Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1: Ponto-Caspian Foraminifera 3031123735, 9783031123733

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Valentina Yanko

Quaternary Foraminifera of the Caspian-Black SeaMediterranean Corridors: Volume 1 Ponto-Caspian Foraminifera

Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1

Valentina Yanko

Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1 Ponto-Caspian Foraminifera

Valentina Yanko Department of Physical, Marine Geology and Paleontology Odessa I. I. Mechnikov National University Odessa, Ukraine Avalon Institute of Applied Science Winnipeg, Manitoba, Canada

ISBN 978-3-031-12373-3 ISBN 978-3-031-12374-0 https://doi.org/10.1007/978-3-031-12374-0

(eBook)

# Springer Nature Switzerland AG 2022 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. Cover illustration: Map in the middle - The Ponto-Caspian- Mediterranean Corridor Around the map foraminiferal species: 1. Canalifera parkerae Yanko, 1974, 2. Ammonia ammoniformis (d’Orbigny), 1826, 3. Ammobaculites ponticus Mikhalevich, 1968, 4. Bolivina pseudoplicata Heron-Allen et Earland, 1930, 5. Elphidium ponticum Dolgopolskaja and Pauli, 1931, 6. Haynesina anglica (Murray), 1965, 7. Bulimina aculeata d’Orbigny, 1826, 8. Eggerelloides scaber (Williamson, 1858) This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Foreword

The region known as the Ponto-Caspian includes the Black Sea, the Sea of Azov, the Caspian Sea, and the Aral Sea and connecting straits. While this region has been well studied and described in the Russian and Ukrainian literature, the details are much less widely known by scientists without access to or knowledge of those works. Valentina Yanko has produced a synthesis of a vast array of research, including her own and that of her collaborators over the past 50 years. While this work focuses on the Pleistocene through Recent foraminiferal assemblages, the implications extend far beyond to interpretation of basin closures throughout geologic history. And such basin closures include some of the major hydrocarbon sources on Earth, including those of the Ponto-Caspian region. Many lessons from geologic history appear in this extensive and detailed description of the repeated cycles of marine flooding and freshening across a series of related, sometimes interconnected, marginal seas. The pioneering micropaleontologists in the region that began in the last decades of the nineteenth century are mentioned in Chap. 1. More importantly, detailed investigations and descriptions of foraminiferal species and assemblages exploded in the mid-twentieth century. With the author’s career beginning in the mid-1970s, she was incredibly productive as a member of the second generation of mid-late twentieth century researchers who were not simply describing the assemblages but also applying their findings to understanding the geologic history of this series of basins. When glaciation reduced sea levels worldwide, basins freshened and oligohaline species prevailed. Subsequent transgressions allowed marineinfluenced taxa to migrate stepwise through the basins. The series of basins are described in Chap. 2, including their geographic locations as well as bathymetric and hydrographic features, while touching upon their geologic histories. The environmental tragedy of the Aral Sea is lamented. Formerly the fourth largest lake in the world, the Aral Sea has largely dried up, eliminating a once prosperous fishing industry resulting in economic hardship and human suffering. In the second part of Chap. 2, the author describes the sources of the more than 30 thousand sediment samples that were summarized and evaluated in subsequent chapters. The numerous organizations that supported collections and the many scientists whose works are utilized in the synthesis are recognized and documented. Samples were collected from grab samples, gravity cores, piston cores, multi cores, box cores, boreholes, and outcrops, including Quaternary stratotypes. Sites and habitats sampled included limans, lagoons, river deltas, shelves, and continental slopes. Analyses included lithological as well as foraminiferal analyses, and often analyses of other shelled biota, especially mollusks. The discussion of foraminiferal taxonomy in Chap. 3 elaborates on the theme that is absolutely central to the overall work. Taxonomy is a specialty in which Prof. Yanko has superb expertise that she has honed throughout her career. As she emphasizes at the beginning of the chapter, precise taxonomic identification is absolutely essential to high-resolution stratigraphy and paleoenvironmental analyses. And understanding the basic characteristics and biological limitations inherent to higher taxa, specifically the major orders, provides vast insight into where lower taxa originated, and why they were able to proliferate, or were unsuited to thrive, across the euryhaline habitats of these marginal seas. v

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Prof. Yanko emphasizes that recognition and consistent understanding of the characteristics of each species at all stages of ontogeny are essential to precise identification. She reminds the reader that a species is composed of many populations that can differ from each other to some degree as a result of founder effects or adaptations to local conditions. If individuals from these populations differ consistently in diagnostic morphological characteristics and are geographically isolated, they are considered as subspecies. Given the frequent isolation of the individual basins through the Pleistocene and Holocene, local adaptations were inevitable, allowing differences to emerge that are recognized as subspecies. This extensive analysis on the relatively recent geologic history of this series of basins provides insight into both the widely recognized process of allopatric speciation and somewhat lesser-known process of reticulate evolution, whereby taxa separated for millennia can diverge somewhat, but when populations are reconnected, new assortments and adaptations can emerge. Thus, reconnection of the basins has allowed subsequent mixing of those subspecies and occurrences in environmental conditions to which they have adapted. For example, salinity can be incredibly variable locally within limans, deltas, and lagoons, on seasonal and interannual time scales, or even daily or weekly cycles depending upon weather. Prof. Yanko recognizes 173 lower taxa (i.e., species, subspecies, some of them in open nomenclature) of Quaternary benthic foraminifera across the Ponto-Caspian basins, providing an excellent review of the principles of taxonomic classification of the orders that occur there. With the essentials of foraminiferal taxonomy established, in Chap. 4 the modern basins are described in regional detail. For example, the Black Sea study sites are subdivided into 14 regions, with environmental conditions described and the lower taxa identified for each. This chapter provides detailed lessons in regional geography, as well as both regional and sometimes obscure terminology. The reader will want to keep bookmarks on location maps and detailed facies, bathymetric and salinity definitions. The reader should also keep in mind that this chapter describes something like 70 years of research on thousands of samples in places that many readers may be unfamiliar. Don’t be discouraged, the details are here for historical reference and to define and support syntheses to come both later in the chapter and in the volume. The fossil assemblages of the Pleistocene and Holocene are presented and interpreted in substantial detail in Chap. 5. The chapter begins by discussing the diversity of stratigraphic schemes used by researchers who have carried out investigations in the region. Prof. Yanko provides the plethora of terminology essential to understand the regional geology and interpretations of the glacial-interglacial (i.e., regressive/ transgressive) cycles of the PontoCaspian Corridor. Biostratigraphic diagnoses of fossil foraminiferal assemblages were based on the appearance of new species/subspecies that characterized an assemblage. Eco-stratigraphy and paleo-events are recognized lithologically and from changes in dominance of foraminiferal, Molluscan, and other fossil taxa, in particular, with their relationships to salinity. This chapter represents a synthesis of multidisciplinary studies of an enormous amount of geological material and, as such, should be a treasure for anyone interested in the region or in the level of detail available from a synthesis of data on a series of interconnected basins during times of sea-level fluctuations. Another potential readership should be anthropologists and archaeologists studying this important corridor of migrations of hominid populations. The paleogeographic origins of the Pleistocene foraminiferal taxa of the Ponto-Caspian basins, and an exploration of why certain taxa were able to thrive in the individual basins in response to opening and closing of corridors, are explored in Chap. 6. Among the species and subspecies, about 40% are calcareous members of the order Rotaliida, nearly a quarter are Miliolida, and about 20% are Lagenida. The other roughly 15% include calcareous Buliminida and members of three agglutinated orders. The foraminiferal species and subspecies found in the Caspian are overwhelmingly endemics (88%), with a few cosmopolitan species that are known to inhabit brackish environments. More than half the species and subspecies found in Caspian are agglutinated. Thus, the Caspian fauna represent descendants of Tethyan taxa that

Foreword

Foreword

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have diverged in isolation. That abundance of endemic species could provide outstanding material for molecular-genetic and morphological comparisons of the endemic species with their likely closest Tethyan relatives with outstanding potential for addressing both rates and degrees of divergence. This is just one small example of the wealth of future studies for which the amazing details documented in this book could provide impetus. In contrast, the species and subspecies found in the Black Sea region (Pont) are predominantly Mediterranean immigrants (83%), with endemics and Caspian immigrants accounting for most of the remainder at less than 10% each. Prof. Yanko documents eight major waves of foraminiferal migration into the Black Sea from the Mediterranean, corresponding to major glacio-eustatic sea-level transgression events. The fourth wave brought 52 lower taxa, many of which are not still found, likely indicating higher salinities than presently occur in the Black Sea. A distinctive feature of the faunas is the predominance of the Ammoniidae-Elphidiidae shelf genera. All Pontic foraminifera are either shallow-dwelling or descendants of shallow-dwelling taxa, reflecting the shallow nature of the straights that connected the Black Sea with the Mediterranean. The influence of anthropogenic pollution—freshwater input that results in low and variable salinities and natural methane emissions— is summarized in Chap. 7, providing examples of applications of meiobenthos, especially foraminifera, to studies of both naturally occurring stresses and those associated with human activities. This chapter uses results from three sets of cruises that sampled more than 240 stations on the northwestern shelf of the Black Sea. A major advantage of studying foraminifera is that their tests are commonly preserved in sediment. And, in studies of environmental stresses, morphological anomalies are commonly seen in the tests of the foraminifera that lived and grew under stressful conditions. Pyritization of tests is another stress indicator. Ammonia tepida, A. compacta, A. ammoniformis, Canalifera parkerae, and Porosononion martkobi ponticus are taxa that can survive in polluted or otherwise stressed environments on the Black Sea shelf, and their tests most frequently exhibit morphological anomalies or pyritization. Chapter 8 represents a volume on its own! Taxonomic descriptions are provided for a vast majority of the genera, species, and subspecies encountered in the region. Morphological descriptions include scanning electron micrographs, statements regarding variability, and comparisons with similar taxa. In most cases, ecological settings and known distributions in the region are also provided. Thus, this chapter will provide a treasure trove of information for others studying modern and Quaternary-fossil foraminifera in this region. This comprehensive presentation and synthesis of the foraminiferal faunas of the PontoCaspian region is encyclopedic. The book will be an essential resource for anyone working on the foraminifera, meiobenthos, ecology, paleoecology, biostratigraphy, and Quaternary history of this region. Ultimately, even more valuable may be the potential insights and future studies that readers of this work, or of individual chapters, may envision within the region or far from it, either geographically or in geologic time. It is my great pleasure to introduce this book. I am in awe of the detailed information from thousands of samples from her own work and that of many other researchers. I am even more impressed by the syntheses and ecostratigraphic interpretations, and the potential for this work to inspire a diverse array of future investigations, ranging from molecular and morphological evolutionary studies of the development of endemic species, to detailed investigation of closing basins in the Permo-Carboniferous, to many other possibilities that are far beyond my imagination. College of Marine Science University of South Florida St Petersburg, FL, USA e-mail: [email protected]

Pamela Hallock

Preface

In these prefatory paragraphs, the author presents essential information that includes the background for the creation of this book, the process by which it came to be, and appreciative words for those who made this publication possible. The study area known as the Ponto-Caspian includes the Black Sea, the Sea of Azov, the Caspian Sea, the Aral Sea (currently dried up), and connecting straits. The Black Sea, with a maximum depth of 2212 m, is the easternmost of the seas of the Atlantic Ocean basin and the most isolated sea of the modern Global Ocean. The Sea of Azov, with depth up to 14 m, is connected to the Black Sea via the Kerch Strait, which is 45 km long, 4.5 km wide, and up to 6 m deep. The shallowness of the strait results in limited water exchange between the two basins. The Bosporus Strait meanders along its 35 km in length and 0.7–3.5 km in width. It is 35.8 m deep, on average, with a few elongate potholes (about 110 m in depth each) on the bottom. Carrying 5–10 times more water flow than the Kerch Strait, waterflow within the strait is stratified and bidirectional, resembling salt-wedge estuarine circulation. Limited transport of low density, lower-salinity (average 18 psu) water from the Black Sea flows southward; while the dominant transport is subsurface, flowing northward, carrying more saline (average 38 psu), denser seawater from the Sea of Marmara into the Black Sea. The Black Sea exhibits the standard oceanic provinces of continental shelf, slope, and abyssal plain. The extensive continental shelf accounts for 25% of the sea area; the isobath 200 m is commonly taken as the shelf boundary. The sea was formed at the end of the Mesozoic as a back-arc basin. The details and taphonomic conditions of the Black Sea are unusual. It is the world’s largest anoxic (oxygen-free) marine basin. Its strongly stratified water column possesses (1) a thin, well-oxygenated surface layer (20–30 m) with low salinity and warm temperatures; (2) a low-oxygen (suboxic) transition layer (30–150 m); and (3) a thick bottom layer of colder, denser, and more saline water lacking oxygen but high in sulfides. The Caspian Sea is completely isolated from the World Ocean and thus is technically an endorheic lake. It is divided into three distinct physical regions: the Northern, Middle, and Southern Caspian. The Northern Caspian only includes the Caspian shelf and is very shallow, with an average depth of only 5–6 m; it accounts for less than 1% of the total water volume. The Middle and Southern Caspian account for 33% and 66% of the total water volume, respectively. The Aral Sea was also an endorheic lake; lying between Kazakhstan and Uzbekistan, it began shrinking in the 1960s and had largely dried up by the 2010s. UNESCO added the historical documents concerning the collapse of the Aral Sea to its Memory of the World Register as a unique resource to study this “environmental tragedy.” The pioneering micropaleontologists in the region began their work in the last decades of the nineteenth century. Detailed investigations and descriptions of foraminiferal species and assemblages exploded in the mid-twentieth century. Since the mid-1970s, the author of this book has extensively studied taxonomy, ecology, paleoecology, paleogeography, and biostratigraphy of the Ponto-Caspian region, specializing in benthic foraminifera. This book contains an introduction, eight chapters, a conclusions section, and an extensive reference list for each chapter, many of which were published in regional languages, and as such are not well known in the west. The Ponto-Caspian Quaternary benthic foraminifera are ix

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Preface

represented by 7 orders, 35 families, 66 genera, 173 species (some in open nomenclature), and subspecies (i.e., lower taxa). Of the 173, 123 lower taxa inhabit the Black and Caspian Seas today. This in-depth study of benthic foraminifera provides invaluable data on taxonomy of benthic foraminifera, sea level, and salinity changes as well as pollution monitoring and contouring methane reservoirs under the sea bottom. The 107 figures present regional maps, data analyses, and other essential information. The 36 plates that present SEM images of foraminifera include images to illustrate descriptions of all species and subspecies of foraminifera found by the author in the Ponto-Caspian region. The coverage encompasses both terrestrial and underwater areas, and a broad approach ranging from geological subjects to environmental applications. Odessa, Ukraine

