Stradner) > 90° Maastrichtian–Campanian 25(26)
nannofossils
10 Calcareous Ceratolithoides verbeekii Perch–Nielsen Campanian 25(27
nannofossils and 28)
10 Calcareous Ceratolithoides kamptneri Bramlette and Maastrichtian 25(29)
nannofossils Martini
10 Calcareous Ceratolithina hamata Martini Early Turonian–Albian 25(30)
nannofossils
10 Calcareous Ceratolithus rugosus Bukry and Bramlette = Extant–Zanclean 25(31)
nannofossils Ceratolithus cristatus (Kamptner)
10 Calcareous Ceratolithus Kamptner Extant–Tortonian 25(32
nannofossils and 33)
10 Calcareous Cretarhabdus Bramlette and Martini Maastrichtian–Oxfordian 26(1)
nannofossils
10 Calcareous Miravetesina Grün Maastrichtian–Kimmeridgian 26(2)
nannofossils
10 Calcareous Retecapsa Black Maastrichtian–Toarcian 26(3)
nannofossils
10 Calcareous Retecapsa crenulata (Bramlette and Martini) Maastrichtian–Berriasian 26(4)
nannofossils
10 Calcareous Retecapsa escaigii (Noël) Late Jurassic–Aalenian 26(5 and
nannofossils 6)
10 Calcareous Flabellites Thierstein Coniacian–Barremian 26(7)
nannofossils
10 Calcareous Helenea Worsley Cenomanian–Oxfordian 26(8)
nannofossils
10 Calcareous Grantarhabdus Black Campanian–Berriasian 26(9)
nannofossils
10 Calcareous Cretarhabdus Bramlette and Martini Maastrichtian–Oxfordian 26(1)
nannofossils
10 Calcareous Cruciellipsis Thierstein Hauterivian–Berriasian 26(11)
nannofossils
10 Calcareous Speetonia Black Hauterivian–Berriasian 26(12)
nannofossils
10 Calcareous Cruciellipsis cuvillieri (Manivit) Late Hauterivian–Early 26(13
nannofossils Berriasian and 14)
(continued)
302 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Speetonia colligata Black Hauterivian–Berriasian 26(15
nannofossils and 16)
10 Calcareous Discoaster Tan Gelasian–Thanetian 27(1–6)
nannofossils
10 Calcareous Discoaster pentaradiatus Tan Gelasian–Tortonian 27(7)
nannofossils
10 Calcareous Discoaster surculus Martini and Bramlette Gelasian–Tortonian 27(8–10)
nannofossils
10 Calcareous Discoaster deflandrei Bramlette and Riedel Tortonian–Ypresian 27(11
nannofossils and 12)
10 Calcareous Discoaster barbadiensis Tan Priabonian–Ypresian 27(13)
nannofossils
10 Calcareous Discoaster lodoensis Bramlette and Riedel Lutetian–Ypresian 27(14
nannofossils and 15)
10 Calcareous Discoaster sublodoensis Bramlette and Lutetian–Ypresian 27(16
nannofossils Sullivan and 17)
10 Calcareous Discoaster variabilis Martini and Bramlette Gelasian–Tortonian 27(18
nannofossils and 19)
10 Calcareous Discoaster challengeri Bramlette and Riedel Early Miocene 27(20
nannofossils and 21)
10 Calcareous Discoaster brouweri Tan Gelasian–Tortonian 27(23–
nannofossils 25)
10 Calcareous Discoaster pentaradiatus Tan Gelasian–Tortonian 27(26
nannofossils and 27)
10 Calcareous Discoaster tani Bramlette and Riedel Rupelian–Ypresian 27(28)
nannofossils
10 Calcareous Discoaster nobilis Martini Ypresian–Thanetian 27(29)
nannofossils
10 Calcareous Discoaster mohleri Bukry and Percival Ypresian–Thanetian 27(3)
nannofossils
10 Calcareous Discoaster saipanensis Bramlette and Riedel Priabonian–Ypresian 27(31)
nannofossils
10 Calcareous Discoaster kugleri Martini and Bramlette Tortonian–Serravallian 27(32)
nannofossils
10 Calcareous Discoaster multiradiatus Bramlette and Riedel Ypresian–Thanetian 27(33
nannofossils and 34)
10 Calcareous Discoaster hamatus Martini and Bramlette Tortonian 27(35)
nannofossils
10 Calcareous Discoaster quinqueramus Gartner Messinian–Tortonian 27(36)
nannofossils
10 Calcareous Discoaster asymmetricus Gartner Gelasian–Zanclean 27(37)
nannofossils
10 Calcareous Discoaster druggii Bramlette and Wilcoxon Burdigalian–Aquitanian 27(38)
nannofossils
10 Calcareous Eiffellithus Reinhardt Maastrichtian–Berriasian 28(1–3)
nannofossils
10 Calcareous Eiffellithus collis Hoffmann Late Albian 28(4)
nannofossils
10 Calcareous Eiffellithus eximius (Stover) Campanian–Turonian 28(5)
nannofossils
10 Calcareous Eiffellithus parallelus Perch–Nielsen Maastrichtian–Campanian 28(6)
nannofossils
(continued)
Species List 303

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Eiffellithus turriseiffelii (Deflandre) Maastrichtian–Albian 28(7)
nannofossils
10 Calcareous Eiffellithus gorkae Reinhardt Maastrichtian–Albian 28(8)
nannofossils
10 Calcareous Manivitella inversus Black Maastrichtian–Tithonian 28(9)
nannofossils
10 Calcareous Tegumentum stradneri Thierstein Late Maastrichtian–Valanginian 28(10)
nannofossils
10 Calcareous Fasciculithus Bramlette and Sullivan Ypresian–Selandian 28(11–
nannofossils 13)
10 Calcareous Fasciculithus tympaniformis Hay and Mohler Ypresian–Selandian 28(14)
nannofossils
10 Calcareous Heliolithus Bramlette and Sullivan Thanetian–Selandian 28(15–
nannofossils 17)
10 Calcareous Heliolithus kleinpelli Sullivan Thanetian–Selandian 28(18–
nannofossils 19)
10 Calcareous Heliolithus riedelli Bramlette and Sullivan Thanetian 28(2)
nannofossils
10 Calcareous Helicosphaera Kamptner Extant–Ypresian 29(1 and
nannofossils 2)
10 Calcareous Helicosphaera carteri (Wallich) Extant–Chattian 29(3–6)
nannofossils
10 Calcareous Helicopontosphaera? subantarctica Edwards Early Oligocene 29(7)
nannofossils and Perch–Nielsen
10 Calcareous Helicosphaera pacifica Müller and Messinian–Serravallian 29(8)
nannofossils Brönnimann
10 Calcareous Helicosphaera papillata (Bukry and Lutetian–Ypresian 29(9)
nannofossils Bramlette)
10 Calcareous Helicosphaera ampliaperta Bramlette and Langhian–Burdigalian 29(1)
nannofossils Wilcoxon
10 Calcareous Helicosphaera scissura Müller Serravallian–Aquitanian 29(11)
nannofossils
10 Calcareous Helicopontosphaera minima Martini Early Oligocene 29(12)
nannofossils
10 Calcareous Helicosphaera walbersdorfensis Muller Tortonian–Burdigalian 29(13–
nannofossils 15)
10 Calcareous Helicosphaera gertae Bukry Burdigalian–Aquitanian 29(16)
nannofossils
10 Calcareous Helicosphaera orientalis Black Messinian–Serravallian 29(17)
nannofossils
10 Calcareous Helicosphaera hyalina Gaarder Extant–Late Miocene 29(18)
nannofossils
10 Calcareous Helicosphaera wallichii (Lohmann) Extant–Tortonian 29(19)
nannofossils
10 Calcareous Helicosphaera granulata (Bukry and Percival) Tortonian–Chattian 29(2)
nannofossils
10 Calcareous Helicosphaera kampteri Hay and Mohler Oligocene–Miocene boundary 29(21)
nannofossils
10 Calcareous Helicosphaera inversa (Gartner) Ionian–Calabrian 29(22)
nannofossils
10 Calcareous Helicosphaera obliqua Bramlette and Serravallian–Rupelian 29(23)
nannofossils Wilcoxon
10 Calcareous Helicosphaera orientalis Black Messinian–Serravallian 29(24)
nannofossils
(continued)
304 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Helicosphaera perch–nielseniae (Haq) Serravallian–Rupelian 29(25)
nannofossils
10 Calcareous Helicosphaera truempyi Biolzi and Perch– Burdigalian–Chattian 29(26)
nannofossils Nielsen
10 Calcareous Helicosphaera wilcoxonii (Gartner) Chattian–Bartonian 29(27)
nannofossils
10 Calcareous Helicosphaera seminulum Bramlette and Bartonian–Ypresian 29(28)
nannofossils Sullivan
10 Calcareous Helicosphaera mediterranea Muller Tortonian–Rupelian 29(29)
nannofossils
10 Calcareous Helicosphaera recta (Haq) Burdigalian–Rupelian 29(3)
nannofossils
10 Calcareous Helicosphaera compacta Bramlette and Chattian–Lutetian 29(31)
nannofossils Wilcoxon
10 Calcareous Helicosphaera rhomba (Bukry) Serravallian–Chattian 29(32)
nannofossils
10 Calcareous Helicosphaera intermedia Martini Zanclean–Priabonian 29(33)
nannofossils
10 Calcareous Helicosphaera euphratis Haq Burdigalian–Priabonian 29(34)
nannofossils
10 Calcareous Helicosphaera bramlettei (Müller) Chattian–Ypresian 29(35)
nannofossils
10 Calcareous Helicosphaera reticulata Bramlette and Rupelian–Lutetian 29(36–
nannofossils Wilcoxon 39)
10 Calcareous Helicosphaera lophota (Bramlette and Priabonian–Ypresian 29(4)
nannofossils Sullivan)
10 Calcareous Helicosphaera vedderi Bukry Serravallian–Aquitanian 29(41)
nannofossils
10 Calcareous Helicosphaera sellii (Bukry and Bramlette) Ionian–Messinian 29(42)
nannofossils
10 Calcareous Marthasterites Deflandre Campanian–Turonian 30(1)
nannofossils
10 Calcareous Marthasterites furcatus (Deflandre) Campanian–Turonian 30(2–6)
nannofossils
10 Calcareous Petrarhabdus copulatus (Deflandre) Maastrichtian–Campanian 30(7 and
nannofossils 8)
10 Calcareous Annulithus arkelli Rood, Hay and Barnard Early Hettangian 30(9)
nannofossils
10 Calcareous Tribrachiatus orthostylus Shamrai Ypresian 30(10)
nannofossils
10 Calcareous Tribrachiatus contortus (Stradner) Ypresian 30(11)
nannofossils
10 Calcareous Lithraphidites carniolensis Deflandre Maastrichtian–Early Berriasian 31(1)
nannofossils
10 Calcareous Lithraphidites carniolensis serratus Maastrichtian–Early Berriasian 31(2)
nannofossils Shumenko
10 Calcareous Lithraphidites carniolensis var. carniolensis Maastrichtian–Early Berriasian 31(3)
nannofossils Deflandre
10 Calcareous Lithraphidites grossopectinatus Bukry Maastrichtian 31(4)
nannofossils
10 Calcareous Lithraphidites quadratus Bramlette and Maastrichtian 31(5–7)
nannofossils Martini
10 Calcareous Lithraphidites kennethii Perch–Nielsen Maastrichtian 31(8)
nannofossils
(continued)
Species List 305

