RECENT FORAMINIFERA

RECENT FORAMINIFERA

by

Esteban Boltovskoy & Ramil Wright

SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. 1976

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ISBN 978-90-481-8510-8 ISBN 978-94-017-2860-7 (eBook) DOI 10.1007/978-94-017-2860-7

© Springer Science+ Business Media Dordrecht 1976 Originally published by Dr. W. Junk b.v.- Publishers- The Hague in 1976

Cover design Charlotte van Zadelhoff

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Contents

Chapter contents Introduction to the second edition Introduction to the first edition

I. The systematic position and importance of the foraminifera II. The living organism

III. The test

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XV

1 5

51 IV. Benthonic foraminifera in normal marine waters 95 V. Benthonic foraminifera of brackish, hypersaline, ultrasaline, and

fresh waters 139 VI. Benthonic foraminifera of marshes, mangrove swamps and lagoons 151

VII. Planktonic foraminifera 159 VIII. Classification 197

IX. Ecology 223 X. Collection of material 275

XI. Preparation of samples 299 XII. Separation of the foraminifera from the sediment 315

XIII. Taxonomic determination 319 XIV. Faunal studies and their application 333 XV. Foraminiferal cultures 391

XVI. Storage of samples and specimens 401 XVII. Synonymy and illustration 407

XVIII. Unresolved problems 413 Bibliography 415 Index of genera and species 4 79 Index of authors 493 Index of subjects 505

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CHAPTER CONTENTS

I. The systematic position and importance of the foraminifera 1

II. The living organism 5 1. General and cytological description 5 2. Life cycle 21 3. Seasonal occurrence 33 4. Biological cycles 38 5. Parasites, epibionts and predators 39 6. Symbiosis and commensualism 44 7. Foraminifera and algae 47 8. Changes produced by human activity 48

III. The test 51 1. Form and size 52 2. Chambers and their position 54 3. Coiling direction 56 4. Chamber wall 57 5. Pores 67 6. Internal skeleton 71 7. Ornamentation 73 8. Aperture 75 9. Colour 79

10. Growth 80 11. Pathology and regeneration 85 12. Dwarfism 90

IV. Benthonic foraminifera in normal marine waters 95 1. General characteristics 95 2. Qualitative horizontal distribution 97 3. Qualitative vertical distribution 113 4. Quantitative horizontal distribution 122 5. Microdistribution 125 6. Quantitative vertical distribution 125 7. Foraminiferal abundance by weight, volume and surface area 130

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V. Benthonic foraminifera of brackish, hypersaline, ultrasaline and fresh waters 139

1. Brackish water 140 2. Hypersaline and ultrasaline water 14 7 3. Fresh water 148

VI Benthonic foraminifera of marshes, mangrove swamps and lagoons 151 1. Marshes 151 2. Lagoons 153

VII. Planktonic foraminifera 159 1. General characteristics 159 2. Horizontal qualitative distribution 166 3. Horizontal quantitative distribution 175 4. Vertical qualitative distribution (stratification) 186 5. Vertical quantitative distribution 189 6. Vertical migrations 192

VIII. Classification 197

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1. Zoological nomenclature 197 2. Biological significance of taxonomic categories 202 3. Development of the idea of a natural foraminiferal classifica-

tion 4. Conditional classification 5. Morphological traits and their taxonomic value 6. Problems of variability

IX. Ecology 1. Temperature 2. Salinity 3. Depth 4. Nutrition 5. Substrate 6. pH 7. Organic content of the substrate 8. Trace elements 9.

10. 11. 12. 13. 14. 15.