Valentina Yanko

Introduction

This monograph is written on the basis of the long-lasting study of the late Quaternary benthic foraminifera of the Ponto-Caspian region (e.g., Yanko 1974, 1989, 1990; Yanko-Hombach 2007; Yanko and Kondariuk 2020). The region is defined here as the large geographical area covering (from west to east) the Black Sea, the Sea of Azov, the ancient Manych Outlet, the Caspian Sea, the Aral Sea, their coasts, and connecting straits. This Pionto-Caspian is of a strategic importance not only for the Caspian (Republic of Azerbaijan, Russia, Iran, Turkmenistan, Kazakhstan) and Black Sea (Bulgaria, Romania, Ukraine, Russia, Georgia, Turkey) countries but also for at least 17 other countries, which share a drainage basin of the Ponto-Caspian Region that is one-third the size of the European continent. The Ponto-Caspian region acts as a paleoenvironmental amplifier and as a sensitive recorder for climatic events where variations in sea level, hydrological regime, and coastline migration are especially pronounced due to its geographical location and semi-isolation from the open ocean (e.g., Fedorov 1978; Yanko 1990). It also provides a linkage between the marine and continental realms. This region is among the basins, which have been cited in literature as having conditions suitable for natural gas hydrate reserves that are increasingly being considered as a potential energy resource (Kvenvolden 1993). Lately, this region has spurred a tremendous international interest as a possible place where the biblical story of the Great Flood (Ryan et al. 1997, 2003; Chepalyga 2007) originated, encouraging a new round of controversial research on paleoenvironmental reconstructions, e.g., the hydrological regime in connecting straits, transition from a lacustrine to a marine environment, an influence of the Black Sea outflow on deposition of the Eastern Mediterranean sapropels as well as past/present/ future adaptation of humans to environmental change (Yanko-Hombach 2007; YankoHombach 2007; Yanko-Hombach et al. 2014). During the Late Quaternary, the Black Sea was repeatedly isolated from the Mediterranean Sea by sea-level fluctuations. Geographical location and periodic connection of the Black Sea either with the Mediterranean or Caspian seas predetermined specific hydrogeological regimes in the basin, making it an excellent paleoenvironmental amplifier and a sensitive recorder of climatic events All this makes the Ponto-Caspian region and its foraminiferal assemblages unique, both from the modern and fossil perspectives, and also in the context of its tectonic history and sea-level changes in isolated and semi isolated from the World Ocean basins. Because these basins are quite different from more oceanic regions, the author is dealing with more subtle distinctions with respect to salinities and estuarine conditions, and therefore at the end of the volume, there is a glossary of terms that includes the Black Sea, the Sea of Azov, the Caspian Sea, Aral Sea (currently dried up), and connecting straits. The potential of benthic foraminifera for paleoenvironmental reconstructions is well known. These hard-shelled microorganisms have tremendous taxonomic diversity enabling a wide range of biological reactions to varied environmental factors, including many species-specific responses to ecological conditions (Fursenko 1978), which adds to their potential as index species for monitoring sea-level and salinity changes. They have very short reproductive cycles—6 months to 1 year (Boltovskoy 1964)—and rapid growth (Walton 1964) that makes even their community structure particularly responsive to environmental change. Their tests are xi

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readily preserved in the sediments and can record evidence of environmental variability through time. They are small and abundant compared to other larger, hard-shelled taxa (such as mollusks), making them particularly easy to recover in statistically significant numbers (Yanko et al. 1999). The main goal of the work is taxonomy and ecology of the Late Quaternary benthic foraminifera with hard tests and their application for biostratigraphy, environmental reconstructions, and paleogeography of the Ponto-Caspian region. To reach the goal the following objectives have been set: (1) detailed description of recent and Late Quaternary benthic foraminifera from bottom sediments and coastal outcrops of the Ponto-Caspian region in order to establish their taxonomic classification; (2) determination of the ecological preferences for all lower taxa in order to supplement the classification with ecological/paleoecological information; (3) determination of biofacial types among the Late Quaternary foraminiferal assemblages as well as their migration in space and time; (4) based on objectives 1–3 high-resolution paleoenvironmental reconstructions of the Ponto-Caspian region supplemented whenever possible by isotopic and OSL records.

References Boltovskoy E (1964) Seasonal occurrences of some living foraminifera in Puerto Deseado’ (Patagonia, Argentina). ICES J Mar Sci 29(2):136–145 Chepalyga AL (2007) The late glacial great flood in the Ponto-Caspian basin. In: Yanko-Hombach V, Gilbert AS, Panin N, Dolukhanov PM (eds) The Black Sea flood question: changes in coastline, climate and human settlement. Dordrecht, Springer, pp 119–148 Fedorov PV (1978) Pleistotsen Ponto-Kaspiya (The Pleistocene of the Ponto- Caspian). Moscow, Nauka (in Russian) Fursenko AV (1978) Vvedenie v izuchenie foraminifer (Introduction to the study of Foraminifera). Trudy Instituta Geologii i Geofiziki 391. Novosibirsk, Nauka, 242 pp (in Russian) Kvenvolden KA (1993) Gas hydrates-geological perspective and global change. Rev Geophys 31(2):173–187 Ryan WBF, Pitman WC III, Major CO, Shimkus K, Maskalenko V, Jones GA, Dimitrov P, Görür N, Sak{nç M, Yüce H (1997) An abrupt drowning of the Black Sea shelf. Mar Geol 138:119–126 Ryan WBF, Major CO, Lericolais G, Goldstein SL (2003) Catastrophic flooding of the Black Sea. Ann Rev Earth Planet Sci 31:525–554 Walton WR (1964) Recent foraminiferal ecology and paleoecology. In: Imbrie J, Newell ND (eds) Approaches to paleoecology, pp. 151–237 Yanko V (1974) Novye vidy bentosnykh foraminifer iz golode-novykh otlozheniy Severo-8apadnogo shelfa Chernogo morya (New species of benthic foraminifera from Holocene sediments of the north-western shelf of the Black Sea). Paleontologicheskiy Sbornik 11:24–30 (in Russian) Yanko V (1989) Quaternary Foraminifera of the Ponto-Caspian Region (the Black Sea, the Sea of Azov, the Caspian Sea and the Aral Sea): Taxonomy, biostratigraphy, history, ecology. Doctoral thesis. Moscow State University, two volumes, 1000 pp (in Russian) Yanko V (1990) Stratigraphy and paleogeography of marine Pleistocene and Holocene deposits of the southern seas of the USSR. Memorie della Società Geologica Italiana 44:167–187 Yanko V, Kondariuk T (2020) Origin and taxonomy of the Neopleistocene-Holocene Ponto-Caspian benthic foraminifera. Geologichnyy zhurnal 1:17–33 Yanko V, Arnold A, Parker W (1999) The effect of marine pollution on benthic foraminifera. In: Sen Gupta BK (ed) Modern Foraminifera. Dordrecht, Kluwer Academic, The Netherlands, pp 217–238 Yanko-Hombach VV (2007) Controversy over Noah’s flood in the Black Sea: geological and foraminiferal evidence from the shelf. In: Yanko-Hombach V, Gilbert AS, Panin N, Dolukhanov PM (eds) The Black Sea flood question: changes in coastline, climate and human settlement. Dordrecht, Springer, pp 149–204 Yanko-Hombach V, Mudie PJ, Kadurin S, Larchenkov E (2014) Holocene marine transgression in the Black Sea: new evidence from the northwestern Black Sea shelf. Quat Int 345:100–118

References

Acknowledgments

The author has studied the Quaternary Ponto-Caspian foraminifera for nearly 50 years, and the results of her research are presented in a significant number of monographs and articles, many of which are published in Russian or Ukrainian. The information presented in this book represents the results of research conducted not only by the author but also by many geologists from an array of scientific and applied geological organizations who collected sediment from throughout the region. Samples for foraminiferal analysis were obtained from multiple organizations (e.g., Siberian Branch of the USSR Academy of Sciences; Southern Branch of the Institute of Oceanology, USSR Academy of Sciences; Institute of Oceanology of the Bulgarian Academy of Sciences; Yuzhmorgeologiya, GeoEcoMar; Odessa I.I. Mechnikov National University; Prichernomor DGRP, and some others with which the author cooperated on the contractual and/or scientific basis). The contributions of individual experts are acknowledged in the text. Here, the author expresses deep gratitude to A.K. Bogdanowicz, V.A. Krasheninnikov, N.N. Subbotina, and A.V. Fursenko, who taught the author the basics of micropaleontology at the beginning of her carrier. Prof. I.Ya. Yatsko instilled a love for foraminifera, which the author carried through her whole life and to whom she is especially grateful. All transliterations of cited sources published in languages using the Cyrillic alphabet comply with the requirements of international standards for bibliographic references according to the US Library of Congress (https://www.loc.gov/catdir/cpso/romanization/russian.pdf). Exceptions are the names of authors, which we have left in their own preferred transliterations, as well as geographical names as presented most commonly in the majority of English papers. The author also is deeply grateful to the following individuals: Prof. Pamela Hallock from University of South Florida, USA, for editing the English text, providing extremely valuable comments, and composing the Foreword for this book; Prof. Ronald Martin, Delaware University, USA, and Prof. Lyudmila Vorobyeva, the Institute of Marine Biology of the National Academy of Science of Ukraine, for review of the book and their very useful comments; Dr. Irena Motnenko (the author’s daughter) for drawing of the inner structure of foraminiferal tests (Chap. 3) and some other pictures; Dr. Revinder Sidhu, Microscopy and Materials Characterization Facility Manager (Manitoba Institute for Materials, University of Manitoba, Canada) for her help in imaging of microfauna by SEM (Paleontological plates 1–36); and the managing team at Springer, especially Mr. Solomon George, Project Co-ordinator (Books) for Springer Nature, for his guidance and patience in awaiting delivery of the finished manuscript. This book is a contribution to IGCP 521 “Black Sea-Mediterranean Corridor during the last 30 ky: sea-level change and human adaptation”; INQUA 0501 “Caspian-Black Sea-Mediterranean Corridor during the last 30 ka: sea-level change and human adaptive strategies”; IGCP 610 “From the Caspian to Mediterranean: environmental change and human response during the quaternary”; the Russian–Ukrainian project No. Φ28/428-2009 “The Northwestern Black Sea region and global climate change: environmental evolution during the last 20 ka and forecast for the 21st century” sponsored by the State Fund for Fundamental Research, Ukraine; “Study the interaction between nature and human society in the north-western Black Sea coast during the late Pleistocene and Holocene,” “To study the xiii

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processes of methane formation and spatial distribution of methane in the Black Sea and to theoretically justify its influence on eco- and geosystems of the basin,” “Development of forecast criteria for searching for hydrocarbons in the Black Sea on the basis of the theory of fluidogenesis”; all four sponsored by the Ministry of Education and Science of Ukraine; COST Action TD0902 SPLASHCOS project “Submerged prehistoric archaeology and landscapes of the continental shelf ”; EU-FP6 project HERMES “Hotspot ecosystems research on the margins of European seas”; EU BLACK SEA ERA.NET-WAPCOAST project “Water pollution prevention options for coastal zones and tourist areas: Application to Danube Delta front area”; INCO-COPERNICUS “Pollution by oil and herbicide of the Black Sea: Novel technologies of detection and biological impact”; EC (AVICENNE Program, AVI CT920007).

Acknowledgments

Abstract

This volume contains a heretofore unavailable compilation of detailed information on the Ponto- Caspian foraminifera. The region as a whole consists of the remnants of the Tethys and Parathethys seaways, totaling in area a drainage basin one-third the size of the entire European continent, and is of strategic importance to the surrounding countries. Foraminifera are highly reliable paleoenvironmental indicators, ubiquitous in marine environments, and taxonomically diverse, which give them the potential for a wide range of biological responses to varied environmental factors. Their tests are readily preserved and can record evidence of environmental change through time, thus providing historical baseline data even in the absence of background studies. The book includes taxonomic descriptions for 152 species and subspecies (i.e., lower taxa) from the Black Sea, Sea of Azov, Caspian Sea, and Aral Sea. For the majority of them, SEM images and descriptions, as well as data on ecology, paleoecology, distribution, location, and material, are provided. The book will be useful to specialists in the Quaternary history of the Caspian-Black Sea-Mediterranean Corridors as well as those in environmental monitoring and risk assessment. It can be used by students studying marine geology and paleontology to serve as a framework for future investigations of the paleoclimatic history of the Ponto-Caspian region.

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Contents

1

State-of-the-Art . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.1 Foraminifera of the Black Sea and Sea of Azov . . . . . . . . . . . . . . . . . . . . 1.2 Caspian and Aral Seas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3 Our Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1 1 3 4 4

2

Study Area, Material, and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1 Study Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Statistical Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.4 Stratigraphic Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

9 10 13 17 18 18

3

Taxonomic Classification of Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Principles of Taxonomic Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 Agglutinated Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 Taxonomic Classification of the Order Miliolida . . . . . . . . . . . . . . . . . . . . 3.5 Taxonomic Classification of Order Lagenida . . . . . . . . . . . . . . . . . . . . . . 3.6 Taxonomic Classification of Order Rotaliida . . . . . . . . . . . . . . . . . . . . . . . 3.6.1 Test Morphology and Accepted Terminology of Rotaliids . . . . . . . 3.6.2 Systematics and Genetic Interrelations . . . . . . . . . . . . . . . . . . . . . . 3.7 Taxonomic Classification of Order Buliminida . . . . . . . . . . . . . . . . . . . . . 3.8 Systematics and Comparison with Other Basins . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

21 21 22 29 30 31 32 33 35 36 36 38

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Modern Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2 Modern Foraminifera of the Black Sea and the Sea of Azov . . . . . . . . . . . 4.2.1 Northwestern Deltas, Limans, and Lagoons. . . . . . . . . . . . . . . . . . 4.2.2 Northwestern (Including Western Crimean) Shelf . . . . . . . . . . . . . . 4.2.3 Eastern Crimean Shelf, Kerch Strait, and Sea of Azov . . . . . . . . . . 4.2.4 Caucasian Shelf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2.5 Bulgarian, Southwestern, and Bosphorus Outlet Shelf . . . . . . . . . . 4.3 Modern Foraminifera of the Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.1 North Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.2 Middle Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.3 South Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.4 Bays and Straits of the Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . 4.4 Modern Foraminifera of the Aral Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.6 Lateral Rows of Modern Foraminiferal Assemblages . . . . . . . . . . . . . . . . . 4.7 Quantitative Distribution and Ecology of Benthic Foraminifera . . . . . . . . .