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Lithraphidites pseudoquadratus Crux Turonian–Cenomanian 31(9)
nannofossils
10 Calcareous Lithraphidites acutus var. acutus Verbeek and Cenomanian 31(10
nannofossils Manivit and 11)
10 Calcareous Lithraphidites alatus var. alatus Thierstein Cenomanian–Albian 31(12)
nannofossils
10 Calcareous Lithraphidites praequadratus Roth Maastrichtian–Campanian 31(13)
nannofossils
10 Calcareous Lithraphidites bollii (Thierstein) Hauterivian 31(14)
nannofossils
10 Calcareous Pseudolithraphidites quattuorbacillatus Early Tithonian 31(15)
nannofossils Keupp
10 Calcareous Pseudolithraphidites multibacillatus Keupp Early Tithonian 31(16)
nannofossils
10 Calcareous Lithostromation Deflandre Ionian–Ypresian 31(17–
nannofossils 19)
10 Calcareous Pseudomicula quadrata Perch–Nielsen Maastrichtian 31(20)
nannofossils
10 Calcareous Micrantholithus undosus 31(21)
nannofossils
10 Calcareous Microrhabdulus Deflandre 31(22–
nannofossils 24)
10 Calcareous Nannoconus truittii var. rectangularis Deres Campanian–Albian 32(1)
nannofossils and Achéritéguy
10 Calcareous Nannoconus truittii var. frequens Deres and Campanian–Aptian 32(2)
nannofossils Achéritéguy
10 Calcareous Nannoconus truittii var. truittii Brönnimann Campanian–Aptian 32(3)
nannofossils
10 Calcareous Nannoconus fragilis Deres and Acheriteguy Albian 32(4)
nannofossils
10 Calcareous Nannoconus donnatensis Deres and Albian 32(5)
nannofossils Acheriteguy
10 Calcareous Nannoconus cornuta Deres and Acheriteguy Hauterivian–Valanginian 32(6)
nannofossils
10 Calcareous Nannoconus quadratus (Noël) Valanginian–Tithonian 32(7)
nannofossils
10 Calcareous Nannoconus farinacciae Bukry Campanian–Coniacian 32(8)
nannofossils
10 Calcareous Nannoconus regularis Deres and Acheriteguy Campanian–Albian 32(9)
nannofossils
10 Calcareous Nannoconus multicadus Deflandre and Campanian–Albian 32(10)
nannofossils Deflandre–Rigaud
10 Calcareous Nannoconus elongatus var. elongatus Campanian–Barremian 32(11)
nannofossils Brönnimann
10 Calcareous Nannoconus colomi (de Lapparent) Barremian–Tithonian 32(12)
nannofossils
10 Calcareous Nannoconus inconspicuus Deflandre and Late Aptian 32(13)
nannofossils Deflandre–Rigaud
10 Calcareous Nannoconus abundans Stradner and Grün Barremian 32(14)
nannofossils
10 Calcareous Nannoconus borealis Perch–Nielsen Barremian 32(15)
nannofossils
10 Calcareous Nannoconus bermudezi Brönnimann Barremian–Valanginian 32(16)
nannofossils
(continued)
306 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Nannoconus quadriangulus var. Aptian 32(17)
nannofossils quadriangulus Deflandre and Deflandre
10 Calcareous Nannoconus minutus Brönnimann Campanian–Hauterivian 32(18)
nannofossils
10 Calcareous Nannoconus quadriangulus apertus Deflandre Aptian 32(19)
nannofossils and Deflandre
10 Calcareous Nannoconus bronnimanni Trejo Valanginian–Tithonian 32(20)
nannofossils
10 Calcareous Nannoconus carniolensis latus Deres and Holocene 32(21)
nannofossils Achersiteguy
10 Calcareous Nannoconus steinmannii Kamptner Aptian–Early Berriasian 32(22)
nannofossils
10 Calcareous Nannoconus steinmannii minor Deres and Early Berriasian 32(23)
nannofossils Acheriteguy
10 Calcareous Nannoconus dolomiticus Cita and Pasquare Berriasian–Tithonian 32(24)
nannofossils
10 Calcareous Nannoconus boletus Deflandre and Deflandre– Late Aptian 32(25)
nannofossils Rigaud
10 Calcareous Nannoconus calpidomorphus Deflandre and Late Aptian 32(26)
nannofossils Deflandre–Rigaud
10 Calcareous Nannoconus circularis Deres and Achéritéguy Aptian–Late Valanginian 32(27)
nannofossils
10 Calcareous Nannoconus globulus var. globulus Aptian–Berriasian 32(28)
nannofossils Brönnimann
10 Calcareous Nannoconus carniolensis var. carniolensis Late Aptian 32(29)
nannofossils Deflandre and Deflandre
10 Calcareous Nannoconus vocontiensis Deres and Cenomanian–Aptian 32(30)
nannofossils Acheriteguy
10 Calcareous Nannoconus boletus var. curtus Deflandre and Mid-Cretaceous 32(31)
nannofossils Deflandre–Rigaud
10 Calcareous Nannoconus wassalli Brönnimann Aptian–Berriasian 32(32)
nannofossils
10 Calcareous Nannoconus aquitanicus Deres and Early Aptian–Barremian 32(33)
nannofossils Acheriteguy
10 Calcareous Nannoconus bucheri Brönnimann Aptian–Late Valanginian 32(34)
nannofossils
10 Calcareous Nannoconus kamptneri Brönnimann Aptian–Early Berriasian 32(35)
nannofossils
10 Calcareous Nannoconus boneti Trejo Early Aptian–Valanginian 32(36)
nannofossils
10 Calcareous Nannoconus grandis Deres and Achéritéguy Early Aptian–Berriasian 32(37)
nannofossils
10 Calcareous Nannoconus dauvillieri Deflandre and Campanian–Albian 32(38)
nannofossils Deflandre
10 Calcareous Eprolithus floralis (Stradner) Campanian–Aptian 33(1–10)
nannofossils
10 Calcareous Eprolithus Stover Campanian–Hauterivian 33(11–
nannofossils 14)
10 Calcareous Eprolithus antiquus Perch–Nielsen Hauterivian 33(15
nannofossils and 16)
10 Calcareous Tegulalithus septentrionalis (Stradner) Hauterivian 33(17
nannofossils and 18)
(continued)
Species List 307