Turbidity Oxygen Illumination Calcium carbonate Currents and tides Other ecological factors Utility of ecological data

205 209 211 214

223 224 232 237 245 248 257 259 260 261 262 265 266 268 270 273

X. Collection of material 1. Fossil material 2. Recent benthonic foraminifera 3. Recent planktonic foraminifera 4. Preservation of material 5. Sample registration

XI. Preparation of samples 1. Bottom samples 2. Planktonic samples 3. Cytological techniques

XII. Separation of the foraminifera from the sediment

XIII. Taxonomic determination I . Difficulties 2. Process of determination 3. Establishment of a new species 4. Measurements 5. External test examination 6. Internal test examination 7. Wall mineralogy 8. Wall lamellae 9. Supplementary taxonomic traits

275 275 276 288 294 297

299 299 307 309

315

319 319 320 322 323 323 325 330 331 331

XIV. Faunal studies and their application 333 1. Qualitative and quantitative data and their display 333

2. Statistical and other mathematical manipulations 340

3. Paleoclimatic interpretations based on foraminiferal species distribution 348

4. Stratigraphic and climatic studies based on coiling direction and other morphological variations 358

5. Paleoclimatic studies based on oxygen isotope ratios 365

6. Relative sedimentation rates 368 7. Absolute sedimentation rates 370

8. Changes in sea level 371

9. Paleobathymetry 373 I 0. Foraminiferal number and bottom topography 374 II. Paleoclimates and planktonic test porosity 375 12. Trace elements and paleosalinity 375

13. Displaced faunas 376 14. Biologic and hydrologic indicators 377 15. Temperature-salinity-plankton diagrams 385

16. Stratigraphy and correlation 387

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XV. Foraminiferal cultures 391

1. Algal cultivation 392

2. Collection of living material 394

3. Separation and transport 395

4. Foraminiferal cultures 396

XVI. Storage of samples and specimens 401

1. Sample storage 401

2. Specimen storage 403

XVII. Synonymy and illustration 407 1. Synonymy 407 2. Illustration 410

XVIII. Unresolved problems 413 Bibliography 415 Index of genera and species 479 Index of authors 493 Index of subjects 505

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INTRODUCTION TO THE SECOND EDITION

The first edition of this book (Los Foraminlferos Recientes; biologia, metodos de

estudio, aplicacion oceanografica: EUDEBA, Buenos Aires, 510 pp, 114 text figs.)

was published in the Spanish language in 1965. The work was received warmly by

colleagues and reviewed favourably in specialized scientific journals in 9 different

countries. One consistent commentary both in reviews and in numerous letters from

colleagues was the desire to see a second edition of this book published in the

English language. This desire is now realized thanks to the kind suggestion of Junk Publishers.

However, because of the 13-year delay between preparation of the first and

second editions, it became necessary to update the book as well as translate it. The

bibliographical explosion in the sciences in the last decade made the renovation of this work a time-consuming task. To aid in this labour the original author invited his colleague, R. Wright, to participate in the revision. The work of bringing the text up

to date consisted of a revision of all chapters by evaluating and summarizing those data which have appeared in more than a thousand articles published since 1963-

1964. Several new sub-chapters have been added to describe new concepts and techniques developed since the first edition appeared. Many illustrations have been

revised and new ones added. This work was done jointly for which reason the book appears under our dual authorship. The translation of data from the first edition was

the responsibility of R. Wright. The revision of the text was completed in January, 1975, and information

coming to our attention after that date is not included in the book. The techniques described in this book for collecting samples, separating the fauna,

analysing the fauna, etc., are not always the most sophisticated available. We recognize that the development of new equipment has led to the development of

more refined and efficient techniques, but we wish to address the book to all students

of foraminifera at all levels, from the beginning student to the academic or industrial researcher. We also address the book to workers in all kinds of institutions, from the

industrial corporations, large universities and government laboratories with large

research budgets to those individuals working in organizations operating on modest funds. Consequently, we have tried to describe techniques and methodology appro­priate at all levels.

We have abandoned the practice followed in the first edition of saving space by excluding pagination of references in the bibliography. In the preparation of this

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edition we found it necessary to review references cited in the first edition. The abbreviated entries were .often less than helpful in locating a reference. Conse­quently, where possible, we have expanded the amount of information included in the bibliography.

Following the practice of the first edition we prefer to cite the generic and trivial name as written by the original authors. Fewer exceptions are made than in the first edition. This is a rather important point and requires some additional explanation.