41 42 42 43 47 54 59 59 64 64 66 69 71 74 75 77 80 xvii

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4.8

Bionomic Groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 4.8.1 Black Sea and Sea of Azov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 4.8.2 Caspian Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 4.8.3 Aral Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 4.9 Ecological Classification of Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . 97 4.10 Use of Modern Foraminifers for Bio- and Ecostratigraphy and Facies Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 5

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Quaternary Bio- and Ecostratigraphy, with Elements of Paleogeography of the Ponto-Caspian Corridors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2 Caspian Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.1 Eopleistocene (Apsheronian Regional Stage) . . . . . . . . . . . . . . . . . 5.2.2 Lower Neopleistocene (Bakinian Stage) . . . . . . . . . . . . . . . . . . . . 5.2.3 Middle Neopleistocene (Urundzhikian and Gyurgyanian Stages) . . . 5.2.4 Upper Neopleistocene (Khazarian and Khvalynian Stages) . . . . . . . 5.2.5 Holocene (Novocaspian Stage) . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3 Black Sea Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.1 Eopleistocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.2 Lower Neopleistocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.3 Middle Neopleistocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.4 Upper Neopleistocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.5 Holocene (ca. 10 ky BP–Present) . . . . . . . . . . . . . . . . . . . . . . . . . 5.4 Correlation of Stratigraphic Units . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.5 Pleistocene Intrusions from the Mediterranean and Caspian Seas into the Black Sea: Reconstructions Based on Foraminifera . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Origin and Taxonomy of the Pleistocene Ponto-Caspian Benthic Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2 Caspian Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3 Black Sea Region (Pont) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Benthic Foraminifera as Indicators of Environmental Change in the Black Sea in Space and Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2 Foraminifera as Indicators of Environmental Stress Caused by Herbicides and Oil Pollution (INCO-COPERNICUS Case Study) . . . . . . . . . . . . . . . . 7.3 Foraminifera as Indicators of Environmental Stress Caused by River Discharge (WAPCOAST Case Study) . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4 Foraminifera as Indicators of Environmental Stress by Methane Emissions (HERMES Case Study) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.5 Foraminifera as Indicators of Environmental Stress on the Neopleistocene-Holocene Boundary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.6 Morphological Deformities of Foraminiferal Tests as Indicators of Environmental Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

111 112 116 116 117 121 124 128 128 130 131 131 139 144 167 175 178 180 189 189 190 190 206 206 209 209 212 213 214 215 217 220

Systematic Part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325

Contents

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Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331 Appendix 1: Alphabetical Index to Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331 Appendix 2: Addresses of Studied Outcrops on the Black Sea Coast . . . . . . . . . . 332 Paleontological Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337 Alphabetical Index to Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409 Alphabetical Index to Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415

About the Author

Photo taken in September, 2019 during field work on the Black Sea coast

Valentina Yanko Doctor of Geological and Mineralogical Sciences; Professor, Head of the Department of Physical, Marine Geology and Paleontology; Director of the Scientific and Educational Center for Geoarchaeology, Marine and Ecological Geology of the Odessa I.I. Mechnikov National University; and President of the Avalon Institute of Applied Sciences, Canada. She graduated from Odessa I.I. Mechnikov National University (former Odessa I.I. Mechnikov State University). Her main scientific interests include: geology, specialty marine geology; paleoecology and ecology; paleontology, specialty paleontology of invertebrates; ecological micropaleontology; foraminifera of intercontinental basins (Mediterranean Sea, Sea of Marmara, Black Sea, Sea of Azov, Caspian Sea, Aral Sea); foraminifera as indicators of natural and anthropogenic stress; ecotoxicology of foraminifera and their chemical defense mechanisms; marine Quaternary geology and paleoceanography; paleoclimatology; global sea-level changes; Quaternary history, paleoceanography, glacial and intraglacial epochs.

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State-of-the-Art

Abstract

The history of research on Ponto-Caspian Quaternary foraminifera at the end of the nineteenth century to the present is introduced. The earliest report from the Crimean region was by Pereialsvtseva (Novorossiysk.ob-va yestestvoisp 10(2):79–114 (in Russian), 1886), who noted nine species. At the very beginning of the twentieth century, Zernov (Yezhegodnik zool. muzeya AN, Sankt Petresburg 13(4):28–35 (in Russian), 1906) wrote that foraminifera are scarce in the Black Sea. In contrast, Dolgoplskaya and Pauli (Trudy Karadagskoy nauchnoy stantsii imeni T.I. Vyazemskogo 4:23–48 (in Russian), 1931) found foraminifera to be abundant and contribute a significant proportion of recent benthic assemblages. Research carried out in the Black Sea, the Kerch Strait, and the Sea of Azov, as well as near the Bosphorus outlet, was conducted by researchers from the USSR, Bulgaria, Romania, and Georgia. The first data on Caspian foraminifera were presented by Ehrenberg in 1873, who reported 23 species. The earliest observations from the Aral Sea found only two species. Intensive study of taxonomy, quantitative distribution, and ecology of live (Rose Bengal stained) foraminifera in the Caspian and Aral Seas and their comparison with recent foraminifera from other basins was carried out by Mayer (Vertikal’noye raspredeleniye foraminifer v Kaspiyskom more. In: Maev YeG (ed) Kompleksnye issledovaniya Kaspiyskogo morya 6, pp 101–107 (in Russian), 1974a, Foraminifery Kaspiyskogo i Aral’skogo morey (Foraminifera of the Caspian and Aral seas). Avtoref. Dis. kand. biol. Nauk. Moscow State University, 24 pp (in Russian), 1979b). In addition to, and often in collaboration with, a variety of other researchers working in the region, the author has carried out more than 50 years of research on the taxonomy, ecology/paleoecology, paleogeography, and biostratigraphy of Ponto-Caspian benthic foraminifera. This chapter introduces extensive exploration of the northwestern Black Sea, along the Ukrainian, Bulgarian, and

Caucasian shelves, as well as the northern exit of the Bosphorus Strait, the Sea of Azov, the Caspian Sea, and the Mediterranean Sea. Fossil assemblages from numerous Pleistocene stratotypes were studied in the coastal zone of the Ponto-Caspian region. The application of benthic foraminifera to reconstruction of sea-level change and coastline migration in regard to the Great Flood Hypotheses in the Black Sea was also investigated. Most recently, the focus has been on the use of benthic foraminifera and other meiobenthic organisms (nematodes, ostracods) to contour reservoirs of methane stored under the seabed. Altogether, 173 benthic and 7 planktonic species and subspecies (i.e., lower taxa) have been recorded, with 18 previously reported species not found. Thirty lower taxa belong to agglutinated foraminifera; the rest are calcareous. A vast majority of them are described, illustrated, and supplemented by the data on biostratigraphy and paleoecology in subsequent chapters. Keywords

Ponto-Caspian · Quaternary · Benthic foraminifera · History of study

1.1

Foraminifera of the Black Sea and Sea of Azov

Initially, nine species (no descriptions, no pictures) of benthic foraminifera were listed by Pereialsvtseva (1886) in the Sevastopol Bay and Crimea and were cited by Ostroumov (1893) and Sovinskiy (1904). Other scientists (Zernov 1901, 1906, 1913; Andrusov 1892; Arkhangel’skiy and Strakhov 1938) just mentioned the presence of foraminifera in the Black Sea without any identification. Chishkoff (1912) found Rotalia veneta M.Sch., Polystomella strigilata d’Orb, and Quinqueloculina fusca Brady.

# Springer Nature Switzerland AG 2022 V. Yanko, Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1, https://doi.org/10.1007/978-3-031-12374-0_1

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On the Bulgarian shelf, Lepsi (1931) identified Ammodiscus incertus d’Orb and Peneriplis sp. This list was later expanded by Velkanov (1957) who identified 26 benthic species (no description, no pictures) and Velkanova (1981) who described 43 recent species (39 of them were pictured) from a water depth above 65 m. Dolgopolskaya and Pauli (1931) pictured and described ten species of live foraminifera from the Karadag Biological Station, Crimean Peninsula, providing also their ecological requirements. While Zernov (1906) considered that foraminifera are scarce in the Black Sea, Dolgopolskaya and Pauli (1931) emphasized that foraminifera are abundant and make a significant proportion of benthic assemblages. On the Romanian shelf, foraminifera were firstly mentioned by Margineanu (1958) who calculated the percentage of Ammonia and Nonion in foraminiferal assemblages. Baĉescu and Margineanu (1959) documented the presence of a few Mediterranean species and even planktonic Globigerina bulloides (d’Orb) in this area. Macarovici et al. (1958) and Macarovici and Cehan-Jonesi (1961, 1962, 1966) described the distribution of Rose Bengal-stained foraminifera in the surface sediments of the Romanian shelf. Among 14 benthic species, Ammonia was dominant. The highest abundance of Ammonia was discovered at a water depth of 20–45 m, salinity 15.0–16.5‰, and temperature 10 °C. On the northwestern Ukrainian shelf, Didkovskiy (1959) described three foraminiferal assemblages dominated by Ammonia beccarii (Linne) var. risilla, var. n, Nonion, and Ammonia beccarii (Linne) at a water depth of 4–48 m between the Danube Delta and Egorlitsky Bay. He documented 29 species and varieties; some of them were considered as new. No pictures and descriptions were provided for these species, bringing them into the category of nomen nudum. In 1969, Didkovskiy documented 36 species and varieties near the northern exit of the Bosphorus. Some of them, such as Florilus boueanum (d’Orbigny), were likely reworked from the Miocene. Twenty-three species and three subspecies of benthic foraminifera were documented by Morozova (1964) from the Crimean and Caucasian shelf at water depth 0–103 m. These taxa were grouped in three geographical assemblages: northeastern with dominance of Rotaliidae, Discorbiidae, and Nonionida distributed at the Caucasian shelf; northwestern with dominance of Rotaliidae, Miliolidae, and agglutinated foraminifera distributed counterclockwise from the northwestern Crimea to the Danube Delta; and western assemblage distributed in the Danube Delta and dominated by Rotalia beccarii. The latter one was most impoverished compared to all others. This author noticed that the diversity of foraminifera increases with depth due to an appearance of Lagenidae and Polymorphinidae below 59-m isobath. Mikhalevich (1968) described 27 species of benthic foraminifera from the southern Crimean shelf. Kirienko (1979)

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State-of-the-Art

tried to interpret warm–cold episodes in the geological section of the Bulgarian shelf using left alterations of left and right coiling of Ammonia tests. The most detailed work on recent Black Sea foraminifera was published by Tufescu (1974a, b). She synthesized previously obtained (1967–1974) data (Tufescu 1967, 1968a, b, 1969a, b, 1970, 1971, 1973). Based on 146 stations (depth 0.2–62 m), she described the ecology, zoogeography, and evolution of 21 species, 17 of which were pictured. Based on morphometrical analysis, she divided A. beccarii into A. beccarii sensu stricto and A. tepida. Makaveeva (1979) briefly described the quantitative distribution of five Elphidium and Quinqueloculina species in the Karkinitskiy, Tendrovskiy, and Egorlitskiy Bays and northern Black Sea. She concluded that these epiphytic foraminifera have their highest abundance (225,000 ind/m2) on some algae. A very preliminary study of the Sea of Azov foraminifera was performed by Didkovskiy (1958) who identified nine benthic species and concluded that foraminifera of the Sea of Azov are significantly impoverished compared to those in the Black Sea. Fossil foraminifera from the ODDP drills 379, 380, and 381 in the Black Sea were studied by Cheorghian (1978) who documented 29 benthic species (no pictures, no description) in the Miocene, Pliocene, and Quaternary sediments. He did not discover foraminifera in the postglacial sediments although reworked foraminiferal tests were present throughout as was also noticed by Khrustalev and Shcherbakov (1974) for different geological age sequences. Interestingly, the number of reworked foraminifera increased with the proportion of coarse sediments. Brief data on Quaternary foraminifera of the Kerch Strait were presented by Suprunova (1980) allowing her to divide Quaternary sediments into Drevneuxinian (Old Euxinian), Neoeuxinian (New Euxinian), Drevnechernomorian (Old Black Sea) and Novochernomorian (New Black Sea) layers. Kitovani (1971) and Barg et al. (1982) found benthic foraminifera in the Pleistocene sediments of the Georgian and Crimean coasts, respectively. Temlekov and Mutchinova (1999) investigated the quantitative distribution of the sublittoral benthic foraminifera from the Bulgarian Black Sea coast (the Kiten inlet) and correlated the number of Ammonia and Elphidium specimens with the water depth and grain size of sediments. The taxonomic composition and distribution of foraminifera in the surface sediment from the Kazachya Bay, Crimea, were studied by Anikeeva (2005). Twenty species of benthic foraminifera were identified at 25 stations. Specimens of the genera Ammonia, Elphidium, and Quinqueloculina were the most common. This author failed to correlate foraminiferal distributions to certain environmental factors, e.g., heavy metals.

1.2 Caspian and Aral Seas

However, Kravchuk (1999, 2004) discovered that in fact benthic foraminifera are excellent indicators of marine pollution in the Black Sea. She identified 33 species from 19 genera and 10 families in the northwestern part of the basin. The highest number of species were Elphidiidae (8 species) and Ammonoidea (4 species). Agglutinated foraminifera are represented by rare specimens of Ab. ponticus Mikhalevich and Di. imperspica Yanko. A decrease in the number of species and specimens as well as an increase in the proportion of morphological deformities, stunting, and pyritization of foraminifera was attributed to pollution by domestic sewage and heavy metals. Temlekov et al. (2006) provided an updated checklist of the recent foraminifera from the Bulgarian Black Sea coast. He provided a list of 107 species from 56 genera, 32 families, and 10 orders largely repeating the list of foraminifera provided by Yanko in 1989. The most common species belonging to the Ammonia, Elphidium, Cribroelphidium, and Fisurina genera were pictured in SEM. This author also provides a synecological characteristic of the foraminiferal communities inhabiting five types of sediment from the upper sublittoral (down to 20-m depth), seashore pools, and river mouths of the Bulgarian South Black Sea area, Temlekov (2008). He determined the species’ frequency of occurrence, the dominant structure, and similarity of the foraminiferal communities. Kondariuk (2018) studied the lateral distribution of morphological deformations in 15 foraminiferal tests from the Romanian shelf of the Black Sea in connection with the influence of the freshwater inflow of the Danube River. She emphasized the possibility of using them as indicators of river inflow influence. Some researchers studied soft foraminifera from the Black Sea (e.g., Sergeeva and Anikeeva 2018). But we never found soft foraminifera in our samples neither among recent nor fossil foraminifera and as so their description is not provided in a given monograph.

1.2

Caspian and Aral Seas

The first data on the Caspian foraminifera were presented by Ehrenberg (1873) who found 23 species in the surface sediments of 132 stations located at 6–836 m water depth. Most of the species were distributed above a 25-m isobath; with depth, their number decreased to 1–2 species. At least two species (Rotalia globulosa Ehrenberg and Textularia globulosa Ehrenberg) were recognized as reworked from the Cretaceous sediments. Much later Shokhina (1937) described and pictured six benthic species. Bening (1937) documented 13 species from the Mertvyy Kultuk and Kaydak. Klenova (1956) and Popov (1955) found three recent species and plenty of reworked Cretaceous

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foraminifera. Gofman (1966) documented seven species of dead benthic foraminifera. Not one of the abovementioned studies contained data on the quantitative distribution of foraminifera in the Caspian Sea. There is only a short note of Klenova (1956) that 75% of foraminiferal assemblages consist of Rotalia beccarii (Linne). The Aral Sea foraminifera were studied even worse than those of the Caspian Sea. Only two species of benthic foraminifera are known from the work of Kiaer (1906), Berg (1908), and Hülsen (1911). These species were documented from a depth of 32 m and the sandy and muddy substrata from almost freshwater of the Taldik Bay to the normal salinity of the Aral Sea (Berg 1908). An intensive study of taxonomy, quantitative distribution, and ecology of live (Rose Bengal-stained) foraminifera in the Caspian and Aral Seas and their comparison with recent foraminifera from other basins was performed by Mayer (1968, 1970, 1972, 1974a, b, 1976, 1979a, b, 1983a, b). A comparison of recent and Pliocene–Quaternary foraminifera from this region was performed by Naydina et al. (1974) and Mayer (1975). Mayer (1979b) described 18 species from the Caspian Sea, two of which, Hemisphaerammina sp. and Saccamina sp., were given in open nomenclature; two, Mi. fusca (Brady) and Tr. aguajoi (Bermudez), are well know from other basins; the rest of the species were initially described by Mayer. Eleven Caspian live in the Aral Sea (Mayer 1979b). The abundance of benthic foraminifera in the sediments of the Southern Caspian Sea from Bahnamir to Babolsar, Iran, was studied by Ghane et al. (2014). A total of five species of benthic foraminifera (A. beccarii caspica Shchedrina, El. littorale caspicus [Shokhina], Mi. fusca [Brady], Ammotium sp., and one unidentified species) were identified in the bottom sediments, and their distribution was correlated with environmental parameters (dissolved oxygen, temperature, pH, organic matter, calcium carbonate concentration, and grain size). A study of benthic foraminifera abundance in the sediments of Southern Caspian Sea from Fereydunkenar to Babolsar was performed by Sadough et al. (2013). A multidisciplinary case study from the southeastern flank of the Caspian Sea on sea-level changes at the end of Little Ice Age and its impacts on the avulsion of the Gorgan River was performed by Naderi Beni et al. (2014). A study of benthic foraminifera in the sediments of Southern Caspian Sea was carried out (Ghane et al. 2014). The paper describes foraminiferal species and provides a correlation of their abundance and environmental factors enabling more information on Caspian Sea ecosystem. The effect of urban pollutants on the distribution of benthic foraminifera in the Southern Caspian Sea was studied by Zarghami et al. (2019). The authors recognized 11 species of benthic foraminifera from 6 genera of 5 families. The

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State-of-the-Art

cosmopolitan A. beccarii caspica was common in the studied area. The density of benthic foraminifera is significantly correlated with seasons, the highest density being observed in winter, and the most dense foraminifera were observed in Babolsar stations that showed a good situation for living there; hence, we can use these foraminifera as a bioindicator for pollutant area. To the best of our knowledge, there are no publications (except ours) on the application of benthic foraminifera to the Ponto-Caspian Late Quaternary stratigraphy and paleogeography.

species, e.g., Miocene Florilus boueanum (d’Orbigny) and Nubecularia novorossica (Karrer), seemed to be reworked, and together with other species in open nomenclature, e.g., Ammorginulina sp. (Baĉescu and Margineanu 1959), were excluded from our analysis. Altogether, 180 benthic and seven planktonic species and subspecies are on the list (Table 3.1). Eighteen species are absent in our material (marked by stars in the table). Thirty low taxa belong to agglutinated foraminifera, and the rest are calcareous. A vast majority of them are pictured, described, and supplemented by data on biostratigraphy and paleoecology.