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Hayesites albiensis Manivit Cenomanian–Albian 33(19–
nannofossils 23)
10 Calcareous Uniplanarius sissinghii (Perch–Nielsen) Early Maastrichtian–Campanian 33(24
nannofossils and 25)
10 Calcareous Uniplanarius trifidus (Stradner) Maastrichtian–Campanian 33(26)
nannofossils
10 Calcareous Lithastrinus grillii Stradner Campanian–Coniacian 33(27–
nannofossils 29)
10 Calcareous Eprolithus moratus (Stover) Santonian–Turonian 33(30
nannofossils and 31)
10 Calcareous Lithastrinus septenarius Forchheimer Santonian–Turonian 33(32
nannofossils and 33)
10 Calcareous Radiolithus planus Stover Early Turonian–Aptian 33(34
nannofossils and 35)
10 Calcareous Micula prinsii Perch–Nielsen Maastrichtian 34(1–4)
nannofossils
10 Calcareous Micula staurophora (Gardet) Maastrichtian–Coniacian 34(5–7)
nannofossils
10 Calcareous Micula concava (Stradner) Maastrichtian–Coniacian 34(8–13)
nannofossils
10 Calcareous Micula murus (Martini) Maastrichtian 34(14–
nannofossils 16)
10 Calcareous Micula praemurus (Bukry) Maastrichtian–Campanian 34(17)
nannofossils
10 Calcareous Micula quadrata (Stradner) Maastrichtian 34(18
nannofossils and 19)
10 Calcareous Micula staurophora (Gardet) Maastrichtian–Coniacian 34(20–
nannofossils 23)
10 Calcareous Quadrum gartneri Prins and Perch-Nielsen Maastrichtian–Turonian 34(24–
nannofossils 26)
10 Calcareous Uniplanarius gothicus (Deflandre) Maastrichtian–Santonian 34(27
nannofossils and 28)
10 Calcareous Uniplanarius sissinghii (Perch-Nielsen) Early Maastrichtian–Campanian 34(29)
nannofossils
10 Calcareous Uniplanarius trifidus (Stradner) Maastrichtian–Campanian 34(29
nannofossils and 30)
10 Calcareous Prediscosphaera Vekshina Maastrichtian–Aptian 35(1–5)
nannofossils
10 Calcareous Prediscosphaera columnata (Stover) Turonian–Albian 35(6–10)
nannofossils
10 Calcareous Prediscosphaera microrhabdulina Perch– Maastrichtian–Santonian 35(11)
nannofossils Nielsen
10 Calcareous Prediscosphaera spinosa (Bramlette and Maastrichtian–Aptian 35(12)
nannofossils Martini)
10 Calcareous Prediscosphaera arkhangelskyi (Reinhardt) Maastrichtian–Santonian 35(13)
nannofossils
10 Calcareous Prediscosphaera serrata Noël Maastrichtian–Santonian 35(14)
nannofossils
10 Calcareous Prediscosphaera bukryi Perch–Nielsen Maastrichtian–Albian 35(15)
nannofossils
10 Calcareous Prediscosphaera stoveri (PerchNielsen) Maastrichtian–Campanian 35(16
nannofossils and 17)
(continued)
308 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Prediscosphaera rhombica (Perch–Nielsen) Maastrichtian–Campanian 35(18)
nannofossils
10 Calcareous Prediscosphaera mgayae Lees Early Maastrichtian–Campanian 35(19)
nannofossils
10 Calcareous Prediscosphaera grandis Perch–Nielsen Maastrichtian–Coniacian 35(20)
nannofossils
10 Calcareous Prediscosphaera columnata (Stover) Turonian–Albian 35(21
nannofossils and 22)
10 Calcareous Prediscosphaera ponticula (Bukry) Maastrichtian–Albian 35(23)
nannofossils
10 Calcareous Prediscosphaera cretacea (Arkhangelsky) Maastrichtian–Albian 35(24
nannofossils and 25)
10 Calcareous Prediscosphaera majungae Perch–Nielsen Maastrichtian–Campanian 35(26)
nannofossils
10 Calcareous Emiliania huxleyi (Lohmann) Extant–Ionian 35(27)
nannofossils
10 Calcareous Pseudoemiliania Gartner Ionian–Zanclean 35(28
nannofossils and 29)
10 Calcareous Pseudoemiliania lacunosa Kamptner Ionian–Zanclean 35(30)
nannofossils
10 Calcareous Rhabdosphaera Haeckel Extant–Ypresian 35(31)
nannofossils
10 Calcareous Reticulofenestra pseudoumbilicus (Gartner) Zanclean–Burdigalian 35(32)
nannofossils
10 Calcareous Reticulofenestra umbilicus (Levin) Rupelian–Lutetian 35(33
nannofossils and 34)
10 Calcareous Syracosphaera Lohmann Extant–Danian 36(1–5)
nannofossils
10 Calcareous Stoverius achylosus (Stover) Turonian–Aptian 36(6–8)
nannofossils
10 Calcareous Stoverius helotatus (Wind and Wise) Albian–Kimmeridgian 36(9)
nannofossils
10 Calcareous Cylindralithus coronatus Bukry Campanian–Cenomanian 36(10)
nannofossils
10 Calcareous Cylindralithus biarcus Bukry Maastrichtian–Cenomanian 36(11
nannofossils and 12)
10 Calcareous Mitrolithus Deflandre Toarcian–Hettangian 36(13)
nannofossils
10 Calcareous Crucirhabdus Rood Hay and Barnard Pliensbachian–Norian 36(14
nannofossils and 15)
10 Calcareous Nephrolithus frequens Górka Late Maastrichtian–Campanian 36(16)
nannofossils
10 Calcareous Grantarhabdus Black Campanian–Berriasian 36(17)
nannofossils
10 Calcareous Retecapsa Black Maastrichtian–Toarcian 36(18)
nannofossils
10 Calcareous Amphizygus Bukry Maastrichtian–Albian 36(19
nannofossils and 20)
10 Calcareous Cretarhabdus Bramlette and Martini Maastrichtian–Oxfordian 36(21)
nannofossils
10 Calcareous Blackites Hay and Towe Chattian–Thanetian 37(1–3)
nannofossils
(continued)
Species List 309

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Blackites gladius (Locker) Lutetian–Ypresian 37(4)
nannofossils
10 Calcareous Calcicalathina oblongata (Worsley) Barremian–Berriasian 37(5)
nannofossils
10 Calcareous Stephanolithion Deflandre Tithonian–Bajocian 38(1–7)
nannofossils
10 Calcareous Stephanolithion hexum Rood and Barnard Callovian–Bathonian 38(8–11)
nannofossils
10 Calcareous Stephanolithion speciosum Deflandre Callovian–Bajocian 38(12–
nannofossils 15)
10 Calcareous Stephanolithion speciosum var. speciosum Callovian–Bajocian 38(16)
nannofossils Deflandre
10 Calcareous Stephanolithion speciosum var. octum (Rood Callovian–Bajocian 38(17)
nannofossils and Barnard)
10 Calcareous Stephanolithion bigotii Deflandre Tithonian–Callovian 38(18)
nannofossils
10 Calcareous Stephanolithion bigotii var. bigotii Deflandre Tithonian–Callovian 38(19)
nannofossils
10 Calcareous Stephanolithion bigotii var. maximum Tithonian–Callovian 38(20)
nannofossils (Deflandre)
Stephanolithion laffittei Noël 38(21)
10 Calcareous Stephanolithion speciosum Deflandre Callovian–Bajocian 38(22)
nannofossils
10 Calcareous Truncatoscaphus hexaporus Moshkovitz and Albian–Bathonian 39(1–2)
nannofossils Ehrlich
10 Calcareous Truncatoscaphus octoporus Moshkovitz and Tithonian–Aalenian 39(3 and
nannofossils Ehrlich 4)
10 Calcareous Truncatoscaphus pauciramosus (Black) Hauterivian–Oxfordian 39(5 and
nannofossils 6)
10 Calcareous Truncatoscaphus intermedius Perch–Nielsen Tithonian–Late Kimmeridgian 39(7)
nannofossils
10 Calcareous Truncatoscaphus delftensis (Stradner and Turonian–Albian 39(8–11)
nannofossils Adamiker)
10 Calcareous Stradnerlithus asymmetricus (Rood, Hay and Tithonian–Aalenian 39(12)
nannofossils Barnard )
10 Calcareous Stradnerlithus tortuosus Noël Turonian–Late Kimmeridgian 39(13–
nannofossils 15)
10 Calcareous Stradnerlithus geometricus (Górka) Maastrichtian–Oxfordian 39(16)
nannofossils
10 Calcareous Stradnerlithus langii (Rood and Barnard) Toarcian 39(17)
nannofossils
10 Calcareous Stradnerlithus comptus Black Tithonian–Toarcian 39(18–
nannofossils 20)
10 Calcareous Stradnerlithus humilis Goy Toarcian 39(21)
nannofossils
10 Calcareous Stradnerlithus callomonii (Rood et al.) Oxfordian 39(22)
nannofossils
10 Calcareous Stradnerlithus fragilis (Rood and Barnard) Tithonian–Callovian 39(23)
nannofossils
10 Calcareous Stradnerlithus escovillensis (Rood and Early Callovian 39(24)
nannofossils Barnard)
10 Calcareous Thurmannolithion Grün and Zweili Oxfordian–Callovian 39(25)
nannofossils
(continued)
310 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Thurmannolithion clatratum Grün and Zweili Early Oxfordian–Early Callovian 39(26)
nannofossils
10 Calcareous Stephanolithion Deflandre Tithonian–Bajocian 39(27)
nannofossils
10 Calcareous Truncatoscaphus Rood, Hay and Barnard Turonian–Aalenian 39(28)
nannofossils
10 Calcareous Corollithion Stradner Maastrichtian–Valanginian 39(29)
nannofossils
10 Calcareous Cylindralithus Bramlette and Martini Maastrichtian–Albian 39(30)
nannofossils
10 Calcareous Rotelapillus Noël Maastrichtian–Tithonian 39(31)
nannofossils
10 Calcareous Rotelapillus crenulatus (Stover) Maastrichtian–Late Tithonian 39(32
nannofossils and 33)
10 Calcareous Rotelapillus munitus (Perch–Nielsen) Maastrichtian 39(34)
nannofossils
10 Calcareous Rotelapillus radians Noël Tithonian 39(35)
nannofossils
10 Calcareous Rotelapillus crenulatus (Stover) Maastrichtian–Late Tithonian 39(36)
nannofossils
10 Calcareous Schizosphaerella Deflandre and Dangeard Kimmeridgian–Hettangian 40(1–3)
nannofossils
10 Calcareous Schizosphaerella punctulata Deflandre and Kimmeridgian–Hettangian 40(4 and
nannofossils Dangeard 5)
10 Calcareous Sphenolithus heteromorphus Deflandre Serravallian–Burdigalian 41(1)
nannofossils
10 Calcareous Sphenolithus pseudoradians Bramlette and Chattian–Lutetian 41(2)
nannofossils Wilcoxon
10 Calcareous Sphenolithus belemnos Bramlette and Burdigalian 41(3)
nannofossils Wilcoxon
10 Calcareous Sphenolithus paratintinnabulum Bergen and Burdigalian–Chattian 41(4)
nannofossils de Kaenel
10 Calcareous Sphenolithus distentus (Martini) Chattian–Rupelian 41(5)
nannofossils
10 Calcareous Sphenolithus abies Deflandre in Deflandre and Gelasian–Serravallian 41(6 and
nannofossils Fert 7)
10 Calcareous Sphenolithus dissimilis Bukry and Percival Burdigalian–Aquitanian 41(8)
nannofossils
10 Calcareous Sphenolithus orphanknollensis Perch–Nielsen Lutetian–Ypresian 41(9)
nannofossils
10 Calcareous Sphenolithus editus Perch–Nielsen Ypresian 41(10)
nannofossils
10 Calcareous Sphenolithus radians Delfandre in Grassé Rupelian–Ypresian 41(11)
nannofossils
10 Calcareous Sphenolithus pseudoradians Bramlette and Chattian–Lutetian 41(12)
nannofossils Wilcoxon
10 Calcareous Sphenolithus predistentus Bramlette and Chattian–Lutetian 41(13)
nannofossils Wilcoxon
10 Calcareous Sphenolithus distentus (Martini) Chattian–Rupelian 41(14)
nannofossils
10 Calcareous Sphenolithus ciperoensis Bramlette and Chattian–Rupelian 41(15)
nannofossils Wilcoxon
10 Calcareous Sphenolithus spiniger Bukry Priabonian–Ypresian 41(16)
nannofossils
(continued)
Species List 311