There are two common ways in the current literature of handling this nomen­clatural problem. One is to cite the original name and to place after it, in parenthesis, the changed name which according to the writer is the correct one. A second way is to simply cite only the changed name, omitting the original designation. Neither of these methods is desirable. They both reflect the taxonomic taste of the writer but without the necessary ingredient of examination of the original specimens whose names are being changed. The first method also has the disadvantage of enlarging the text.

To better understand why we disagree with both of these methods we cite the following examples. In the recent past several hundred species ascribed to (or which would have been ascribed to) Bolivina were divided among Bolivina, Bolivinita, Bolivinitella, Brizalina, Coryphostoma, Sigmavirgulina and Suggrunda and there is no uniformity or agreement in the placement of these previous species of Bolivina in the other genera. We can not place the species which used to be (or would have been) in Bolivina into the genera now in use unless we can examine the specimens ourselves. In the case of trivial names the situation is yet worse as can be seen in the following example. Sagrina virgula, which was established by Brady (1884), is divided into three species, Rectobolivina virgula (Brady), Stilostomella bradyi (Cushman) and S. antillea (Cushman), according to Barker (1960). If someone cites Sagrina virgula (Brady) how can we decide to what species these specimens should be assigned in the modern classification without first examining the specimens?

Therefore, we are strongly convinced that, as a general rule, the original names should be maintained. Consultation of the species index, which contains the original author's name, should clarify any confusion.

However, as we said above, in this book some exceptions were made to this practice. They were made in the case of very old names not in use for several decades, and only in such cases when the change could not create even the most minimal of misunderstanding. Such a case is mentioned in the footnote on page 140. They are also encountered in the discussion of data from Ehrenberg, Rhumbler and some other authors, mainly from the last Century. There are only a few of these exceptions.

A number of persons helped make this book possible and we wish to acknowledge their contributions. To Haydee Lena we owe our greatest debt of gratitude for her immeasurable help in the preparation of the sections on cytology (Chapter 11,1 and XI,3). We wish to acknowledge A. Be, F. Haake, C. Hemleben, H. Lena, G. Lutze, R. Rottger, P. Thompson and K. Towe who graciously provided illustrative material for the book.

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For her help in drafting we thank Isabel Riob6 de Magaldi and in typing, Ana Marfa Leverone, Estela Orma and Luis Maria Mendez. We also wish to thank Cynthia Wright for long hours of editorial assistance.

Buenos Aires, February, 1976

E. Boltovskoy & R. Wright

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INTRODUCTION TO THE FIRST EDITION

The great development of study of the oceans resulted in the investigation of marine organisms. The importance placed on these investigations is reflected in the creation of numerous marine biological stations and the organization of various oceano­graphic expeditions by nations of both hemispheres.

The Protozoa, particularly the Foraminiferida, play a significant role in the organic world of the oceans. These typically marine organisms are numerous in both the benthos and the plankton.

Unfortunately, the bibliography in Spanish dedicated to these organisms is rela­tively small. There are works which analyse certain regions, faunas or specific problems, but there is no manual or text of a general character dedicated principally to the Recent forms. The work of Colom (1946) and the recently published work by Charlton de Rivero Palacio & Bermudez (1963) are excellent analyses of fossil foraminifera and well serve their geologic purpose. The summary of Douglass (1960) has the same goal. However, this last work is only available in mimeographed form as it was prepared for internal use in the University of Chile. Copies, therefore, are difficult to obtain. These circumstances stimulated me to write the present work.

Some words about the content and purpose of this book: 1. It is designed particularly for those interested in biological oceanography in general and in Recent foraminifera in particular. At the same time, it should prove of some value to micropaleontologists, since conclusions based on study of Recent species are com­monly of great interest to those who study fossil specimens. Moreover, there are various methods of investigation that are similar for both groups of foraminifera. Consequently, some of the pages of this book are devoted to methods of treating fossil foraminifera. 2. This book does not include a systematic section dealing with the description of the families and genera that comprise the Order Foraminiferida. Such descriptions are easily found in several manuals published in various foreign languages. I do not think that comprehension of systematics presents any great language difficulties since taxonomic descriptions use a limited number of often repeated terms.'