1.3

References

Our Study

The taxonomy, ecology/paleoecology, paleogeography, and biostratigraphy of Ponto-Caspian benthic foraminifera have been carried out by the author of this book in the northwestern (e.g., Yanko 1973, 1974a, b, c, d, e, 1975, 1982a, b, c, d; Voskoboynikov et al. 1985; Sulimov et al. 1983; Vorobyova and Yanko 1986), Bulgarian (Yanko 1979, 1982c; Dimitrov et al. 1979), and Caucasian (Yanko et al. 1983; Yanko and Gramova 1990) and across the shelf (Yanko and Troitskaya 1987) and northern exits of the Bosphorus Strait (Yanko and Vorobyova 1991; Yanko-Hombach 2007a), the Sea of Azov (Yanko 1982d; Yanko and Aleevskaya 1982; Gudina and Yanko 1989; Yanko and Vorobyova 1990), Caspian Sea (Svitoch et al. 1992, 1997; Yanko 1989, 1990), and Mediterranean Sea (Yanko et al. 1994, 1998). In addition, foraminifera from numerous Pleistocene stratotypes were studied in the coastal zone of the Ponto-Caspian region (e.g., Yanko 1989, 1990; Yanko et al. 1990; Svitoch et al. 1992). The application of benthic foraminifera in the reconstruction of sea-level change and coastline migration in regard to the Great Flood Hypotheses in the Black Sea is present in Yanko-Hombach (2003, 2004, 2007a, b) and Yanko-Hombach et al. (2007, 2011, 2014). Lately, attention has been paid on the use of benthic foraminifera along with other meiobenthos organisms (nematodes, ostracods) for contouring reservoirs of methane stored under the seabed (Yanko et al. 2017; Yanko-Hombach et al. 2017; Shnyukov and Yanko 2014; Shnyukov and Yanko-Hombach 2020). Our analysis of published data developed a general list of benthic species for the Ponto-Caspian region. This list includes 183 low taxa (Yanko 1989): 88 species have pictures (e.g., Tufescu 1973, 1974b), and some of them are supported by brief (e.g., Mikhalevich 1968; Temlekov et al. 2006) or more detailed (e.g., Dolgopolskaya and Pauli 1931) description. For other species only location and some ecological remarks (e.g., Vlkanova 1981) are provided. These 88 species are considered in our study. From the rest of the 97 species, which do have neither pictures nor description, only 33 species could be identified in our material. Seventy

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6 Gilyarov MS, Zevina GB (eds) Biologicheskie resursy Kaspiyskogo morya, pp 21–43 (in Russian) Mayer EM (1983b) Raspredeleniye foraminifer v razreze golotsenovykh otlozheniy Aral’skogo morya (Distribution of foraminifers in the section of the Holocene deposits of the Aral Sea). In: Mayev YeG Paleogeografiya Kaspiyskogo i Aral’skogo morey v kaynozoye, Moscow, pp 153–164 (in Russian) Mikhalevich VI (1968) Otryad foraminifery (Order foraminifera). In: Vodyanitskiy VA (ed) Opredelitel’ fauny Chornogo i Azovskogo morey, Kiev, pp 9–21 (in Russian) Morozova VG (1964) Foraminifery biotsenozov shel’fa Chernogo morya (Foraminifera of biocenoses of the Black Sea shelf). Byull. MOIP. Otd. geol. Nov. ser. 69(6):148 (in Russian) Naderi Beni A, Lahijani H, Pourkerman M et al (2014) Caspian Sea-level changes at the end of Little Ice Age and its impacts on the avulsion of the Gorgan River: a multidisciplinary case study from the southeastern flank of the Caspian Sea. Méditerranée 12:145–155 Naydina IN, Mayer EM, Muromtseva TL (1974) Novyye dannyye k faunisticheskoy kharakteristike verkhnepliotsenovykh otlozheniy Severnogo Prikaspiya (New data on the faunistic characteristics of the Upper Pliocene deposits of the Northern Caspian). In: Leont’yev OK, Mayev YEG (eds) Kompleksnyye issledovaniya Kaspiyskogo morya. Moscow, vyp. 4, pp 183–190 (in Russian) Ostroumov AA (1893) Poyezdka na Bosfor (A trip to the Bosphorus). Zapiski Imper. AN LXXII, 55 pp (in Russian) Pereialsvtseva SM (1886) Protozoa Chornogo morya (Protozoa of the Black Sea) Zap. Novorossiysk.ob-va yestestvoisp 10(2):79–114 (in Russian) Popov GI (1955) O stratigraficheskom raschlenenii i sopostavlenii chernomorskikh i kaspiyskikh chetvertichnykh otlozheniy (On stratigraphic division and comparison of the Black Sea and Caspian Quaternary deposits). Dokl. AN SSSR 101(1):143–146 (in Russian) Sadough M, Ghane F, Manouchehri H et al (2013) Identification and abundance of benthic foraminifera in the sediments from Fereidoonkenar to Babolsar of Southern Caspian Sea. Turk J Fish Aquat Sci 13(1):79–86 Sergeeva NG, Anikeyeva OV (2018) Myagkorakovinnyye foraminifery Chornogo i Azovskogo morey (Soft-shelled foraminifera of the Black and Azov Sea). AO Kovalevsky Institute of Marine Biological Research. Simferopol, PP “AR-IAL”, 156 pp (in Russian) Shnyukov EF, Yanko VV (2014) Gazootdacha dna Chernogo morya: geologo- poiskovoye, ekologicheskoye i navigatsionnoye znacheniye (Degasing from the bottom of the Black Sea: geological prospecting, environmental and navigational significance). Visnyk Odes’koho natsional’noho universytetu. Heohrafichni ta heolohichni nauky 19, 3(23):225–241 (in Russian) Shnyukov E, Yanko-Hombach (=Yanko) V (2020) Mud Volcanoes of the Black Sea Region and their environmental significance—2020. Springer, 494 pp Shokhina VA (1937) kornenozhki Mertvogo Kultuka i Kaydaka (Rhizomes of Dead Kultuk and Kaydak). In: Orlov YUA (ed) Osnovy Paleontologii, v. 1 (in Russian) Sovinskiy VK (1904) Vvedeniye v izucheniye fauny Ponto-KaspiyskoAral’skogo morskogo basseyna (Introduction to the study of fauna in the Ponto-Caspian-Aral sea basin). Zap. Kiyevsk. ob-va estestvozn. 18:1–13 (in Russian) Sulimov IN, Blagodarov MI, Yanko VV et al (1983) O roli biogennogo materiala v donnykh osadkakh (On the role of biogenic material in bottom sediments). Tez.dokl.Vsesoyue.soveshch. Biosedimentatsiya v moryakh i okeanakh, 25 September–2 October 1983, Moscow, pp 12–13 (in Russian) Suprunova NI (1980) Etapnost’ razvitiya fauny ostrakod i foraminifer kak osnova biostratigraficheskogo raschleneniya chetvertichnykh otlozheniy Kerchenskogo proliva (Stages in the development of the fauna of ostracods and foraminifers as the basis for the

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State-of-the-Art

biostratigraphic subdivision of the Quaternary deposits of the Kerch Strait). Voprosy mikropaleontologii 23:171–173 (in Russian) Svitoch AA, Yanina TA, Yanko VV et al (1992) Biostratigrafiya razreza Gora bakinskogo yarusa (Biostratigraphy of the Outcrop Gora Bakinskogo Yarusa). Izvestiнa Izvestiya Rossiyskoy Akademii Nauk 2:128–130 (in Russian) Svitoch AA, Selivanov AO, Yanina TA (1997) Paleogeograficheskiye sobytiya pleystotsena Ponto-Kaspiya i Sredizemnomor’ya (materialy po rekonstruktsii i korrelyatsii) (Pleistocene paleogeographic events of the Ponto-Caspian and the Mediterranean (reconstruction and correlation materials)). Moscow, RASKHN, 288 pp (in Russian) Temlekov BK (2008) Ecological characteristics of the foraminiferal fauna (Protozoa: Foraminifera) of the Bulgarian South Black Sea Area. Acta Zoolooica Bulgarica, Suppl. 2:275–282 Temlekov BK, Mutchinova PE (1999) About the quantative distribution of sublittoral foraminifera in the inlet of Kiten, Bulgarian Black Sea Coast. Acta Zoolooica Bulgarica 51 (2/3):73–78 Temlekov BK, Golermansky VG, Todorov MT (2006) Updated checklist of the recent foraminifera from the Bulgarian Black Sea Coast. Acta Zoolooica Bulgarica 58(1):17–36 Tufescu M (1967) Esquisse de dynamique saisonniere du elute Ammonia Brimmed dans la zone de ressac sur le littoral Romain. An. Univ. Bucuresti. Ser. Biol., pp 145–150 Tufescu M (1968a) Probremes de biogeographie concernant les fo-raminiferes pontiques. Rev. roum. geol. geophys. et geogr. Stir. geogr. 14(2):253–265 Tufescu M (1968b) Ammonia tepida (Cushman) (ord. Foraminifera). Some features of its variability in the Black Sea Basin. Revue Roumaine de Biologie et Zoologie 13:169–177 Tufescu M (1969a) Foraminifera of the zone of the Romanian littoral lakes, lagoons and mouths of the Danube—Sulina. Hidrobiologia 10:291–302 Tufescu M (1969b) Sur la presence de Trichochyalis aguajoi (Bermudez) dans la Mer Noire. Rev. micropaleontol 12(1):46–52 Tufescu M (1970) Problemes de boigeographic concernant les foraminiferes pontiques. Rev. roum. Geol. Geophys. Geogr, Ser. Geogr. 14(2):253–265 Tufescu M (1971) Structure topografica a populatiilor de foraminifere din apele litorale romanesti. Dari de seams 57(3):103–113 Tufescu M (1973) Les associations de foraminiferes du Nord-Quest de la Mer Noire. Rev Espanola de Micropaleontol 5(1):15–32 Tufescu M (1974a) The dual origin of the Black Sea foraminifera. Internationale Revue der gesamten Hydrobiologie und Hydrographie 59(3):353–365 Tufescu M (1974b) Populatiile de foraminifere din apele litorale romanesti. Edit. Acad. Rep. Soc. Romania. Bucuresti, 75 pp Velkanov A (1957) Katalog na nashate chernomorska fauna. Tr. Morsk. Biologist. st. Varna 19:1–61 Vlkanova IKH (1981) Retsentni foraminiferi (Rhizopodea, Foramіnіferida) pred blgarskiya bryag na Cherno more. Acta Zoologica Bulgarica 18:3–14 Vorobyova L, Yanko VV (1986) O raspredelenii sovremennykh foraminifer v severo-zapadnoy chasti Chornogo morya (On the distribution of modern foraminifers in the northwestern part of the Black Sea). Zoological zhurnal 1(8):I250–1254 (in Russian) Voskoboynikov VM, Krakovskiy BI, Konnikov EG, Yanko VV (1985) Litologiya chetvertichnykh otlozheniy: Tekstury donnykh otlozheniy (Lithology of Quaternary sediments: textures of the bottom sediments). In: Shnyukov E F (ed) Geologiya shelfa USSR: Litologiya. Naukova Dumka, Kiev, pp 93–130 (in Russian) Yanko VV (1973) Foraminifery dzhemetinskikh otlozheniy severozapadnoy pasti Chornogo morya. Tezisy dokladov VI Vsesoyuznogo mikropaleontologicheskogo soveshchaniya, Novosibirsk, p 196 (in Russian)

References Yanko VV (1974a) Pozdnechetvertichnyye foraminifery severozapadnogo shel’fa Chornogo morya (Late Quaternary foraminifera of the North-West Shelf of the Black). PhD thesis, Odessa I. I. Mechnikov State University, 227 pp (in Russian) Yanko V (1974b) Stratigrafiya i nekotoryye paleogeograficheskiye osobennosti razvitiya severo-zapadnogo shel’fa Chornogo morya v pozdnetsetvertichnoye vremya po bentosnym foraminiferam (Stratigraphy and some paleogeographical features of the development of the north-western shelf of the Black Sea in the Late Quaternary according to benthic foraminifers). In: Veklich MF et al (ed) Paleogeografiya i inzhenernaya geologiya (pozdniy kaynozoy). Kiev, pp 93–96 (in Russian) Yanko V (1974c) Pozdnechetvertichnyye foraminifery severozapadnogo shel’fa Chornogo morya (Late Quaternary foraminifers of the Northwestern shelf of the Black Sea). Byull. MOIP. Otd. Geol. 5:149–150 (in Russian) Yanko V (1974d) Deyaki dani pro foraminifery donnykh vidkladiv okremykh’ dilyanok Pivnichno-3akhidnoho ta Kavkaz’koho shel’fu Chornoho morya (Some data on foraminifera donnik deposits of separate parts of the North-Eastern and Caucasian shelf of the Black Sea). In: Babynets AE (ed) ta in. Geologiya uzberezhzhya i dna Chornoho ta Azovs0 kogo moriv u mezhakh URSR 7:39–43 (in Ukrainian) Yanko V (1974e) Novyye vidy bentosnykh foraminifer iz golotsenovykh otlozheniy Severo-Zapadnogo shel’fa Chornogo morya (New species of benthic foraminifers from the Holocene deposits of the north-western shelf of the Black Sea). Paleontologicheskiy sbornik 1(11):24–30 (in Russian) Yanko V (1975) Foraminifery sovremennykh donnykh otlozheniy severoØzapadnoy akvatorii Chornogo morya (Foraminifers of modern bottom sediments of the northwestern water area of the Black Sea). In: Fursenko AV (ed) Obraz zhizni i zakonomernosti rasseleniya sovremennoy i iskopayemoy mikrofauny. Tr. IGiG; Vyp.333, pp 73–79 (in Russian) Yanko V (1979) Stratigrafiya i geochronologoya donnykh otlozheniy: Stratigraficheskie kompleksy bentocnykh foraminifer. Obschaya stratigraficheskaya skhema (Stratigraphy and geochronology of the bottom sediments: stratigraphic complexes of benthic foraminifera. General stratigraphic scheme). In: Malovitskiy Y (ed) Geology and hydrology of the Western Part of the Black Sea. Ac. Sc. Bulg., Sofia, pp 82–95 (in Russian) Yanko V (1982a) Paleoekologiya pozdnetsetvertichnykh foraminifer severo-zapadnogo shel’fa Chornogo morya i ikh znacheniye dlya vyyasneniya usloviy obrazovaniya osadkov (Paleoecology of Late Quaternary foraminifers of the Northwestern shelf of the Black Sea and their significance for elucidating the conditions for the formation of sediments). Moscow, Deposited in VINITI 720–81, pp 131–137 (in Russian) Yanko V (1982b) Stratigrafiya verkhnechetvertichnikh otlozheniy severo-zapadnogo shel’fa Chernogo morya po bentosnim foraminiferam (Stratigraphy of the upper Quaternary sediments of the Black Sea north-western shelf based on benthic foraminifera). In: Zhuze AP (ed) Morskaya mikropaleontologiya. Nauka, Moscow, pp 126–131 (in Russian) Yanko V (1982c) Sistematicheskiy sostav i kolichestvennoye rasØpredeleniye foraminifer v poverkhnostnom sloye donnykh osadkov Bolgarskogo shel’fa Chornogo morya (Systematic composition and quantitative distribution of foraminifers in the surface layer of bottom sediments of the Bulgarian shelf of the Black Sea). Moscow, Deposited in VINITI 720-82, pp 73–78 (in Russian) Yanko V (1982d) Stratigrafiya donnykh otlozheniy Chornogo i Azovskogo morey po bentosnym foraminiferam (Stratigraphy of bottom sediments of the Black and Azov Seas according to benthic foraminifers). Abstracts of the XI International Congress INQUA. Moscow, pp 314–315 (in Russian)