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Sphenolithus conicus Bukry Burdigalian–Rupelian 41(17)
nannofossils
10 Calcareous Sphenolithus anarrhopus Bukry and Bramlette Ypresian–Selandian 41(18)
nannofossils
10 Calcareous Sphenolithus conspicuus Martini Ypresian 41(19)
nannofossils
10 Calcareous Sphenolithus belemnos Bramlette and Burdigalian 41(20)
nannofossils Wilcoxon
10 Calcareous Sphenolithus heteromorphus Deflandre Serravallian–Burdigalian 41(21)
nannofossils
10 Calcareous Sphenolithus delphix Bukry Chattian 41(22)
nannofossils
10 Calcareous Sphenolithus elongatus Perch–Nielsen Middle Eocene 41(23)
nannofossils
10 Calcareous Sphenolithus tribulosus Roth Rupelian–Priabonian 41(24)
nannofossils
10 Calcareous Sphenolithus abies Deflandre Gelasian–Serravallian 41(25)
nannofossils
10 Calcareous Sphenolithus primus Perch Nielsen Ypresian–Danian 41(26)
nannofossils
10 Calcareous Sphenolithus paratintinnabulum Bergen and Burdigalian–Chattian 41(27)
nannofossils de Kaenel
10 Calcareous Sphenolithus neoabies Bukry and Bramlette Zanclean–Burdigalian 41(28)
nannofossils
10 Calcareous Sphenolithus moriformis (Bronnimann and Zanclean–Selandian 41(29)
nannofossils Stradner)
10 Calcareous Sphenolithus grandis Haq and Berggren Tortonian–Serravallian 41(30)
nannofossils
10 Calcareous Sphenolithus furcatolithoides Locker Bartonian–Lutetian 41(31)
nannofossils
10 Calcareous Sphenolithus capricornutus Bukry and Aquitanian–Chattian 41(32)
nannofossils Percival
10 Calcareous Sphenolithus celsus Haq Rupelian–Priabonian 41(33)
nannofossils
10 Calcareous Sphenolithus obtusus Bukry Priabonian–Bartonian 41(34)
nannofossils
10 Calcareous Sphenolithus intercalaris Martini Priabonian–Bartonian 41(35)
nannofossils
10 Calcareous Thoracosphaera Kamptner Extant–Carnian 42(1–3)
nannofossils
10 Calcareous Triquetrorhabdulus Martini Serravallian–Rupelian 42(4)
nannofossils
10 Calcareous Triquetrorhabdulus carinatus Martini Burdigalian–Chattian 42(5)
nannofossils
10 Calcareous Isthmolithus Deflandre in Deflandre and Fert Rupelian–Ypresian 42(6 and
nannofossils 7)
10 Calcareous Isthmolithus recurvus Deflandre Rupelian–Priabonian 42(8)
nannofossils
10 Calcareous Reinhardtites levis Prins and Sissingh Maastrichtian–Campanian 42(9 and
nannofossils 10)
10 Calcareous Zeugrhabdotus embergeri (Noël) Selandian–Tithonian 42(11)
nannofossils
10 Calcareous Tranolithus orionatus (Reinhardt) Maastrichtian–Albian 42(12)
nannofossils
(continued)
312 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
10 Calcareous Zygodiscus Bramlette and Sullivan Ypresian–Selandian 42(13)
nannofossils
10 Calcareous Ansulasphaera helvetica Grün and Zweili Callovian–Bathonian 43(1 and
nannofossils 2)
10 Calcareous Lotharingius sigillatus (Stradner) Oxfordian–Pliensbachian 43(3)
nannofossils
10 Calcareous Lotharingius barozii Noël Aalenian–Toarcian 43(4)
nannofossils
10 Calcareous Lotharingius primitivus (Rood Hay and Toarcian 43(5)
nannofossils Barnard)
10 Calcareous Lotharingius crucicentralis (Medd) Oxfordian–Toarcian 43(6)
nannofossils
10 Calcareous Lotharingius hauffii Grün and Zweili Callovian–Pliensbachian 43(7)
nannofossils
10 Calcareous Cyclagelosphaera reinhardtii (Perch–Nielsen) Ypresian–Albian 43(8)
nannofossils
10 Calcareous Cyclagelosphaera margereliii Noël Thanetian–Bajocian 43(9)
nannofossils
10 Calcareous Cyclagelosphaera rotaclypeata Bukry Danian–Albian 43(10)
nannofossils
10 Calcareous Watznaueria fossacincta (Black) Late Maastrichtian–Bajocian 43(11)
nannofossils
10 Calcareous Watznaueria barnesiae (Black in Black and Maastrichtian–Bathonian 43(12)
nannofossils Barnes)
10 Calcareous Watznaueria biporta Bukry Maastrichtian–Tithonian 43(13)
nannofossils
10 Calcareous Watznaueria britannica (Stradner) Cenomanian–Bajocian 43(14–
nannofossils 16)
10 Calcareous Watznaueria ovata Bukry Maastrichtian–Callovian 43(17
nannofossils and 18)
10 Calcareous Watznaueria reinhardtii Rood, Hay and Tithonian–Oxfordian 43(19)
nannofossils Barnard
10 Calcareous Watznaueria fasciata Wind and Cepek Aptian–Berriasian 43(20)
nannofossils
10 Calcareous Watznaueria fossacincta (Black) Late Maastrichtian–Bajocian 43(21–
nannofossils 23)
10 Calcareous Watznaueria britannica (Stradner) Cenomanian–Bajocian 43(24)
nannofossils
10 Calcareous Ellipsagelosphaera plena Grün and Zweili Middle Callovian 43(25)
nannofossils
Calpionellids
11 Calpionellids Calpionella Lorenz Late Tithonian–Late Berriasian 1(1)
11 Calpionellids Calpionellites Colom Early Valanginian 1(2)
11 Calpionellids Calpionellopsis Colom Late Berriasian–Early 1(3)
Valanginian
11 Calpionellids Crassicollaria Remane Late Tithonian–Early Berriasian 1(4)
11 Calpionellids Tintinnopsella Colom Late Tithonian–Early 1(5)
Valanginian
11 Calpionellids Lorenziella Knauer and Nagy Late Berriasian–Early 1(6)
Valanginian
11 Calpionellids Calpionella alpina Lorenz Late Tithonian–Late Berriasian 1(7)
11 Calpionellids Calpionellites darderi (Colom) Early Valanginian 1(8)
11 Calpionellids Remaniella cadischiana (Colom) Berriasian 1(9)
(continued)
Species List 313