'In 1956 a special dictionary was published to help those who work with the foraminiferal bibliography in foreign languages: E. Boltovskoy, Diccionario foraminiferologico plurilingue (in 5 languages): English, Spanish, German, French and Russian (with alphabetic indicies): Ministerio de Marina, Servicio de Hidrografia Naval, Argentina, Pub!. Misc. No. 1001.

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3. The text contains many generic and specific names obtained from the works of many authors. In several cases more than one name has been employed for the same taxon. In those cases where the generic names are undoubtedly synonyms; e.g. Ammonia - Streblus - Rotalia, I have not considered it convenient to follow the authors choice since this might produce difficulties in understanding the text; I chose the most common name. When there were doubts about synonomy of generic names, I cited the taxon as the original author did. The specific names have been conserved in every case as originally cited. 4. A great deal of attention has been given to the practical side of foraminiferal study. It has become clear that recent university graduates possess a good under­standing of biological and protozoological theory, but are often at a loss to handle the more common tasks arising from the treatment of foraminiferal material. Because of this lack, this book contains detailed counsel on different procedures and techniques used by the foraminiferologist. Moreover, some advice is tendered which is not directly related to foraminiferal study but which may prove of great value to those working with these organisms; e.g. the construction of simple plankton nets and elementary devices for the extraction of bottom sediments, instructions on the cultivation of algae for the maintenance of foraminiferal cultures, etc. 5. The bibliography dedicated to the description of various methods used in the different stages of foraminiferal study is rather ample. There are numerous articles dealing with the procedures of sample disaggregation, test removal, etc. It is practically impossible to describe all the techniques available. I have therefore limited myself to describing only the most common techniques and ones which I have used in my laboratory. 6. When treating various themes, especially those poorly understood (ecology, distribution, etc.), I have described the various observations and opinions from the literature before presenting conclusions. In each case I have indicated the origin (author, date) of this information so that the reader may examine the original source to obtain a deeper understanding. 7. For reasons of space and because of the purpose of this book, some types of investigations, indirectly associated with foraminifera, have been omitted. Among these omissions are: the exposition of various zoological nomenclatural problems (e.g. types), advice on photomicrography, etc. Nevertheless, bibliographic sources of information on these omitted topics are always given. 8. When it became necessary to employ illustrative examples to best explain certain themes, I preferred to cite examples from South America. There are two reasons for this preference. Above all, this region is not well known from a foraminiferal perspective. In addition, this work is designed principally for those marine biologists who speak Spanish, the majority of whom reside in Central and South America. 9. All species names are cited in the text without reference to their authors. However, a taxonomic index is provided at the end of the book which contains this data. 10. The bibliolography requires two clarifications: (a) Unfortunately, there is no

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universally accepted system for the transliteration of names written in Cyrillic script. There is, therefore, confusion over the transcriptions of the Slavic authors. Some of

these authors may appear to be different persons because of different transcriptions of their names. To lend some uniformity to this situation I have adopted the system that was published in the Bulletin of Zoological Nomenclature, London. 2 The only exceptions are made for those Slavic authors whose names have appeared frequently in Latin characters based on a different transliteration system. (b) In order to save space, the bibliographic entries are abbreviated, without pagiTlation and illustration data.

This work has been in preparation for several years but the lack of some information has postponed its publication. In the year 1961, thanks to a subsidy granted by the Consejo Nacional de Investig~ciones Cientificas y Technicas (Argen­tina) I was able to visit various institutions in Europe and finish the manuscript. I consider it a very agreeable obligation to express my sincere appreciation to the Consejo. In addition, I most cordially thank Dr. R. Ruhstaller, Srtas. Haydee Lena and Anna-Maria Coronado and Sr. F. Theyer for their help in the preparation of the manuscript. I also wish to thank Sr. J. C. Riob6 for the drafting of the majority of the illustrations.

Buenos Aires, April, 1963

E. Boltovskoy

2 Almasov, A. & Boltovskoy, E. 1965. On the treatment of words written with cyrillic charac­ters, for the purposes of zoological nomenclature, bibliography, references, indicies, etc.: Bull. Zoo!. Nomencl., v. 11. no. 1.

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