7 Yanko V (1989) Quaternary foraminifera of the Ponto-Caspian Region (the Black Sea, the Sea of Azov, the Caspian Sea and the Aral Sea): taxonomy, biostratigraphy, history, ecology. Doctoral thesis, Moscow State University, two volumes, 1000 pp (in Russian) Yanko V (1990) Stratigraphy and paleogeography of marine Pleistocene and Holocene deposits of the southern seas of the USSR. Memorie Società Geologica Italiana 44:167–187 Yanko-Hombach V (2003) “Noah’s flood” and the late Quaternary history of the Black Sea and its adjacent basins: a critical overview of the flood hypotheses. GSA topical session T104 “Noah’s flood” and the Late Quaternary geological and archaeological history of the Black Sea and adjacent basins. Geological Society of America, September 2003. Abstracts with Programs 35(6):460 Yanko-Hombach V (2004) The Black Sea controversy in light of the geological and foraminiferal evidence. In: Yanko-Hombach V, Görmüs M, Ertunç A et al (eds) 4th EMMM 2004, Program and extended abstracts of the fourth international congress on environmental micropalaeontology, microbiology, and meiobenthology (13–18 September 2004, Isparta, Turkey), pp 224–227 Yanko-Hombach V (2007a) Controversy over Noah’s Flood in the Black Sea: geological and foraminiferal evidence from the shelf. In: Yanko-Hombach V, Gilbert AS, Panin N, Dolukhanov PM (eds) The Black Sea flood question: changes in coastline, climate and human settlement. Springer, Dordrecht, pp 149–203 Yanko-Hombach V (2007b) Late Quaternary history of the Black Sea: an overview with respect to the Noah’s Flood hypothesis. In: Erkut G, Mitchell S (eds) The Black Sea: past, present and future: proceedings of the international, BIAA monograph series 42, pp 5–20 Yanko V, Aleevskaya N (1982) Pozdnechetvertichnyye bentosnyye foraminifery Azovskogo morya (Late Quaternary benthic foraminifers of the Sea of Azov). Moscow, Deposited in VINITI 3910–82, pp 79–84 (in Russian) Yanko V, Gramova L (1990) Stratigraphy of the Quaternary sediments of the Caucasian shelf and continental slope of the Black Sea on microfauna (foraminifera and ostracoda). J Sov Geol 2: 60–72 (in Russian) Yanko V, Troitskaya T (1987) Pozdnechetvertichnye foramifery Chernogo morya (Late Quaternary foraminifera of the Black Sea). Nauka, Moscow (in Russian) Yanko V, Vorobyova LV (1990) Sovremennye foraminiferi Azovskogo morya (Recent Foraminifera of the Sea of Azov). Ekologia Moria 35: 29–34 (in Russian) Yanko V, Vorobyova LV (1991) Foraminiferi bosphorskogo rayona Chernogo morya (Foraminifera of the Bosphorus Region of the Black Sea). Ekologia morya 39:47–50 (in Russian) Yanko V, Berdnikova VG, Katzuk AV (1983) Stratigraphiya donnykh otlozheniy Kavkazskogo shelfa Chernogo morya po molluskam, foraminiferam i ostracodam (Startigraphy of the bottom sediments of the Black Sea Caucasian shelf on mollusks, foraminifera and ostracods). VINITI, no 2117–83, pp 45–50 (in Russian) Yanko V, Frolov VT, Motnenko IV (1990) Foraminifery i litologiya stratotipicheskogo gorizonta (antropogen Kerchenskogo poluostrova) (Foraminifera and lithology of the stratotypical horizon (Anthropogene of the Kerch peninsula)). Bulletin of the Moscow Society of the Investigators of Nature, Geology 65(3):85–97 (in Russian) Yanko V, Kronfeld A, Flexer A (1994) The response of benthic foraminifera to various pollution sources: implications for pollution monitoring. J Foraminifer Res 24:1–17 Yanko V, Ahmad M, Kaminski M (1998) Morphological deformities of benthic foraminiferal tests in response to pollution by heavy metals: implications for pollution monitoring. J Foraminifer Res 28(3): 177–120

8 Yanko V, Kravchuk A, Kulakova I (2017) Meiobentos myetanovykh vykhodov Chyernogo morya (Meiobenthos of methane outlets of the Black Sea). Phenix, Odessa, 241 pp (in Russian) Yanko-Hombach V, Gilbert A, Dolukhanov P (2007) Critical overview of the Flood Hypotheses in the Black Sea in light of geological, paleontological, and archaeological evidence. Quat Int 167–168:91– 113 Yanko-Hombach V, Mudie P, Gilbert AS (2011) Was the Black Sea catastrophically flooded during the post-glacial? Geological evidence and archaeological impacts. In: Benjamin J, Bonsall C, Pickard DRC, Fischer A (eds) Underwater archaeology and the submerged prehistory of Europe. Oxbow Books, pp 245–262 Yanko-Hombach V, Mudie PJ, Kadurin S, Larchenkov E (2014) Holocene marine transgression in the Black Sea: new evidence from the northwestern Black Sea shelf. Quat Int 345:100–118 Yanko-Hombach V, Schnyukov E, Pasynkov A et al (2017) Late Pleistocene-Holocene environmental factors defining the AzovBlack Sea Basin, and the identification of potential sample areas

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State-of-the-Art

for Seabed Prehistoric Site prospecting and landscape exploration on the Black Sea Continental Shelf. In: Flemming F et al (eds) Submerged landscapes of the European Continental Shelf: Quaternary paleoenvironments. Wiley-Blackwell, Chichester, pp 431–478 Zarghami M, Al-Maliky THY, Nazarhaghighi F, Sohrabi Mollayousefi M (2019) Effect of urban pollutants on distribution of benthic foraminifera in the Southern of Caspian Sea. In: Proceedings of the International Academy of Ecology and Environmental Sciences Zernov SA (1901) Plankton Azovskogo morya i yego limanov (Plankton of the Sea of Azov and its estuaries). Yezhegodnik zool. muzeya AN, Sankt Petresburg 6:38–I26 (in Russian) Zernov SA (1906) Penilia shamacheri Rich. v Chornom more v Karkinitskom zalive (Penilia shamacheri in the Karkinitian Bay, Black Sea). Yezhegodnik zool. muzeya AN, Sankt Petresburg 13(4):28–35 (in Russian) Zernov SA (1913) K voprosu ob izuchenii zhizni Chornogo morya (On the issue of studying the life of the Black Sea). Seria VIII, Phiz.-mat. otd., vol XXXII, no 1, 299 pp (in Russian)

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Study Area, Material, and Methods

Abstract

This chapter provides an overview of the study area known as the Ponto-Caspian, which includes the Black Sea, the Sea of Azov, the Caspian Sea, the Aral Sea (currently dried up), and connecting straits. The Black Sea, with a maximum depth of 2212 m, is the easternmost of the seas of the Atlantic Ocean basin and can be considered as the most isolated sea of the modern Global Ocean. The Sea of Azov is connected to the Black Sea via the Kerch Strait. On average, the level of the Black Sea is 7–11 cm lower than that of the Sea of Azov and 30 cm higher than that of the Sea of Marmara. The Black Sea exhibits the standard oceanic provinces of the continental shelf, slope, and abyssal plain. The extensive continental shelf accounts for 25% of the sea area; the 200-m isobath is commonly taken as the shelf boundary of the Global Ocean. The bottom relief is largely smooth due to the distribution of sediment discharge from major lowland European rivers, such as the Danube, Dnieper, Dniester, and the Southern Bug, that together discharge 56.8 million tons of sediments annually. The average bottom-water salinity of the Black Sea (17 psu) is only half that of the Eastern (39 psu, practical salinity units) and Western (34 psu) Mediterranean. Salinity varies spatially, temporally, and with depth, in association with freshwater discharge into the basin and the influence of subsurface inflow from the Sea of Marmara with salinity of ~35 psu. The maximum salinity of the Sea of Azov is 13 psu. The Caspian Sea is the world’s largest inland body of water, variously classed as an isolated lake without an outflow (i.e., an endorheic basin). It lies at the junction of Europe and Asia, with the Caucasus Mountains to the west and the steppes of Central Asia to the east. The Caspian Sea, like the Black Sea, is a remnant of the ancient Paratethys Sea. It became landlocked about 5.5 million years ago due to tectonic uplift and falling sea level.

The Caspian Sea is divided into three distinct physical regions: the Northern, Middle, and Southern Caspian. The Northern Caspian only includes the Caspian shelf and is very shallow, with an average depth of only 5–6 m; it accounts for less than 1% of the total water volume. More than 130 rivers flow into the Caspian, with the Volga River being the largest. The Northern Caspian water is almost fresh, becoming more brackish toward the south. The sea is most saline off the Iranian shore, where the watershed contributes little inflow. Currently, the mean salinity of the Caspian is 12.8 psu on average, varying from 1 psu near the Volga outlet to a high of 200 psu in the Kara-Bogaz-Gol, where intense evaporation occurs. The Aral Sea was also an endorheic lake; lying between Kazakhstan and Uzbekistan, it began shrinking in the 1960s and had largely dried up by the 2010s. Formerly it was the fourth largest lake in the world with an area of 68,000 km2. The shrinking of the Aral Sea has been called “one of the planet’s worst environmental disasters.” In the Black Sea, the Sea of Azov, and the Kerch Strait, sediment samples have been collected since mid-1970s using various research vessels. In limans (marshes) and river deltas, the samples were collected from small vessels by hand corer or diving. As a rule, the sampling campaigns took place in May–June. Samples for foraminiferal analysis were obtained from multiple organizations of the Black Sea countries. In total, approximately 32,000 samples from 1500 grabs, multicorers, box corers, 4300 gravity/piston cores, and 56 boreholes (up to 40 m in length) were investigated. The samples were obtained in limans, lagoons, river deltas, shelf, and continental slope of the Black Sea, the Kerch Strait, the Sea of Azov, and the Caspian Sea. The total length of investigated sediment cores reached 8000 m. Marine research was supplemented by the study of 112 Quaternary outcrops including stratotypes located on the Crimean and

# Springer Nature Switzerland AG 2022 V. Yanko, Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1, https://doi.org/10.1007/978-3-031-12374-0_2

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Kerch Peninsula, Caucasian, and northwestern coast of the Caspian Sea. At most marine stations, environmental (e.g., salinity, depth, temperature, dissolved oxygen) and foraminiferal parameters were measured. The total assemblage, including live (stained) and dead (empty) tests of foraminifera, was determined and expressed as the number of tests (abundance) per 50 g of dry sediment. To identify possible interrelations among foraminiferal characteristics and environmental parameters, a range of statistical methods were applied. The inner structure of foraminiferal tests and their wall ultrastructure were studied in 320 thin sections and 120 artificial models. All species of benthic foraminifera were morphologically examined, taxonomically identified, and SEM imaged. Particular attention was paid to morphological deformities of foraminiferal tests as indicators of environmental stress. The collections of benthic foraminifera from the Black Sea, Caspian Sea, and Sea of Azov are stored in the Paleontological Museum of Odessa National University, Ukraine. The stratigraphic work follows the Russian subdivision of the Quaternary System (Zhamoida, Stratigr Geol Correl, 12:321–330, 2004) into the Eopleistocene (1.8–0.8 Ma), Neopleistocene (0.8–0.01 Ma), and Holocene (0.01–0.0 Ma). The resulting high-resolution stratigraphy and geochronology of the Late Quaternary (Neopleistocene and Holocene) sediments are based on the combination of bio- and ecostratigraphic criteria supplemented by absolute dating. Biostratigraphic criteria include precise taxonomic analysis of benthic foraminifera to trace species evolution and to discover species indicators for certain time intervals. Due to the shortness of the studied time interval, the application of these criteria is limited and so is supplemented by ecostratigraphic criteria. Keywords

Black Sea · Sea of Azov · Caspian Sea · Aral Sea · Drillholes · Outcrops

2.1

Study Area

The study area is known as the Ponto-Caspian, which includes the Black Sea, the Sea of Azov, the Caspian Sea, the Aral Sea (currently dried up), and connecting straits (Fig. 2.1). The Black Sea is the easternmost of the seas of the Atlantic Ocean basin. Considering the ratio of the sea volume to the summary area of the cross sections of all its straits (which is 0.04 km2 for the Bosphorus and 0.02 km2 for the Kerch