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
11 Calpionellids Tintinnopsella carpathica (Murgeanu and Late Tithonian–Early 1(10)
Filipescu) Valanginian
11 Calpionellids Lorenziella hungarica Knauer and Nagy Late Berriasian–Early 1(11)
Valanginian
11 Calpionellids Tintinnopsella longa (Colom) Late Tithonian–Early 1(14 and
Valanginian 15)
11 Calpionellids Calpionellopsis simplex (Colom) Late Berriasian–Early 1(16–18)
Valanginian
11 Calpionellids Tintinnopsella carpathica (Murgeanu and Late Tithonian–Early 1(19–25)
Filipescu) Valanginian
11 Calpionellids Calpionella alpina Lorenz Late Tithonian–Late Berriasian 2
11 Calpionellids Calpionella elliptica Cadisch Late Tithonian–Late Berriasian 2
11 Calpionellids Calpionella grandalpina Nagy Late Tithonian–Late Berriasian 2
11 Calpionellids Calpionella minuta Houša Late Tithonian–Late Berriasian 2
11 Calpionellids Calpionellites caravacensis Allemann Early Valanginian 2
11 Calpionellids Calpionellites coronatus (Trejo) Early Valanginian 2
11 Calpionellids Calpionellites darderi (Colom) Early Valanginian 2
11 Calpionellids Calpionellites major (Colom) Early Valanginian 2
11 Calpionellids Calpionellites uncinata Cita and Pasquar Early Valanginian 2
11 Calpionellids Calpionellopsella maldonadoi Trejo Early Albian 2
11 Calpionellids Calpionellopsis oblonga (Cadisch) Late Berriasian–Early 2
Valanginian
11 Calpionellids Calpionellopsis simplex (Colom) Late Berriasian–Early 2
Valanginian
11 Calpionellids Chitinoidella bermudezi (Furrazola– Middle Tithonian 2
Bermudezj)
11 Calpionellids Chitinoidella boneti Doben Middle Tithonian 2
11 Calpionellids Crassicollaria colomi Doben Middle Tithonian 2
11 Calpionellids Chitinoidella cubensis (Furrazola-Bermudez) Middle Tithonian 2
11 Calpionellids Chitinoidella dobeni Borza Middle Tithonian 2
11 Calpionellids Chitinoidella slovenica (Borza) Middle Tithonian 2
11 Calpionellids Chitinoidella tithonica Borza Middle Tithonian 2
11 Calpionellids Colomiella mexicana Bonet Early Albian 2
11 Calpionellids Colomiella recta Bonet Early Albian 2
11 Calpionellids Colomiella semiloricara Early Albian 2
11 Calpionellids Crassicollaria Remane Late Tithonian–Early Berriasian 2
11 Calpionellids Crassicollaria brevis Remane Late Tithonian–Early Berriasian 2
11 Calpionellids Crassicollaria colomi Doben Late Tithonian–Early Berriasian 2
11 Calpionellids Crassicollaria intermedia (Durand Delga) Late Tithonian–Early Berriasian 2
11 Calpionellids Crassicollaria massutiniana (Colom) Late Tithonian–Early Berriasian 2
11 Calpionellids Crassicollaria parvula Remane Late Tithonian–Early Berriasian 2
11 Calpionellids Deflandronella tenius Trejo Aptian–Albian 2
11 Calpionellids Deflandronella veracruzana (Trejo) Aptian–Albian 2
11 Calpionellids Lorenziella hungarica Knauer and Nagy Late Berriasian–Early 2
Valanginian
11 Calpionellids Lorenzlella plicata Late Berriasian–Early 2
Valanginian
11 Calpionellids Parachitinoidella cuvillieri Trejo Aptian–Albian 2
11 Calpionellids Parachitinoidella ornata Trejo Aptian–Albian 2
11 Calpionellids Praecalpionellites murgeanui (Pop) Valanginian 2
11 Calpionellids Praecalpionellites siriniaensis Pop Valanginian 2
(continued)
314 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
11 Calpionellids Praecolomiella boneti Borza Aptian–Albian 2
11 Calpionellids Praecolomiella trejoi Borza Aptian–Albian 2
11 Calpionellids Praetintinnopsella andrusovi Borza Late Tithonian 2
11 Calpionellids Remaniella borzai Pop Late Tithonian–Late Berriasian 2
11 Calpionellids Remaniella cadischiana (Colom) Late Tithonian–Late Berriasian 2
11 Calpionellids Remaniella ferasini (Catalano) Late Tithonian–Late Berriasian 2
11 Calpionellids Remaniella filipescui Pop Late Tithonian–Late Berriasian 2
11 Calpionellids Sturiella oblonga Borza Valanginian 2
11 Calpionellids Tintinnopsella carpathica (Murgeanu and Late Tithonian–Early 2
Filipescu) Valanginian
11 Calpionellids Tintinnopsella doliphormis (Colom) Late Tithonian–Early 2
Valanginian
11 Calpionellids Tintinnopsella emanei (Borza) Late Tithonian–Early 2
Valanginian
11 Calpionellids Tintinnopsella longa (Colom) Late Tithonian–Early 2
Valanginian
11 Calpionellids Tintinnopsella subacuta (Colom) Late Tithonian–Early 2
Valanginian
11 Calpionellids Borzaiella slovenica (Borza) Tithonian 3
11 Calpionellids Carpathella rumanica Pop Late Tithonian–Early Berriasian 3
11 Calpionellids Chitinoidella boneti Doben Middle Tithonian 3
11 Calpionellids Chitinoidella elongata Pop Middle Tithonian 3
11 Calpionellids Daciella danubica Pop Middle Berriasian–Late 3
Berriasian
11 Calpionellids Daciella svintensis Late Tithonian 3
11 Calpionellids Dobeniella bermudezi (Furrazola–Bermudez) Middle Berriasian–Late 3
Berriasian
11 Calpionellids Dobeniella colomi (Borza) Early Tithonian 3
11 Calpionellids Dobeniella cubensis (Furrazola–Bermudez) Late Berriasian 3
11 Calpionellids Dobeniella tithonica (Borza) Late Tithonian–Early Berriasian 3
11 Calpionellids Longicollaria dobeni (Borza) Tithonian 3
11 Calpionellids Longicollaria insueta (Řehánek) Middle Berriasian 3
11 Calpionellids Popiella oblongata Reháková Tithonian 3
11 Calpionellids Chitinoidella boneti Doben Middle Tithonian 4(1–2)
11 Calpionellids Chitinoidella carthagensis Sallouhi Boughdiri Middle Tithonian 4(3)
and Cordey
11 Calpionellids Chitinoidella elongata Pop Middle Tithonian 4(4–5)
11 Calpionellids Chitinoidella sp. indet Middle Tithonian 4(6)
11 Calpionellids Chitinoidella hegarati Sallouhi Boughdiri and Middle Tithonian 4(7–8)
Cordey
11 Calpionellids Chitinoidella popi Sallouhi Boughdiri and Middle Tithonian 4(9–10)
Cordey
11 Calpionellids Chitinoidella sp. indet Middle Tithonian 4(11)
11 Calpionellids Chitinoidella boneti Doben Middle Tithonian 4(12–18)
11 Calpionellids Chitinoidella carthagensis Sallouhi Boughdiri Middle Tithonian 4(19–21)
and Cordey
11 Calpionellids Chitinoidella elongata Pop Middle Tithonian 4(22–25)
11 Calpionellids Chitinoidella bermudezi (Furrazola– Middle Tithonian 4(26)
Bermudezj)
11 Calpionellids Chitinoidella cristobalensis (Furrazola– Middle Tithonian 4(27)
Bermudezj)
(continued)
Species List 315

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
11 Calpionellids Calpionella alpina Lorenz—large form Late Tithonian–Middle 5(1)
Berriasian
11 Calpionellids Calpionella alpina Lorenz—medium form Late Tithonian–Middle 5(2)
Berriasian
11 Calpionellids Calpionella alpina Lorenz—large form Late Tithonian–Middle 5(3)
(C. “grandalpina”) Berriasian
11 Calpionellids Calpionella sp. (C. “elliptalpina”) Late Tithonian–Middle 5(4)
Berriasian
11 Calpionellids Calpionella alpina Lorenz—medium spherical Late Tithonian–Middle 5(5)
form Berriasian
11 Calpionellids Calpionella alpina Lorenz—small slightly Late Tithonian–Middle 5(6)
elliptical form Berriasian
11 Calpionellids Calpionella alpina Lorenz Late Tithonian–Middle 5(7–14)
Berriasian
11 Calpionellids Calpionella elliptica Cadisch Late Tithonian–Middle 5(15–17)
Berriasian
11 Calpionellids Calpionella minuta Houša Late Tithonian–Middle 5(18 and
Berriasian 19)
11 Calpionellids Crassicollaria brevis Remane Late Tithonian–Early Berriasian 6(1 and
2)
11 Calpionellids Crassicollaria colomi Doben Late Tithonian–Early Berriasian 6(3 and
4)
11 Calpionellids Crassicollaria intermedia Durand Delga Late Tithonian–Early Berriasian 6(5–8)
11 Calpionellids Crassicollaria massutiniana (Colom) Late Tithonian–Early Berriasian 6(9)
11 Calpionellids Crassicollaria parvula Remane Late Tithonian–Early Berriasian 6(10–13)
11 Calpionellids Crassicollaria intermedia Durand Delga Late Tithonian–Early Berriasian 6(14)
11 Calpionellids Praetintinnopsella andrusovi Borza Late Tithonian 6(15)
11 Calpionellids Remaniella ferasini (Catalano) Late Tithonian–Late Berriasian 7(1–3)
11 Calpionellids Remaniella catalanoi Pop Late Tithonian–Late Berriasian 7(4–5)
11 Calpionellids Remaniella borzai Pop Late Tithonian–Late Berriasian 7(6)
11 Calpionellids Remaniella duranddelgai Pop Late Tithonian–Late Berriasian 7(7–10)
11 Calpionellids Remaniella filipescui Pop Late Tithonian–Late Berriasian 7(11)
11 Calpionellids Remaniella cadischiana (Colom) Late Tithonian–Late Berriasian 7(12 and
13)
11 Calpionellids Remaniella colomi Pop Late Tithonian–Late Berriasian 7(14 and
15)
11 Calpionellids Sturiella oblonga Borza Valanginian 7(16)
11 Calpionellids Chitinoidella sp. indet. Middle Tithonian 8(1)
11 Calpionellids Dobeniella colomi (Borza) Early Tithonian 8(2)
11 Calpionellids Daceilla danubica Pop Late Tithonian–Early 8(3 and
Valanginian 4)
11 Calpionellids Tintinnopsella carpathica (Murgeanu and Late Tithonian–Early 9(1–12)
Filipescu) Valanginian
11 Calpionellids Tintinnopsella longa (Colom) Late Tithonian–Early 9(13–17)
Valanginian
11 Calpionellids Lorenziella gr. hungarica Knauer and Nagy Late Berriasian–Early 9(18)
Valanginian
11 Calpionellids Lorenziella hungarica Knauer and Nagy Late Berriasian–Early 9(19 and
Valanginian 20)
11 Calpionellids Lorenzlella ruggierii (Catalano) Late Berriasian–Early 9(21)
Valanginian
11 Calpionellids Praecalpionellites dadayi (Knauer) Valanginian 9(22)
(continued)
316 Species List

Chapter Chapter name Species illustrated Age Fig. Nos.

No.
11 Calpionellids Praecalpionellites murgeanui (Pop) Valanginian 9(23 and
24)
11 Calpionellids Praecalpionellites dadayi (Knauer) Valanginian 9(25)
11 Calpionellids Praecalpionellites siriniaensis Pop Valanginian 9(26)
11 Calpionellids Praecalpionellites filipescui (Pop) Valanginian 9(27)
11 Calpionellids Praecalpionellites murgeanui (Pop) Valanginian 9(28)
11 Calpionellids Calpionellites major (Colom) Early Valanginian 10(1 and
2)
11 Calpionellids Calpionellites coronatus (Trejo) Early Valanginian 10(3 and
4)
11 Calpionellids Calpionellites darderi (Colom) Early Valanginian 10(5–8)
11 Calpionellids Calpionellites caravacensis Allemann Early Valanginian 10(9)
11 Calpionellids Calpionellopsis oblonga (Cadisch) Late Berriasian–Early 10(10)
Valanginian
11 Calpionellids Calpionellopsis simplex (Colom) Late Berriasian–Early 10(11
Valanginian and 12)
11 Calpionellids Dobeniella bermudezi (Furrazola–Bermudez) Middle Berriasian–Late 11(1 and
Berriasian 2)
11 Calpionellids Dobeniella cubensis (Furrazola–Bermudez) Late Berriasian 11(3)
11 Calpionellids Borzaiella atava Grün and Blau Tithonian 11(4 and
5)
11 Calpionellids Borzaiella slovenica (Borza) Tithonian 11(6)
11 Calpionellids Longicollaria dobeni (Borza) Late Tithonian–Early Berriasian 11(7)
11 Calpionellids Colomiella mexicana Bonet Albian 11(8)
11 Calpionellids Colomiella recta Bonet Albian 11(9 and
10)
11 Calpionellids Daciella danubica Pop Early Tithonian 12(1 and
2)
11 Calpionellids Dobeniella colomi (Borza) Early Tithonian 12(3)
11 Calpionellids Longicollaria dobeni (Borza) Tithonian 12(4)
11 Calpionellids Dobeniella cf. tithonica (Borza) Late Tithonian–Early Berriasian 12(5)
11 Calpionellids Borzaiella slovenica (Borza) Middle Berriasian–Late 12(6)
Berriasian
11 Calpionellids Longicollaria cf. insueta (Řehánek) Middle Berriasian 12(7)
11 Calpionellids Popiella oblongata Reháková Tithonian 12(8)
11 Calpionellids Praetintinnopsella andrusovi Borza Late Tithonian 12(9)
 Index

A Axial cross (Abb +), 218

Abdomen, 121 Axis, 10, 15–17, 45, 48, 56, 57, 118, 132, 134, 175, 178,
Aboral, 8, 13, 107, 261 202, 216, 227, 263
Aboral and Adoral, 261 Axopodia, 117
Aboral pole, 11, 13
Aboral scar, 13
Acanthomorphs, 27, 29–31, 33, 34, 37 B
Acanthomorphs (acantha = thorn), 33 Baculate, 58
Additional taxon-based terminology, 222 Band or segment, 134
Adductor muscles, 143, 147, 148 Bar, 28, 37, 96, 100, 103, 117–121, 143, 216, 218, 227
Adductor muscle scar group, 147, 148 Bar-type conodonts, 96
Adont (or unipartite), 143 Basal, 9, 13, 15, 24, 58, 94, 100, 103, 104, 118, 121, 123,
Agglutinated, 182, 183, 185 149, 178, 180, 181, 216, 221
Alate, 104, 108 Basal body, 94
Amphidont, 145 Basal callus, 15
Anicephalic lobe, 120 Basal cavity, 94, 103, 104
Antapical series, 76, 77 Basal edge, 15–17
Anterior, 75–77, 80, 85, 100, 110, 145, 148, 224, 227 Basal–extraumbilical, 178, 180
Anterior/posterior, 224 Basal interiomarginal, 178, 180
Anti-apertural pole, 13 Basal margin, 9, 15
Anticlockwise/clockwise imbrication, 212 Basal pore, 15
A or V -shaped, - chambers, 178, 179 Basal rim, 17
Apertural plug, 19 Basal scar, 13, 15
Apertural pole, 8, 9, 15, 17 Base, 3, 4, 7, 8, 11–13, 15–19, 58, 104, 120, 145, 181,
Aperture, 3, 8, 9, 11, 13, 15–18, 44, 48, 50, 52, 53, 213, 215, 250
55–57, 60, 121, 171, 175, 178–181, 228, 261 Biconvex, 178, 179
Aperture form, 181 Bifurcate tip, 225
Aperture not visible, 181 Bipennate, 104, 106, 108
Apex, 8, 12, 15, 16, 18, 58, 76, 77 Biserial, 175, 177, 178
Apical axis, 132, 134, 140 Blade, 96, 100, 103, 215, 228
Apical cycle, 225, 228 Blade-type conodonts, 96
Apical plane, 134 Blanket, 208, 218, 227
Apical region, 224 Block, 196, 208, 210, 213
Apical spine, 120, 228 Boss, 18, 219, 225
Apical structure, 11, 15–17 Braarudosphaeraceae, 219, 222, 223, 235
Appendices, 15, 18 Breviform, 104, 108
Arch, 85, 224 Brick, 178, 179
Archeopyle, 83, 85 Bridge, 215, 216, 218, 227
Arcuate lunear–arch, 181 Brief note on classification and illustration of major
Areal, 179–181 families, 229
Areolae, 132, 134, 141 Bulb, 228
Arm, 218, 219, 224 Butting, 212
Asymmetrical, 104, 205, 218

© Springer Nature India Private Limited 2020 317

S. Jain, Fundamentals of Invertebrate Palaeontology, Springer Geology,
https://doi.org/10.1007/978-81-322-3962-8
318 Index

C Coccolith outline and axial ratio, 205

Calicalith, 215 Coiled chain (=helicoidal chain), 16
Calicate, 219 Coiled spirally, 175, 176
Callus, 9, 11, 15 Collar, 15–17, 34, 120, 210, 211, 261, 263
Callus (basal), 15 Collarette, 3, 4, 11, 12, 15–19, 34, 120, 211, 261
Calpionellidae Bonet, 261, 265 Collarette (=collar), 16
Calymma, 117 Collar structure, 120
Calyptrolith, 215 Colomiellidae Toso, 265
Calyx, 219 Colpoidorate, 55
Canal, 74, 145, 147, 210, 219, 228 Colporoidate, 55
Carapace and associated structures, The, 143 Colporus, 53
Carapace length, 148 Colpus, 52–54
Carapace orientation, 148 Column cycle, 225
Cardinal margin (hinge margin), 145, 148 Compact, 219
Carina, 9–11, 15–18, 100 Composite elements, 96
Carminate and angulate, 107, 110 Compound, 1, 33, 50, 53, 54, 96, 228
Catenary structure (chain-like structure), 11, 16, 17 Cones, 11, 93, 113, 118, 175, 216, 218, 219, 225
Caudal horn, 261, 263 Conical, 8, 175, 211, 218, 219, 234
Cavate, 79, 82, 84, 87, 215 Conical to subconical, 175
Cavity, 16, 84, 94, 103, 104, 145, 210, 213, 215, 218, Coniform elements, 104, 110
228, 263 Conjunct bar, 216, 227
Cell, 11, 18, 27, 43, 44, 67, 74, 75, 77, 117–119, Conochitinidae, 19, 22, 24
131–134, 136, 139, 140, 147, 171, 172, 178, 183, Constriction, 126, 134, 228
193, 194, 265 Contact-surface, 208, 212, 225
Cell wall structure, 134 Convex outline, 222
Central area, 195, 196, 202, 205, 207, 216–219, 224, 225, Copula, 3, 9, 11, 12, 15, 16
227, 229 Core, 221, 228
Central area and rim, 202, 205 Costae, 37, 184, 185
Central body, 28, 33, 34, 221, 222, 225 Crescentic, 178, 179
Central capsule, 117 Crests, 11, 16, 19
Central cavity, 1, 16, 228 Cribrate, 181, 182, 218
Central muscle scar field, 147 Cribrum, 134
Central nodule, 132 Crimp, 100
Central opening, 205, 218, 227, 228 Cross, 103, 119, 213, 218, 263
Central opening/central cavity, 228 Cross-bar, 218
Cephalus, 120, 122 Crown, 11, 19, 29, 94, 104, 211
Ceratolith, 224 Cryptopylome, 34
Ceratolithaceae, 223, 224, 239 Crystallite, 194, 196, 213, 215
Chain, 9, 11, 16, 17, 19, 24, 193 Crystallite arrangement, 213
Chamber, 1, 4, 8, 9, 11, 12, 15–19, 24, 132, 171, 175, Crystallographic orientation, 219, 220
177–179, 181, 183 Crystallography, 207, 219
Chamber arrangement, 171, 175, 177, 178 Crystallolith, 215
Chamber form, 171, 178, 179 Crystal unit, 196, 207, 208, 212
Chamber shapes, 4, 9 Cup, 104, 265
Chitinoidellidae Trejo, 265 Curved, 16, 17, 34, 104, 108, 134, 183, 184, 207, 210,
Chitinozoan affinity, 19 261
Chorate, 79, 81–84, 87, 88 Cusp, 104, 106
Cicatricose, 60 Cycle, 43, 131, 171, 205, 208, 210, 212, 225, 228
Cingular series, 76, 77 Cylindrical, 4, 8, 16, 18, 24, 57, 58, 221
Cingulum, 75–77, 79, 81, 87, 132, 134 Cyst types, 79, 84, 87
Circular (centric), 133 Cyst wall (phragma), 84
Classification, 19, 34, 74, 78, 84, 110, 135, 136, 140, 151,
152, 171, 194, 229, 230, 265
Clavate, 58, 127 D
Claviform, 8, 219 Dendroid, 175, 176
Closed central area, 218 Dentate keel, 224
Cluster, 11, 12, 16, 107 Denticle, 94, 96, 145
Coccolith arrangement, 212 Depressed, 149, 183, 184
Coccolith orientation, 196 Depression, 55, 60, 74, 76, 183, 210, 215, 225
Index 319