Strait) as a measure of isolation of a sea basin, then the Black Sea can be considered the most isolated sea of the modern Global Ocean (Zubov 1956). Its maximum length (along 42
290 N lat) and width are 1148 km and 611 km, respectively. Its surface area (excluding estuaries, such as the Dnieper-Bug liman—liman is a local term for ancient estuaries in the Black Sea and Sea of Azov) and its volume are about 416,790 km2 and 535,430 km3, respectively, and the maximum depth is 2212 m (Ivanov and Belokopytov 2013). The Sea of Azov is connected to the Black Sea via the Kerch Strait and has an area of 39,000 km2 and a volume of 290 km3. The maximum length, width, and depth of the Sea of Azov are 360 km, 180 km, and 14 m, respectively. On average, the level of the Black Sea is 7–11 cm lower than that of the Sea of Azov and 30 cm higher than that of the Sea of Marmara. The details and taphonomic conditions of the Black Sea are unusual. It is the world’s largest anoxic (oxygen-free) marine basin. Its strongly stratified water column possesses (1) a thin, well-oxygenated surface layer (20–30 m) with low salinity and warm temperatures, (2) a low-oxygen (suboxic) transition layer (30–150 m), and (3) a thick bottom layer of colder, denser, and more saline water lacking oxygen but high in sulfides. The Black Sea lies within the Anatolian sector of the Alpine-Himalayan orogenic system, located between the Eurasian plate to the north and the African-Arabian plates to the south. Global plate models (DeMets et al. 1990) and recent space geodetic measurements (Smith et al. 1994; Reilinger et al. 1997) indicate that, in the surrounding region, the northward-moving African and Arabian plates are colliding with the Eurasian plate. From this collision, the Anatolian block is moving westward with a rotation pole located approximately to the north of the Sinai Peninsula (Tari et al. 2000). The northward movement of the Arabian plate and westward escape of the Anatolian block along the North and East Anatolian faults have been accompanied by several episodes of extension and shortening since the Permian (Yilmaz 1997; Robertson et al. 2004), as can be seen in seismic-reflection data (McKenzie 1972; McClusky et al. 2000). The Black Sea exhibits the standard oceanic provinces of the continental shelf, slope, and abyssal plain (Fig. 2.2). The extensive continental shelf accounts for 25% of the sea area; the 200-m isobath is commonly taken as the shelf boundary of the Global Ocean. The northwestern shelf extends 220 km outward and occupies 16% of the sea area (68,390 km2) and 0.7% of the water volume (3555 km3) between the Chersonesus and Kaliakra capes. In the flattened and gently sloping part of the shelf adjacent to the shore, depths are 30–40 m, and the bottom slope is 1–2
. Its steepness increases toward the shelf break to 10–12
. Against the flat plain of the shelf, several large, shallow paleo-river valleys are visible in Fig. 2.2, separated by low underwater

2.1 Study Area

11

Fig. 2.1 The Ponto-Caspian. Manych Outlet (currently Manych Depression in red) presumably connected the Caspian and Black Seas at 18 ka BP to 10 ka BP

hills (Ivanov and Belokopytov 2013). The bottom relief is largely smooth due to the distribution of sediment discharge provided by major lowland European rivers, such as the Danube, Dnieper, Dniester, and Southern Bug, that together discharge 56.8 million tons of sediments annually (Panin and Jipa 2002). No known expressions of active tectonic movements have influenced the ancient shoreline positions and deposition of sediments in any appreciable way. The other, less extensive shelf areas of the Black Sea include the coastal zone of Bulgaria and western Turkey from Cape Kaliakra to the city of Ereğli (shelf width up to 50 km), the Kerch–Taman shelf (shelf width up to 50 km), the central Anatolian coast from Cape Kerempe to the city of Giresun (shelf width up to 35 km), the southern Crimean coast between capes Chersonesus and Ai-Todor (shelf width up to 30 km), and the Gudauta Bank in the vicinity of Ochamchira town (shelf width up 20 km) (Ivanov and Belokopytov 2013). Narrow shelves with widths of several kilometers are located along the Caucasian and Anatolian coasts, as well as along the southern Crimean coast from Yalta to Cape Meganom. Their slopes are considerably steeper compared to the broader shelves, ranging from 5–6
to 30
. The shelf break lies at depths from 100 to 200 m, and the slope is 1–2
. The depth of the shelf break is close to 100 m, compared to areas with broader shelves where the break can exceed 200 m. The predominantly flat bottom of the Sea of Azov descends gradually to the depression at its center. At the

bottom, there are a few positive relief forms, the largest of them being the Pischana Bank. The continental slope descends down to 1600–1900 m of water depth with a considerable gradient from 11
to 13
, sometimes reaching 38
in the regions along the southern Crimean and Turkish coasts. The surface of the continental slope is complicated with blocks of the Earth’s crust that often give it a graduated profile, revealing underwater canyons of different origins. They can begin in the coastal zone at depths of 10–15 m and extend as deep as 1600 m. These canyons are the most important route for the transfer of sedimentary material from the coast to the abyssal depression of the Black Sea (Fig. 2.2). In the deepest part of the canyons, at depths of 1600–1900 m, sedimentary material forms extensive cones. Individual cones can coalesce to form the continental subslope. Thus, the morphogenesis of the slope is directly linked to selective erosion and denudation of rocks with different physical and mechanical properties. Erosive and denudation activities in the canyons caused the emergence of huge underwater amphitheaters forming deepwater fans and plumes of terrigenous sediments on the footslope. The abyssal plain is bounded by the 2000-m isobaths and occupies about 35% of the total sea area. It is a relatively flat, accumulative plain with a slight slope to the south. The bottom of the abyssal basin is characterized by hilly relief; slope angles vary from 0
to 1
. According to echo-sounding surveys, significantly large features of submarine relief are absent (Fig. 2.2). Deposits covering the abyssal plain form

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Fig. 2.2 Digital high-resolution map of relief. Modified after Bagrov et al. 2012. 1–24 canyons: (1) Dunaisky, (2) Dniestrovsky, (3) Peleokalanchaksky, (4) Donuslavsky, (5) Al’minsky, (6) Kachinsy, (7) Forossky, (8) Yaltinsky, (9) Khapkhal’sky, (10) Meganomsky, (11) Kubansky, (12) Novorossiysky, (13) Tuapsinsky, (14) Sochinsky, (15) Sukhumsky, (16) Rioniisky, (17) Batumsky, (18) Trabzonsky, (19) Ordossky, (20) Samsunsky, (21) Kysyl-Irmansky, (22) Sakar’iaksky, (23) Bosphorsky, and (24) Burgazsky. Red, blue, and yellow triangles ¼ mud volcanoes reliably detected, discovered based on geophysical data, and expected, respectively. Blue lines ¼ river

paleovalleys. Yellow cylinders ¼ gas seeps. The map was compiled based on results of digitizing bathymetric maps of different scales (1: 50,000, 1:100,000, 1:200,000, 1:500,000, 1:1,125,000) produced by Gosgidrografiya USSR. The Crimean shelf and slope, as well as the southern part of the northwestern shelf and slope, were adjusted based on the results of our own sonar and hydroacoustic soundings obtained during marine expeditions on R/Vs Kiev, Professor Vodyanitsky, Mikhail Lomonosov, Ichthyander, and Vernadsky. Reproduced with permission from the Research Center of Sustainable Development, Ukraine. (From Yanko-Hombach et al. 2017, their Fig. 16.14)

11 material-genetic types. Six types are shallow, and five are deep water. Between all types of deposits, there is a continuous transfer, conditioned by gradual change in grain size and composition. The mean rate of accumulation at the bottom of the central abyssal depression is 30–40 mm/kyr. Today, the Bosphorus Strait (Figs. 2.1 and 2.2) is the only passage for exchange of water and organisms between the Black Sea and Sea of Marmara. This zigzagging strait is about 35 km in length, 0.7–3.5 km in width, and 35.8 m deep, on average, with a few elongate potholes (about 110 m in depth each) on the bottom. The strait possesses two sills, one in the north at a water depth of 59 m and one in the south at a water depth of 34 m, each located about 3 km from the corresponding entrance to the strait. The two directions of water flow within the strait overlap each other: the northward underflow (inflow) from the Sea of Marmara has an average

salinity of 38 psu and a velocity of 5–15 cm/s, and the southward overflow (outflow) from the Black Sea has an average salinity of 18 psu and a velocity of 10–30 cm/s. Due to the sills, the interface between the two flow directions rises from 50 m at the northern end to 20 m at the southern end. The underflow is initiated by the difference in water density between the Black Sea and the Sea of Marmara; the pressure gradient pushes against the Black Sea and powers the underflow. The outflow is initiated by two main factors: (1) the 30-cm elevation of the Black Sea surface above that of the Sea of Marmara, which, in turn, is 5–27 cm above the level of the northern Aegean Sea and (2) the positive balance of the Black Sea, where precipitation (575 km3/year) exceeds evaporation (350 km3/year), producing a discharge of about 600 km3 of brackish water annually (Yanko-Hombach 2007a).

2.2 Material and Methods

The Kerch Strait connects the Black Sea with the Sea of Azov (Figs. 2.1 and 2.2) and is 45 km long, 4.5 km wide, and up to 6 m deep. The shallowness of the strait results in reduced water exchange between the two basins, which is five to ten times smaller than that of the Bosphorus. The Black Sea average bottom water salinity (17 psu) is only half that of the Eastern (39 psu) and Western (34 psu) Mediterranean. The use of “psu” (practical salinity units) instead of the former ‰ is explained in Yanko-Hombach et al. (2013). Salinity varies spatially, temporally, and with depth, in association with freshwater discharge into the basin and the influence of subsurface inflow from the Sea of Marmara with salinity of ~35 psu. Surface salinity ranges from 1 to 3 psu in the Danube delta to 26.2 psu in the Bosphorus outlet area. The maximum salinity of the Sea of Azov is 13 psu. The Caspian Sea is the world’s largest inland body of water, variously classed as an isolated lake without an outflow (i.e., an endorheic basin). It lies at the junction of Europe and Asia, with the Caucasus Mountains to the west and the steppes of Central Asia to the east. The sea is bordered by Russia to the northwest, Azerbaijan to the west, Iran to the south, Turkmenistan to the southeast, and Kazakhstan to the northeast. The Caspian Sea, like the Black Sea, is a remnant of the ancient Paratethys Sea. Thus, the Caspian Sea is underlain by oceanic basalt and not by continental granitic rock. It became landlocked about 5.5 million years ago due to tectonic uplift and a fall in sea level. The Caspian Sea has a surface area of 371,000 km2 (excluding the Gara-Bogaz-Gol lagoon) and a volume of 78,200 km3. The lake has a north–south orientation and consists of two deep basins that occupy its central and southern areas leading to both spatial and depth differences in temperature, salinity, and ecology. The Caspian Sea spreads out over nearly 1200 km from north to south, with an average width of 320 km. With associated wetlands, it covers a region of around 386,400 km2 and its surface is about 27 m below sea level. The Caspian Sea is divided into three distinct physical regions: the Northern, Middle, and Southern Caspian. The northern–middle boundary is the Mangyshlak Threshold, which runs through Chechen Island and Cape Tiub-Karagan. The middle–southern boundary is the Apsheron Threshold, a sill of tectonic origin between the Eurasian continent and an oceanic remnant that runs through Zhiloi Island and Cape Kuuli. Differences among the three regions are dramatic. The Northern Caspian only includes the Caspian shelf and is very shallow, with an average depth of only 5–6 m; it accounts for less than 1% of the total water volume. The Northern Caspian freezes in the winter, and in the coldest winters, ice forms in the south as well. The sea floor deepens toward the Middle Caspian, where the average depth is 190 m. The Southern Caspian is the deepest, with oceanic depths of over 1000 m.

13

The Middle and Southern Caspian account for 33% and 66% of the total water volume, respectively. More than 130 rivers flow into the Caspian, with the Volga River being the largest. The Northern Caspian water is almost fresh, becoming more brackish toward the south. The sea is most saline off the Iranian shore, where the watershed contributes little inflow. Currently, the mean salinity of the Caspian is 12.8 psu on average, varying from 1 psu near the Volga outlet to a high of 200 psu in the Kara-BogazGol, where intense evaporation occurs. In the open sea, the vertical distribution of salinity is markedly uniform; from the surface to the bottom it increases by only 0.1–0.2 psu. Caspian waters differ chemically from typical ocean waters and resembles other evaporative lake waters in having higher concentrations of calcium and magnesium cations, and sulfate and carbonate cations, and lower chloride content. The Aral Sea was also an endorheic lake; lying between Kazakhstan and Uzbekistan it began shrinking in the 1960s and had largely dried up by the 2010s. Formerly it was the fourth largest lake in the world with an area of 68,000 km2. The shrinking of the Aral Sea has been called “one of the planet’s worst environmental disasters” (Daily Telegraph 2010). The region’s once-prosperous fishing industry has been devastated, bringing unemployment and economic hardship. The Aral Sea region is also heavily polluted, with consequential serious public health problems. UNESCO added the historical documents concerning the collapse of the Aral Sea to its Memory of the World Register as a unique resource to study this “environmental tragedy.”

2.2

Material and Methods

In the Black Sea, the Sea of Azov, the Kerch Strait and the Caspian Sea sediment samples have been collected since the mid-1970s using various research vessels, including the R/V Professor, R/V Akademik, R/V Antares, R/V Vladimir Parshin, and R/V Mare Nigrum. In limans (marshes) and river deltas, the samples were collected from small vessels by hand corer and/or diving. As a rule, the sampling campaigns took place in May– June (e.g., Yanko 1979, 1989, 1990; Yanko and Troitskaya 1987; Yanko and Gramova 1990; Yanko-Hombach 2007a; Yanko-Hombach et al. 2014, 2017; Yanko et al. 2019; Yanko and Kondariuk 2020). Samples for foraminiferal analysis were obtained from multiple organizations (e.g., Siberian Branch of the USSR Academy of Sciences; Southern Branch of the Institute of Oceanology, USSR Academy of Sciences; Institute of Oceanology of the Bulgarian Academy of Sciences; Yuzhmorgeologiya, GeoEcoMar; Odessa I.I. Mechnikov National University; Prichernomor DGRP M.V. Lmonosov Moscow State University and some others with which the author cooperated on the contractual and/or scientific basis). In this regard, the following scientists, V. M. Voskoboynikov,

14

A. Yu. Glebov, V. I. Dmitrienko, Ya A. Izmailov, G. I. Karmishina, V. A. Karpov, E. G. Konnikov, E. M. Mayer, I. V. Pogrebnyak, E. F. Shnyukov, A. A. Svitoch, G. G. Tkachenko, T. S. Troitskaya, and K. M. Shimkus, must be mentioned, with deep gratitude to all. The location maps of the studied materials are provided in Yanko (1989, 1990) and Yanko-Hombach et al. (2014, 2017), as well as in Figs. 2.3 and 2.4. At the majority of the sampling stations, salinity, temperature, pH, and DO of bottom water were taken using the Neil Brown Instrument Systems (CTD) with a General Oceanic rosette equipped with 6–11 Niskin bottles and electronic sensors, as described in Yanko et al. (1998) and Yanko-Hombach et al. (2017). In addition, in some areas (e.g., Romanian shelf), additional hydrological parameters were measured: conductivity (U), transparency (Tr), oxygen saturation index (SI), and oxygen-reduction potential (Eh) later normalized to standard pH ¼ 7 (for building the Purbae diagram). Transparency was measured by Secchi disk. Although salinity based on conductivity measurements are now considered to be unitless (e.g., https://unesdoc. unesco.org/ark:/48223/pf0000065031), salinity data reported

2 Study Area, Material, and Methods

in this book were taken by a variety of methods, including conductivity and refractometry, and were originally reported as ‰ or psu. For readers who may be unfamiliar with recent recommendations, the author will use psu when reporting or discussing salinity. Concentrations of phosphates (РО43) and dissolved silica (SiO2) in the water column were calculated with the help of the molybdovanadate method by acid persulfate digestion (HASH equipment); the concentration of CaCO3 was measured by titration. For the calculation of total carbon (C) and total nitrogen (N), the ground samples from the superficial (0–1 cm) sediment layer collected by a multicorer were analyzed using a CNS elemental analyzer Carlo Erba NA 1500 and gas chromatography. Organic carbon (Corg) was calculated after threefold removal of inorganic carbon. The analyses were performed at the Institute of Biochemistry and Marine Chemistry of the Hamburg University in Germany (Yanko-Hombach et al. 2017). The grain-size analysis of the superficial (0–2 cm) sediment layer was performed by sieving and elutriation methods described in Logvinenko and Sergeeva (1986). Based on the results, the median diameter (Md) and coefficient of sorting

Fig. 2.3 Sketch of geological materials obtained in the Black Sea, the Sea of Azov, and their coasts and studied by foraminiferal analysis

2.2 Material and Methods

15

Fig. 2.4 Sketch of geological materials obtained in the Caspian Sea and its coast and studied by foraminiferal analysis

(So) were calculated for each sample. Cores were split and the working half was examined in the uppermost 2 cm of each 10-cm interval of the sediment column. Some of the gravity/piston cores were examined in 2-cm intervals (e.g., Core 1136 in Yanko-Hombach 2007a, here Fig. 8) to provide high-resolution bio- and ecostratigraphy. In total, approximately 32,000 samples from 1500 grabs, multicorers, box corers, 4300 gravity/piston cores, and

56 boreholes (up to 40 m in length) were investigated. It should be noted that the method of sampling is very important. Some authors recommend not to use grabs for sampling because “grab samplers create a strong bowwave when they touch the ground. Furthermore, grabs may only scrape the surface, distort the structure of the underlying sediments, and often do not close accurately. A large part of the sample is washed out when the grab is hoisted through the water

16

column, and an intact sediment surface is rarely preserved” (Schönfeld et al. 2012, 4). This is not always correct and cannot be recommended without exceptions. The yield of a sample very much depends on the grab construction, serviceability of the equipment, and the experience of the working team. In our case, all samples retained their undisturbed structure and were covered (in the delta front and partially in the prodelta) by an undisturbed thin (50% of a given assemblage) and accessory species. According to their ecological preferences, foraminifera were divided into oligohaline (1–5 psu), strictoeuryhaline (11–26 psu), polyhaline (18–26 psu), euryhaline (1–26 psu), shallow-dwelling (0–30 m), relatively deep-dwelling (31–70 m), and deep-dwelling (71–220 m) species (Yanko and Troitskaya 1987; Yanko 1989, 1990).