Desmochitinidae, 19, 21, 24 External/Outer/Outward, 202

Dextral/sinistral, 212, 227 External structures, 10, 11, 16
Dextral/sinistral obliquity, 212 Extracardinal margin (free margin), 145, 149
Dextrogyral face, 225 Extraumbilical, 178–180
Dextrogyre, 210
Diacromorphs (akron = summit), 33
Diagonal, 139, 216, 218, 227 F
Diagonal cross (Abb X), 218 Fasciculiths, 225, 227
Diameter (= D), 16 Faveolate, 60
Dibrachiate, 219, 224 Felt-like, 11
Digyrate, 104, 108, 110 Fenestra, 55, 225
Directions on the rim, 211 Fenestrate, 16, 55
Disc, 193, 196, 202, 205, 215, 221, 224, 225 Fistulose, 184, 185
Discoasters, 194, 205, 210, 221, 222, 224, 225, 241 Flagella, 74
Disjunct bar, 216, 227 Flange, 33, 37, 100, 107, 210, 213, 227, 228
Distal, 9, 11, 17, 44, 45, 51–53, 202, 210–213, 215, 216, Flank, 8, 9, 11, 15–17, 19
218–220, 225, 227, 239, 242, 249, 252 Flare and taper, 211
Distal cover, 213, 215, 227 Flare surfaces, 212
Dolabrate, 104 Flexure, 7, 17, 19
Dorsal muscle scar field, 147 Flosculolith, 215
Dorsal view, 80, 81, 83, 86, 148, 156, 158–169 Foot, 60, 122, 218
Duocrystalline, 228 Foramen, 134, 175, 178
Duplicatures, 143, 145 Fossulate, 29, 58
Foveolate, 11
Fragariolith, 215
E Free ray, 221, 222, 224, 225
Earliest plants, 60 Frontal group, 148
Echinate/spinate, 58 Frontal view, 148, 242, 256
Ectophragm, 33, 34, 37, 84 Frustule, 132–134, 140, 141
Ectophragm (Periphragm), 33 Furrow, 52, 53, 74, 75, 104
Ectoplasm, 117, 119, 171, 172, 178 Fusiform, 29, 33, 34, 175, 176
Element arrangement as seen in plan view, 212
Element curvature, 210
Element modifications, 208, 209 G
Element notation and apparatus, 107 Gemmate, 58
Elements, 1, 11, 15, 17, 18, 29, 57, 58, 60, 84, 93, 94, 96, General history, 24, 151
100, 104–108, 110, 113, 117, 118, 134, 143, 145, General history and distribution, 24
149, 154, 183, 194–196, 205, 207–213, 218, General morphology, 1, 29, 74, 93, 143, 235, 261
220–222, 224, 225, 227, 228, 263 Geniculate, 104
Element shapes, 208, 209 Geological history, 34, 87, 113, 122, 134, 171, 185, 186,
Elevated, 183, 205, 216, 218 230, 265
Elliptical, 37, 132, 133, 141, 196, 202, 207, 250, 269 Geological history and distribution, 34
Elliptical (Pennate), 133 Germ denticles, 94
Elongate, 19, 34, 67, 93, 96, 118, 122, 132, 136, 145, Girdle, 75, 77, 85, 132, 134, 137, 138, 141
175, 176, 208, 210, 218, 228 Glabrous, 11, 17, 18
End, 1, 8, 9, 13, 15, 16, 18, 24, 37, 38, 52, 53, 57, 76, Gliscolith, 215
100, 118, 132, 148, 149, 154, 185, 194, 202, 207, Globose-ovate, 175, 176
208, 219, 224, 225, 227, 228, 261 Granule, 11, 58, 84, 183, 184, 208
Endophragm, 33, 34, 84 Grill, 216, 218
Endoplasm, 117, 119, 171, 172, 178 Gross morphology, 134
Endopylome, 34, 35 Growth lamellae, 94, 100, 104
Episome, 75, 76
Epitheca, 75–78, 85, 132
Epityche, 34, 35 H
Eucephalic lobe, 120 Helicoid catenary linkage, 16
Excystment opening, 31, 33–35 Helicosphaera, 205, 208, 215, 216, 218, 227, 243
Exine ornamentation, 56, 58 Helladolith, 215
Extensiform, 104, 110 Hemisolenic, 149
External morphological features, 171 Hemispherical, 4, 151
320 Index

Herkomorphs (herkos = wall), 33 Leaf-like, 175, 176, 215

Heterococcoliths, 193, 194, 196, 205 Left/right, 224, 227
Heterocolpate, 54 Length/width/height, 202
Heteromorphic, 33 Lenticular, 4, 207
Hexagonal, 119, 196, 213 Lete, 52
Hexagonal meshwork, 213 Ligament, 143
High angle, 216 Linear chain, 17
Hinge types, 146 Linear linkage arrangements, 11, 13
Hispid to pustulose, 184, 185 Linkage by double adhesion, 12
Hole, 18, 58, 85, 145, 182, 183, 210 Linkage by simple juxtaposition, 11
Holococcolith, 194, 196, 213–215, 229 Linkage structures, 17
Holosolenic, 149 Lip, 11, 12, 15, 17, 19, 181, 182, 229, 265
Homomorphic, 33 Lobe, 100, 120, 151
Horizontal, 18, 57, 104, 175, 202, 211, 212, 215, 228 Long free rays, 222, 225
Horns, 24, 76, 84, 122, 151, 154, 261, 263 Longitudinal, 17, 45, 74, 110, 132, 202, 216, 218, 225
Hyaline, 94, 110, 182, 183, 243, 261, 265 Longitudinal axis, 17
Hyposome, 75, 76 Longitudinal ridge, 225
Hypotheca, 75–77, 132 Loop-shaped, 181, 182
Lophate/fenestrate, 55
Lorica, 261–263, 265
I Low angle, 212, 216
Imbrication, 212 Lumber structure, 121
Imbrication angle, 212
Inner lamella, 94, 145
Inner layer (=ectoderm/endoderm), 17 M
Intercalaries, 76, 77, 80, 81, 83, 85, 86, 138, 255 Main spine, 119
Intercalary bands, 77 Mandibular group, 148
Interlocking, 143, 193, 212 Marginal structures (spines, denticles or ridges), 145
Internal/inner/inward, 16–19, 45, 52, 56, 57, 60, 81, 84, Margin (= basal edge), 17
94, 106, 117, 119, 145, 171, 172, 195, 196, 202, Margin (=Basal margin), 9
205, 211, 212, 263 Median bar, 120, 121
Inversely oriented bar, 227 Median split, 34
Irregular, 11, 58, 60, 175–179 Merodont, 145
Irregularly elliptical, 207 Mesh-like, 11, 17, 218
Irregular streptospiral, 177, 178 Mesophragm, 33
Microgranular, 182, 183, 261, 265
Miliolid, 178
K Monocrystalline, 228
Keel, 100, 104, 110, 171, 184, 210, 224 Monoporate, 53
Keeled, 183, 184 Monosulcate, 45, 53
Kink, 208 Morphogroups, 29
Kofoid’s alphanumeric system, 77 Morphological types, 1, 43, 196, 214, 215
Morphology, 1, 12, 18, 29, 31, 33, 34, 43–45, 74, 93, 117,
118, 120, 121, 131, 134, 143, 149, 151, 194, 196,
L 222, 229, 235, 261, 262
Labiatiform, 218 Morphology of coccoliths, 194
Laciniated, 10, 17 Mucron, 11, 12, 15, 17
Laevogyral face, 225 Multi orate, 54
Laevogyre, 210 Multiradiate, 221
Lagenochitinidae, 23, 24 Murolith, 195, 205, 210
Lamellae, 94, 100, 103, 104, 145 Muscle scars, 147–149, 156, 159, 162, 164, 165, 167, 168
Lamellae and cardinals, 145
Lamina, 208, 224
Laminolith, 215 N
Lateral bar, 218 Nannoconids, 227, 228
Lateral cycles, 228 Nannolith shapes, 219, 221
Lateral or side view, 148 Narrow central area, 216
Lateral or transverse split, 34 Nassellarians, 118, 120–122, 126, 129
Lath, 183, 208, 210, 224 Neck, 8, 9, 11, 12, 15–19, 24, 34, 181, 182
Index 321

Neck (= apertural tube), 9, 17 Perforation, 3, 118, 132, 134, 171, 210, 213, 215, 218
Net, 131, 184, 218 Pericolpus, 53
Netromorphs (netro = spindle), 33 Periphragm, 33, 84
Node, 107, 149, 210, 224 Periporus, 52
Non-geniculate, 104, 110 Pervalvar axis, 132
Non-overlapping, 212 Petal/petaloid element, 208
No primary but multiple apertures, 179–181 Pit, 17, 18, 60, 94, 100, 183, 210, 215
Normal central-area, 216 Placolith, 195, 205
Normal (lateral), 145 Planar, 216
Normally oriented bar, 227 Planispiral, 175, 177, 178
Nuclei, 27, 117, 119, 132, 171, 178 Planolith, 195, 205
Plate, 37, 76, 77, 79–81, 84, 85, 100, 132, 145, 193, 215,
216, 218, 224, 227
O Platform-type conodonts, 100
Obliquity, 212 Plug, 11, 15, 18, 19, 34, 213
Oblong, 207, 261 Polarity, 44, 45, 52, 136, 138
Offset, 81, 208, 212, 218 Pollen morphology, 44
Offset cross, 218 Pollen units, 44, 47
Oomorphs (oon = egg), 33 Polygonal, 33, 60, 196, 207
Open central area, 218 Polygonomorphs (gonia = angle), 33
Open (simple), 145 Ponticuli, 183, 184
Operculum, 8, 9, 11, 12, 15, 17–19, 24, 34, 35, 85 Porcelaneous, 182, 183
Oral, 8, 11, 16, 18, 19, 94, 261 Pore, 15, 18, 29, 37, 52–55, 58, 74, 119, 121, 122, 132,
Oral opening (=aperture), 261 134, 136, 145, 147, 181–183, 213, 215, 219, 250
Oral scar, 18 Pore canals, 145, 147
Organic, 1, 19, 27, 29, 30, 84, 117, 118, 182, 183, 196, Pores in rows, 181
261, 265 Porocolpate, 55
Orientation, 45, 84, 148, 183, 195, 196, 208, 209, 211, Pororus, 53
212, 216, 219, 220, 224, 225, 227, 228, 263 Porus, 52
Orientation in plan view, 216 Position of primary aperture, 171, 178, 180
Orientation in profile, 216 Post-abdominal segments, 121
Origin, 1, 3, 27, 28, 63, 219, 227 Postcingular series, 76
Ornamentation, 9–11, 15–19, 29, 30, 32–34, 37, 44, 51, Posterior, 3, 75–77, 81, 83, 110, 145, 148, 151, 227
52, 56, 58, 81, 84, 151, 171, 175, 184, 185 Posterior view, 3, 148
Ornamentation and surface patterns, 9 Precingular series, 76, 77, 85
Other useful terms, 225 Prismatomorphs (prisma = prism), 33
Outer lamella, 145 Process, 9, 11, 15, 17–19, 29, 31, 33, 37, 58, 79, 81,
Outer layer (=periderm), 18 83–85, 87, 104, 106, 107, 122, 136, 140, 215, 218,
Overlapping, 132, 151, 212 219
Ovoid, 4, 207 Processes (=appendices), 18
Processes/spines, 9, 33
Prosome, 11, 15, 18, 19
P Protrusion, 29, 33, 34, 100, 184, 210, 213, 218, 219, 225
Parasyncolpate, 53 Proximal, 44, 45, 51, 52, 132, 195, 205, 211–213, 215,
Partial split (cryptosuture), 34 216, 219, 225, 227, 228, 232, 242, 249, 250, 252
Parts, 11, 13, 15–18, 34, 45, 52, 58, 60, 63, 84, 93, 94, Proximal cycle, 228
104, 120, 134, 138, 143, 148, 149, 154, 193, 195, Proximal extension, 225
196, 205, 208–210, 212–215, 218, 219, 224, 225, Proximal flange, 213
227, 228, 263 Proximal plate, 215, 227
Parts of rims, 209, 210 Proximal pore, 219
Pastinate, 104, 107, 110 Proximal ring, 213
Pectinform elements, 104, 106 Proximal surface, 227
Peduncle, 11, 15, 18 Proximate, 79, 82, 84, 87
Pentalith, 222, 224 Proximochorate, 79, 82, 84, 87
Pentaradiate, 221 Pseudopylome, 34, 35
Perforate, 16, 58, 117, 132, 183 Pseudoraphe, 132, 133
Perforated, 10, 18, 57, 132, 134, 182, 218 Psilate/laevigate, 58
Perforated plate, 132, 218 Pteromorphs (pteros = wing), 33
322 Index

Punctae, 132 Sexual dimorphism, 149, 151

Pylome, 33–35 Shape, 1, 4, 7, 9, 11, 29, 33, 34, 44, 46, 51, 53, 56–58, 60,
Pyriform oval convex, 178, 179 67, 79, 80, 84, 96, 104, 107, 119, 120, 122, 133,
136, 141, 145, 149, 151, 175, 176, 181, 183, 193,
194, 196, 205–208, 213, 216, 219, 221, 222, 263,
Q 265
Quadriramate, 104 Shield, 195, 205, 210–212, 227
Short free rays, 222, 225
Shoulder, 4, 8, 17, 19, 122
R Side, 56, 75, 100, 148, 178, 195, 202, 207, 211, 218, 219,
Radial, 44, 117–119, 132, 136, 145, 196, 208, 211–213, 225, 227, 249
219, 221 Sieve plate, 132
Radial (marginal), 145 Sieve-type, 145, 147
Radiate, 44, 117, 181, 182, 219, 221 Simple elements, 104
Raised, 17, 44, 149, 183, 184 Simple tip, 225
Ramiform elements, 104 Sinistral/dextral, 216
Raphe, 131–133, 136, 138, 140, 141 Size, 29, 33, 43, 44, 55, 57, 94, 118, 143, 149, 151, 171,
Ray, 208, 221, 222, 224, 225 183, 193, 194, 196, 208, 213, 229, 243, 263
Ray-related terms, 224 Skeletal structures, 110, 118–121, 193
Ray tip, 225 Skeleton, 27, 93, 110, 117–120
Reinforced linkage, 12 Sleeve, 8, 11, 16, 17, 19
Relative width, 216 Slit, 44, 52, 132, 181, 182, 210
Reniform, 207 Smooth keel, 224
Reticulate, 18, 29, 60, 134, 184, 185 Smooth or scabrate, 11
Reticulate/Reticulum, 60 Solid, 18, 33, 104, 118, 120, 215
Rhomboid, 213 Special, 52, 123, 181, 182
Rica, 11, 18 Sphaeromorphs, 27–30, 34, 35
Ridge, 11, 44, 55, 60, 107, 134, 143, 145, 149, 183, 184, Sphaeromorphs and acanthomorphs, 29
210, 225 Sphaeronorphs (spharia = ball), 33
Rim, 17, 34, 195, 196, 202, 205, 207, 210, 212, 213, 215, Sphenoliths, 228, 255
216, 218, 227, 229 Spheres, 118, 119
Ring-shaped, 207 Spherical, 4, 29, 33, 34, 37, 58, 67, 117, 119, 122, 269
Rod, 57, 117, 148, 183, 208, 210, 224, 250 Spherical–globular, 178, 179
Rod-shaped, 210, 221, 224 Spicules, 117, 118, 120, 183
Rosette-shaped, 222, 225 Spine, 9, 11, 18, 19, 24, 28–30, 33, 37, 58, 76, 84,
Round and flat, 132, 178, 179 117–122, 132, 140, 141, 149, 184, 208, 213, 219,
Round, flat, and discoidal, 175, 176 228
Round–oval–reniform, 181 Spinose, 184, 185
Rugulate, 29, 60 Spinose projections, 184, 185
R-unit, 219 Spiny ornamentation, 16–19
Spiral to serial, 177, 178
Spiraperturate, 53
S Spongy or microgranulous, 11, 18, 120
Sacculiform sac-like, 218 Spore morphology, 43
Salpingiform, 219 Sporoderm stratification, 44, 56, 57
Scabrate, 11, 29, 58 Spumellaria, 118, 119, 122
Scar, 11, 13, 15, 18, 44, 60, 147–149, 156, 159, 162, 164, Spur, 224, 227
165, 167, 168 Star-shaped, 222, 225
Sculpture, 9, 10, 18, 29, 31, 33, 34, 58, 79, 149–151, 184 Stellate, 104, 106, 222
Segment, 121, 122, 134, 207, 208, 224, 225 Stem, 219, 225, 249, 250
Segminate, 104, 107 Straight, 8, 17, 34, 45, 57, 104, 107, 148, 183, 184, 207,
Selvage, 145 210, 261
Semicircular cyclical, 178, 179 Striate, 60, 184, 185
Septa, 18, 84, 85, 215 Structural types, 216
Septate, 215 Structures closing central area, 218
Septum, 134, 215 Styliform, 219
Index 323

Sulculus/sulculi, 53 U
Sulcus, 52, 53, 74–77, 79, 81, 148 Ulcrus, 53
Surface patterns, 9–11, 28 Ultrastructure, 30, 63, 207, 261
Sutural ridge, 225 Umbilical, 178–181
Suture, 85, 171, 183, 184, 208, 211, 212, 225 Umbilical–extraumbilical, 178, 180, 181
Symmetry, 15, 34, 44, 54, 56, 57, 117, 118, 132, 133, Uniserial, 175, 177, 178
178, 193, 205, 207, 210, 219, 221 Upper/lower quadrant, 228
Syncolpate, 53
Synorate or svnclinorate, 54
V
Vacuoles (pusules), 74
T Valve closure, 149
Tabular, 84, 178, 179 Valve view, 132–134, 137–141
Tabulation, 77, 79–81, 87 Vaulted, 215, 216
Tangential, 117, 211, 212, 224 Velum, 134
Taper surfaces, 211 Ventral view, 80, 81, 137, 148, 156, 158, 159, 161, 164,
Terminal, 122, 138, 143, 145, 178, 180 165, 168
Termination, 31, 84, 227 Vermiculate, 11
Tertiopedate, 104, 106 Verrucate, 18, 29, 58
Test shape, 171, 175, 176 Vertical, 11, 16, 57, 132, 175, 178, 202, 211, 212
Tetraradiate, 221 Vesicle, 1, 3, 7–13, 15–19, 24, 27, 29, 30, 33, 34, 37
Theca (Amphiesma, Cell covering, or Cell wall), 77 Vestibulum (valve cavity), 145
Thickened, 15, 17, 34, 58, 183, 184, 261 V-unit, 219
Thorax, 120, 121
Tier, 208
Tile, 77, 183, 208 W
Tooth, 93, 181, 182, 210, 224 Wall, 1, 7, 9–11, 15–19, 27, 29, 30, 33, 34, 37, 43, 56, 57,
Torsion, 75 63, 74, 77, 81, 82, 84, 119, 132–134, 149, 171, 175,
Transapical axis, 132, 134 181–183, 205, 210, 213, 215, 228, 261–263, 265
Transverse, 34, 53, 74, 75, 110, 129, 202, 216, 218 Wall structure and test composition, 182
Triangular to trapezoid, 178, 179 wedge, 208, 210
Trichotomosulcate, 53 White matter, 94, 103, 110
Tricolpate/tricolporate/triporate, 53 Wide central area, 216
Triradiate, 52, 221 Wing, 33, 37, 122, 205, 224, 227
Triserial, 175, 177, 178
Trochospiral (trochoid, rotaliform), 175, 177, 178
Tube, 9, 16, 19, 121, 175, 185, 205, 211 Z
Tubercles, 11, 149, 169 Zeugoid rim, 212
Tubular, 9, 16, 17, 34, 175–177 Zonocolporate (=stephanocolporate), 54
T-unit, 219 Zonoporate (=stephanoporate), 54
Types of openings, 209, 210 Zygolith, 215