2.3 Statistical Treatment

Particular attention was paid to morphological deformities of foraminiferal tests as indicators of environmental stress. In fossil foraminiferal tests they have been noted by researchers since the last century (e.g., Carpenter 1856; Rhumbler 1911; Bogdanowicz 1952, 1960, 1971; Pflum and Frerichs 1976). In recent years, reports of deformities have become increasingly more common. Deformities have been linked to a number of environmental factors, such as (1) changes in temperature and reduced or elevated salinity, which also affects the size of foraminifera; (2) the lack or overabundance of food, which causes aberrant growth and affects the size of foraminifera; (3) the type of substrate, which affects the outline and shape of foraminiferal tests; (4) low dissolved oxygen content, which may create dwarfed, thin-walled, less ornamented and aberrant forms; (5) insufficient light, which may affect the size of foraminifera; (6) and pollution of marine environment (Yanko et al. 1999). The percentage of deformed foraminifera has been reported to increase dramatically in polluted areas (e.g., Lidz 1965) where foraminifera display a wide variety of deformities, including extreme compression, double apertures, twisted coiling, aberrant chamber shape, and protuberances. Bresler and Yanko (1995), using sulfaflavine fluorescence and chlortetracycline fluorescence, distinguished morphological deformities caused by mechanical damage from those caused by pathological morphogenesis. Noted that the UNESCO Practical Salinity Scale of 1978 (PSS78) is recommended for use in preference to parts per thousand (‰). The PSS defines salinity in terms of a conductivity ratio and so is dimensionless. On the PSS, normalmarine salinity is generally in the range 30–40, while brackish seas/waters have salinity in the range 0.5–12. Approximately equivalent values expressed in ppt are 30–50‰ (open sea) and 0.5–30‰ (brackish sea) (Mudie et al. 2011). We use psu (practical salinity unit) in this document. The collections of benthic foraminifera from the Black Sea, Caspian Sea, and Sea of Azov are stored in the Paleontological Museum of Odessa I.I. Mechnikov National University, Ukraine.

2.3

Statistical Treatment

To identify possible interrelations between foraminiferal characteristics and environmental parameters, cluster, correlation, factor, and multidimensional scaling analyses were applied using the “Statistica 10” package. For Q-mode cluster analysis, Ward’s method was used to optimize the minimum variance within clusters. Pearson’s correlation coefficient was used as a measure of similarity. Factor analysis of hydrological parameters was applied using the method of principal components followed by the varimax orthogonal-rotation procedure (varimax normalized). Correlations between parameters were considered as significant at p < 0.05 and 95% confidence limits.

17

For fossil assemblages, the residue obtained by CCl4 flotation was weighed; a portion of 0.02 g was taken for the analysis where the number of foraminiferal specimens of each species ( f ) was calculated using the equation f ¼ kn where n ¼ number of specimens for each low taxon in 0.02-g residue, g ¼ weight of the total residue, and k ¼ g/0.02. An 11-key medical counter was used to speed up the count which is especially convenient with low taxonomic diversity. For live (Bengal Rose-stained) foraminifera that have usually much lower quantity, quartering was not usually used, and all live specimens were counted. The total number of foraminifera (F) in the sample was calculated using quotation F ¼ f1 + f2 +. . . .fn. The concentration (c) of each species per sample was calculated using the equation: c ¼ f

100% F

The average concentration (C) of each lower taxon per area/stratigraphic unit, which defines a role of the species in the assemblage, was calculated using the following: C¼

c1 þ c2 þ c 3 . . . þ c n N

or

C ¼ F1

100 F

where N is the number of samples. The spatial occurrence (O) of a given species is calculated as a percentage from N in a given area/stratigraphic unit equivalent to 100%. Based on C and O, all species are divided into dominant and accessory ones. The dominant species with the highest C forms the core of an assemblage. Accessory species define its variability. If O  50%, the species are considered as widely distributed, 49–10% as often occurring, 9–1% as rare, and > 1% as single. If any species, independently from its quantity, occurs in the majority of samples, such a species is considered to be characteristic. Even if such species do not occur in high concentrations in assemblages, they can be still species indicators of a certain environment. For the evaluation of sedimentation rate and taphonomic considerations, the ratio between live and dead specimens was calculated using the following equations: P¼

F1  100% Fl þ Fd

e¼

f 1,2 . . . nl f 1,2 . . . nd

where Fl and is the total number of live and dead specimens of all species, respectively, P is the ratio between live and total specimens, and e is the proportion of live to dead specimens of all species for a given sample. Coefficient P indicates the sedimentation rate; a lower P indicates slower sedimentation (Phleger 1960). Coefficient e characterizes taphonomic conditions, with the lower e indicating better taphonomic conditions for a given species.

18

2 Study Area, Material, and Methods

An influence of environmental factors on foraminifera is estimated using the ratio between megalospheric and microspheric (d ) as well as between juvenile and adult (z) specimens of a given species using equations:

d¼

f meg f mic

z¼

f f ad1

If d > 1 and z > 1 or ¼ 0, the life conditions likely were not favorable and a species lived without reproduction. This is based on an assumption that, in extreme conditions, any organism tries to devote its energy to protecting itself and, therefore, prefers mitosis to meiosis (Effrussi and Farber 1975). To compare assemblages, the method of Cabioch (1979) was implemented using quotations:

1 K ¼ pffiffiffi 2

r ffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi  2  xa 2 ya þ x y

θ ¼ Arctg

xð y  aÞ yð x  aÞ

where x and y are the number of species in compared assemblages, a is the number of common species, K is the coefficient of similarity, and θ is the coefficient of enrichment. If K  0.707, the compared faunas are completely different; the smaller K is, the more similar are the faunas: weak similarity (K ¼ 0.707–0.507), medium similarity (K ¼ 0.506–0.207), and strong similarity (K < 0.207). If K ¼ 0, the assemblages are identical. For θ ¼ 60
, the fauna is strongly enriched with species; if θ ¼ 30
, the fauna is impoverished, and intermediate numbers show exchange of one fauna by another. The methods described enabled us to establish the lateral rows of live foraminifera (Yanko 1990) that reflect migration succession of foraminiferal assemblages (MSFA) after changeable environmental conditions. Fossil analogues of the MSFA, expressed as an alternation of foraminiferal assemblages and their ecological characteristics, provide reliable background for the high-resolution bio- and ecostratigraphy and paleoenvironmental reconstructions in an area under study.

2.4

Stratigraphic Techniques

The stratigraphic work follows the Russian subdivision of the Quaternary System (Zhamoida 2004) into the Eopleistocene (1.8–0.8 Ma), Neopleistocene (0.8–0.01 Ma), and Holocene (0.01–0.0 Ma). The resulting high-resolution stratigraphy and geochronology of the Late Quaternary (Neopleistocene and Holocene) sediments are based on the combination of bioand ecostratigraphic criteria supplemented by absolute dating. Biostratigraphic criterion includes precise taxonomic

analysis of benthic foraminifera to trace species evolution and to discover species indicators for certain time intervals. Due to the shortness of the studied time interval, the application of this criterion is limited and so is supplemented by ecostratigraphic criteria. Ecostratigraphy is the biostratigraphic application of ecological and paleoecological principles to develop an understanding of the global external-forcing agents that drive ecological change. The ecostratigraphy of the Black Sea addresses biotic responses to isolation from and connection to the neighboring Sea of Marmara and Caspian Sea and to related sea-level changes and salinity oscillations. This ecostratigraphic technique is based largely on the alternation of foraminiferal assemblages and their ecological characteristics in geological sections, supported by 14C and palynological assays. An increase in the number of Mediterranean immigrants, especially strictoeuryhaline and polyhaline species, in sediment sequences indicates an increase of Mediterranean influence and salinity and vice versa. The complete replacement of Mediterranean immigrants by oligohaline Caspian species shows the separation between the Black Sea and Mediterranean, followed by the desalination of the Black Sea. This conclusion is based on a generally accepted observation, fully supported by our ecological study (Yanko 1989, 1990), that foraminifera are not well adapted to freshwater environments (Sen Gupta 1999).

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19 Reilinger RE, McClusky SC, Souter BJ (1997) Preliminary estimates of plate convergence in the Caucasus collision zone from GPS measurements. Geophy Res Lett 24:1815–1818 Rhumbler L (1911) Die Foraminiferen (Thalamophoren) der Plankton Expedition, Pt. 1, Die allgemeinen Organizationsverhaltnisse der Foraminiferen: Lipsius & Tisher, Kiel und Leipzig, 331 pp Robertson AHF, Ustaömer T, Pickett EA, Collins AS, Andrew T, Dixon JE (2004) Testing models of Late-Palaeozoic-Early Mesozoicorogeny in Western Turkey: support for an evolving open-Tethys model. J Geol Soc 161:501–511 Schönfeld J, Alve E, Geslin E, Jorissen F, Korsun S, Spezzaferri S, Members of the FOBIMO group (2012) The FOBIMO (FOraminiferal BIo-MOnitoring) initiative – towards a standardized protocol for soft-bottom benthic foraminiferal monitoring studies. Mar Micropaleontol 94–95:1–13 Semenenko VN, Kovalyukh NN (1973) Absolutniy vozrast verchnechetvertichnykh otlozeniy azovo- chernomorskogo basseyna po dannym radiouglerodnogo analiza (Radiocarbon age of Upper Quaternary sediments of the Azov-Black Sea basin). Geologicheskiy Zhurnal 33(6):91–97. (in Russian) Sen Gupta BK (1999) Foraminifera in marginal marine environments. In: Sen Gupta BK (ed) Modern Foraminifera. Kluwer Academic Publishers, Dordrecht, pp 141–160 Smith DE, Kolenkiewics R, Robbins PJW, Dunn J, Torrence MH (1994) Horizontal crustal motion in the central and eastern Mediterranean inferred from satellite laser ranging measurements. Geophys Res Lett 21:1979–1982 Svitoch AA, Yanina TA, Menabde IV (1992) Paleogeografiya pozdnego pleystotsena Ponto-Kaspiya (Paleogeography of the Late Pleistocene of the Ponto-Caspian). Vestnik Mosk. un-ta. Ser. 5. Geografiya 6(68–76) (in Russian) Tari E, Sahin M, Barka A et al (2000) Active tectonics of the Black Sea with GPS. Earth Planets Space 52:747–751 Voskoboynikov VM, Krakovskiy BI, Konnikov EG, Yanko V (1985) Litologiya chetvertichnykh otlozheniy: Tekstury donnykh otlozheniy (Lithology of Quaternary sediments: textures of the bottom sediments). In: Shnyukov EF (ed) Geologiya shelfa USSR: Litologiya. Naukova Dumka, Kiev, pp 93–130. (in Russian) Yanko V (1979) Stratigrafiya i geochronologoya donnykh otlozheniy: Stratigraficheskie kompleksy bentocnykh foraminifer. Obschaya stratigraficheskaya skhema (Stratigraphy and geochronology of the bottom sediments: stratigraphic complexes of benthic foraminifera. General stratigraphic scheme). In: Malovitsky Y (ed) Geology and hydrology of the Western part of the Black Sea. Ac. Sc. Bulg, Sofia, pp 82–95. (in Russian) Yanko V (1989) Quaternary Foraminifera of the Ponto-Caspian Region (The Black Sea, the Sea of Azov, the Caspian Sea and the Aral Sea): taxonomy, biostratigraphy, history, ecology. Doctoral thesis. Moscow State University, two volumes, 1000 pp (in Russian) Yanko V (1990) Stratigraphy and paleogeography of marine Pleistocene and Holocene Deposits of the Southern Seas of the USSR. Mem Soc Geol Ital 44:167–187 Yanko V, Gramova L (1990) Stratigraphy of the quaternary sediments of the Caucasian shelf and continental slope of the Black Sea on microfauna (foraminifera and ostracoda). J Sov Geol 2:60–72. (in Russian) Yanko V, Frolov V, Motnenko I (1990) Foraminifery i litologiya karangatskogo gorizonta (Antropogen Kerchenskogo poluostrova) Foraminifera and lithology of the Karangatian horizon (Quaternary of the Kerchenian Peninsula). Bull Mos Soc Nat Geol Branch 65:83– 97. (in Russian) Yanko VV, Kondariuk TO (2020) Origin and taxonomy of the Neopleistocene-Holocene Ponto- Caspian benthic foraminifera. Geologichnyy zhurnal 1:17–33

20 Yanko V, Troitskaya T (1987) Pozdnechetvertichnye foramifery Chernogo morya (Late Quaternary Foraminifera of the Black Sea). Nauka, Moscow (in Russian) Yanko et al. 1995 Yanko V, Ahmad M, Kaminski M (1998) Morphological deformities of benthic foraminiferal tests in response to pollution by heavy metals: implications for pollution monitoring. J Foraminiferal Res 28(3): 177–200 Yanko V, Arnold A, Parker W (1999) The effect of marine pollution on benthic foraminifera. In: Sen Gupta BK (ed) Modern Foraminifera. Kluwer Academic Publishers, Dordrecht, pp 217–238 Yanko VV, Kondariuk TO, Kadurin SV (2019) Istoriya heolohichnoho rozvytku pivnichno- zakhidnoho shel’fu Chornoho morya v pizn’omu neopleystotsena-holotseni (History of geological development of the north-western shelf of the Black Sea in the late NeoPleistocene-Holocene). Zbirnyk naukovykh prats0 Instytutu geolohichnykh nauk NAN Ukrayiny 12:123–136 (in Ukranian) Yanko-Hombach V (2007a) Controversy over Noah’s flood in the Black Sea: geological and foraminiferal evidence from the shelf. In: Yanko-Hombach V, Gilbert AS, Panin N, Dolukhanov PM (eds) The Black Sea flood question: changes in coastline, climate and human settlement. Springer, Dordrecht, pp 149–204 Yanko-Hombach V (2007b) Table of radiocarbon dates from USSR and non-USSR sources. In: Yanko-Hombach V, Gilbert AS, Panin N, Dolukhanov PM (eds) The Black Sea flood question: changes in coastline, climate and human settlement. Springer, Dordrecht, pp 861–877 Yanko-Hombach V, Yanina TA, Motnenko I (2013) Neopleistocene stratigraphy of the Ponto-Caspian Corridors. In: Gilbert A and

2 Study Area, Material, and Methods Yanko-Hombach V (eds) Proceedings of IGCP 610 First Plenary Meeting and Field Trip. 12th–19th October 2013, Tbilisi, Georgia, pp 170–176 Yanko-Hombach V, Mudie PJ, Kadurin S, Larchenkov E (2014) Holocene marine transgression in the Black Sea: new evidence from the northwestern Black Sea shelf. Quat Int 345:100–118 Yanko-Hombach V, Schnyukov E, Pasynkov A, Sorokin V, Kuprin P, Maslakov N, Motnenko I, Smyntyna O (2017) Late PleistoceneHolocene environmental factors defining the Azov- Black Sea basin, and the identification of potential sample areas for seabed prehistoric site prospecting and landscape exploration on the Black Sea continental shelf. In: Flemming NC, Harff J, Moura D, Burgess A, Bailey GN (eds) Submerged landscapes of the European continental shelf: quaternary Paleoenvironments. WileyBlackwell, Chichester, pp 431–478 Yilmaz Y (1997) Geology of Western Anatolia. Active tectonics of northwestern Anatolia. In: Schindler C, Pfister M, Aksoy A (eds) Active tectonics of Northwestern Anatolia: the MARMARA polyproject: a multidisciplinary approach by space-geodesy, geology, hydrogeology, Geothermics and seismology. Vdf, Hochschulverlag AG an der ETH, Zurich, pp 210 Zhamoida AI (2004) Problems related to the international (standard) stratigraphic scale and its perfection. Stratigr Geol Correl 12:321– 330. (Translated from Stratigrafiya. Geologicheskaya Korrelyatsiya 12:3c–13) Zubov NN (1956) Osnovy izuchenya prolivov Mirovogo okeana (Fundamentals of the studies of World Ocean straits). Geographgiz, Moscow (in Russian)

3

Taxonomic Classification of Foraminifera

Abstract

The higher taxonomic classifications proposed by Saidova (O sovremennom sostoyanii sistemy nadvidovykh taksonov kaynozoyskikh bentosnykh foraminifer (On the current state of the system of supraspecific taxa of Cenozoic benthic foraminifers). AN SSSR, Moscow, 720 pp (in Russian), 1981), Mikhalevich (Micropaleontology 59:493–527, 2013), and Pawlowski et al. (Mar Micropaleontol 100:1–10, 2013), in conjunction with classifications suggested by the World Register of Marine Species (WoRMS), were used to provide an authoritative and comprehensive list of names of marine organisms, including information on synonymy. Unfortunately, in many cases the WoRMS classifications did not fit with the author’s concept for the higher classification of the Ponto-Caspian Quaternary foraminifera; among the editors of WoRMS, there is no one with specific expertise in the fauna of this unique geographic region. As a consequence of the specific geographic position and its long semi- or complete isolation from the World Ocean, the Ponto-Caspian biota, including foraminifera, is quite specific. Also, purely taxonomic work was not the main goal of this investigation of the Ponto-Caspian Quaternary foraminifera. This book focuses on applied components enabling the use of foraminifera for stratigraphy and paleogeographic reconstructions, as well as for environmental monitoring. A polytypic concept of species recognizes that any species is composed of many allopatric populations that can differ to a certain degree from each other. If individuals from these populations differ in diagnostic morphological characteristics and are geographically and temporally isolated, they can be considered as subspecies. The identification of species is based on the set of criteria for zoological systematics developed by Mayr et al. (Metody i printsipy zoologicheskoy sistematiki (Methods and principles of zoological systematic). Izd-vo insstr.lit., Moscow, 49 pp (in Russian), 1953) and Mayr (Printsipy

zoologicheskoy sistematiki (Principles of zoological systematic). Mir, Moscow, 454 pp (in Russian), 1971) for the entire fauna as a whole and refined by Fursenko (Vvedenie v izuchenie foraminifer (Introduction to the study of foraminifera). Trudy Instituta Geologii i Geofiziki 391. Nauka, Novosibirsk, 242 pp (in Russian), 1978) for foraminifera. His concept is based on the assumption that only combinations of morphological (or comparative morphological), geographical, ecological, geochronological, and discriminative criteria can provide correct identification of a species. All taxa of the Quaternary foraminifera of the PontoCaspian are classified as belonging to Kingdom Protoctista, Class Foraminifera d’Orbigny, 1826. The classification of the Quaternary foraminifera of the Ponto-Caspian includes 180 and 7 lower taxa of benthic and planktonic foraminifera, respectively. Nineteen species are given in open nomenclature. Among benthic foraminifera, 30 are agglutinated; the remaining 150 lower taxa are characterized by calcium carbonate tests. Keywords

Phenetic species concept · Criteria for species identification · Principles of taxonomic classification

3.1

General Remarks

Application of foraminifera to high-resolution stratigraphy and paleoenvironmental reconstructions requires precise taxonomic identification of lower taxa and their further integration into the general system of foraminifera. As for today, foraminiferal taxonomy relies on a phenetic species concept (e.g., Loeblich and Tappan 1988) that classifies foraminifera by their morphological similarities, which can be also caused by environmental variations. As a consequence, it is often difficult to make a decision whether morphological variations

# Springer Nature Switzerland AG 2022 V. Yanko, Quaternary Foraminifera of the Caspian-Black Sea-Mediterranean Corridors: Volume 1, https://doi.org/10.1007/978-3-031-12374-0_3

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are genotypic or ecophenotypic. Classifying foraminifera by morphological criteria may lead to appearance of species with wide geographic and stratigraphic distribution, e.g., Ammonia beccarii described by Linnaeus (1758). Another concept is needed to resolve taxonomic problems. Molecular concepts (e.g., Holzmann 2000) opens new perspectives for the taxonomy of recent foraminifera because molecular systematics enables relatively clear definitions of existing taxa and investigation of hidden genetic subdivisions (sibling species). The main disadvantage of this concept is its applicability to living organisms only, as they comprise only a small proportion of the total number of known species. Thus, molecular concepts are not useful for stratigraphers who are dealing largely with fossil species that often have no analogues among recent foraminifera. For the Quaternary, the molecular classification is useful, though a variety of limitations must be recognized. At the current stages of development, no sufficient data bank is yet available, some groups are very resistant to genomic techniques, and the wide distributions of cryptobiotic propagules result in the appearance of sequences far outside their viable ranges. Thus, morphological criteria for identification remain essential both in traditional studies and for comparisons with environmental applications of molecular classifications.

3.2

Principles of Taxonomic Classification

The author has carefully analyzed the higher taxonomic classifications suggested by Saidova (1981), Mikhalevich (2013), and Pawlowski et al. (2013) and have tried to adopt those schemes to classifications suggested by the World Registered Marine Species (WoRMS) (WoRMS Editorial Board 2021), the aim of which is to provide an authoritative and comprehensive list of names of marine organisms, including information on synonymy. Unfortunately, in many cases the WoRMS classifications did not fit with the author’s concept for a higher classification of the PontoCaspian Quaternary foraminifera. As a consequence of the specific geographic position of the Ponto-Caspian and its long semi- or complete isolation from the World Ocean, its biota, including foraminifera, is quite specific. Unfortunately, among the editors of WoRMS, there is no one with specific expertise in the fauna of the unique Ponto-Caspian geographic region (see map “Our editors around the world” at http://www.marinespecies.org/about.php). Also, purely taxonomic work was not the main goal of this investigation of the Ponto-Caspian Quaternary foraminifera. This book focuses on applied components enabling use of foraminifera for stratigraphy and paleogeographic reconstructions (Chap. 6), as well as for environmental monitoring (Chap. 7).

Taxonomic Classification of Foraminifera

The basic taxonomic unit of the system for any group of organisms is the “species.” Therefore, the primary task of a researcher dealing with issues of taxonomy is the accurate understanding of the species characteristics and the establishment of criteria to consistently identify individuals at all stages of ontogeny. This work adopts a polytypic concept of the species, that is, that any species is composed of many allopatric populations that can differ to a certain degree from each other. If individuals from these populations differ in diagnostic morphological characteristics and are geographically and temporally isolated, they are considered as subspecies. The identification of species is based on the set of criteria for zoological systematics developed by Mayr et al. (1956) and Mayr (1971) for the entire fauna as a whole and refined by Fursenko (1978) for foraminifera. His concept is based on the assumption that only a combination of morphological (or comparative morphological), geographical, ecological, geochronological, and discriminative criteria can provide correct identification of a species. The morphological criteria describe external and internal features of foraminiferal tests. The geographical, ecological, and geochronological criteria associate species with certain zoogeographical areoles and environmental conditions required for their existence and adaptation at certain chronological time interval. The discriminative criteria specify parameters useful to distinguish foraminiferal species from each other. The above criteria correspond to the main philosophical categories of time (geochronological), space (geographical), evolution (morphological), and metabolism (ecological) of organisms. Being isolated from each other inevitably leads to the emergence of distinct features. In our material they are represented by a group of species previously united under the name A. beccarii, as will be discussed below. Using this concept, this work classifies and uses stratigraphy for some taxa (e.g., Ammonia spp., Yanko 1990a) that were considered by some authors as useless for biostratigraphic purposes because of difficulties and uncertainties in their identification (e.g., Holzmann 2000). The main characteristics are morphological, geochronological, geographic, and ecological, primarily morphological criteria. All taxa of the Quaternary foraminifera of the PontoCaspian are classified as belonging to Kingdom Protoctista, Class Foraminifera d’Orbigny, 1826, Subclass Sarcodina Eichwald, 1830 (Table 3.1). Classification of the Quaternary foraminifera of the PontoCaspian includes altogether 180 benthic and 7 planktonic species and subspecies. Among benthic foraminifera, 30 are agglutinated, and the remaining lower taxa are characterized by calcium carbonate tests. The classification of Loeblich and Tappan (1988) provides the basis for the generic classification used in this study. The taxonomic

3.2 Principles of Taxonomic Classification

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Table 3.1 Taxonomic classification of the Late Quaternary foraminifera of the Ponto-Caspian Region Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Order Astrorhizida Lankester, 1885 Family Astrorhizidae Brady, 1881 Genus Astrammina Rhumbler in Wiesner, 1931 Astrammina sphaerica (Brady, 1871)*** Genus Bathysiphon Sars, 1872 Bathysiphon hirudinea (Herron-Allen and Earland, 1932)*** Family Psammosphaeridae Haeckel, 1894 Subfamily Psammosphaerinae Haeckel, 1894 Genus Psammosphaera Schultze, 1875 Psammmosphaera sp. Family Saccaminidae Brady, 1884 Subfamily Saccamininae Brady, 1884 Genus Saccammina Sars, in Carpenter, 1869 Saccammina sp.** Genus Proteonella Lukina, 1969 Proteonella atlantica (Cushman), 1944 Genus Ovammina Dahlgren, 1962 Ovammina leptoderma Mayer** Genus Hemisphaerammina Loeblich and Tappan, 1957 Hemisphaerammina sp.** Order Ammodiscida Fursenko, 1958 Family Ammodiscidae Reuss, 1862 Genus Glomospira Rzehak, 1885 Glomospira glomerata Hoglund, 1947* Glomospira gordialis (Jones and Parker, 1860)*** Family Discamminidae Mikhalevich, 1980 Genus Discammina Lacroix, 1932 Discammina imperspica Yanko, 1974 Family Lituolidae De Blainville, 1825 Genus Lituola Lamarck, 1804 Lituola nautilus Brady * Genus Labrospira Höglund, 1947 Labrospira sp.* Genus Ammobaculites Cushman, 1910 Ammobaculites exiguus contractus Mayer, 1972 Ammobaculites ponticus Mikhalevich, 1968 Genus Haplophragmoides Cushman, 1910 Haplophragmoides tenuicutis (Mayer), 1972 Genus Ammoscalaria Höglund, 1947 Ammoscalaria verae Mayer, 1968 Ammoscalaria sp., in Yanko, 1989 Genus Haplophragmium (Reussina) Gryzbowski, 1895 Haplophragmium maync Loeblich and Tappan, 1952* Family Textulariidae Ehrenberg, 1838 Genus Textularia Defrance, 1824 Textularia deltoidea Reuss* Textularia conica d’Orbigny, 1839 (¼ Textularia sp. in Yanko, 1989) Order Schlumbergerinida Mikhalevich, 1980 Family Miliamminidae Saidova, 1981 Genus Birsteiniolla Mayer, 1974 Birsteiniolla macrostoma Yankovskaya and Mikhalevich, 1972*** (continued)

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Taxonomic Classification of Foraminifera

Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Genus Miliammina (Brady, 1870) Miliammina fusca (Brady, 1870) Miliammina groenlandica Cushman**** Family Siphonapertinae Saidova, 1975 Genus Siphonaperta Vella, 1957 Siphonaperta agglutinans (d’Orbigny, 1839) Order Ataxophragmiida Fursenko, 1958 Family Trochamminidae Schwager, 1877 Genus Entzia von Daday, 1884 Entzia polystoma caspica (Mayer), 1968** Entzia polystoma dacica (Tufescu), 1973**** Genus Rotaliammina Cushman, 1924 Rotaliammina intermedia Rhumbler 1938* Rotaliammina ochracea (Williamson), 1858 Family Verneulinidae Cushman, 1911 Genus Spiroplectinata Cushman, 1927 Spiroplectinata perexilis (Mayer), 1968** Family Eggerellidae Cushman, 1937 Genus Eggerelloides Cushman, 1933 Eggerelloides scaber (Williamson, 1858) Order Miliolida Lankester, 1885 Family Cornuspiridae Shultze, 1854 Genus Cornuspira Shultze, 1854 Cornuspira minuscula (Mayer, 1968) Cornuspira planorbis Shultze, 1854 Superfamily Milioloidea Ehrenberg, 1839 Family Miliolidae Ehrenberg, 1839 Subfamily Miliolinae Ehrenberg, 1839 Genus Ammomassilina Cushman, 1933 Massilina alveoliniformis (Millett, 1898)* Subfamily Quinqueloculininae Cushman, 1917 Genus Quinqueloculina d’Orbigny, 1826 Quinqueloculina angulata (Williamson, 1858) Quinqueloculina bicornis (Walker and Jacob), 1798 Quinqueloculina consobrina (d’Orbigny), 1846 Quinqueloculina curvula Yanko, 1989 Quinqueloculina delicatula Bogdanowicz, 1952 Quinqueloculina inflata (d’Orbigny), 1826 Quinqueloculina laevigata (d’Orbigny), 1826 Quinqueloculina lamarckiana d’Orbigny, 1839 Quinqueloculina lata Terquem, 1878 Quinqueloculina milletti (Wiesner), 1912 Quinqueloculina oblonga (Montagu), 1803 Quinqueloculina reussi (Bogdanowicz), 1947 Quinqueloculina seminulum (Linne), 1767 Quinqueloculina venusta Karrer, 1868 Quinqueloculina vulgaris d’Orbigny, 1826 Quinqueloculina ex gr. gracilis Karrer, in Yanko, 1989 Genus Dentostomina Carman, 1933 Dentostomina bermudiana Carmann, 1933* Genus Massilina Schlumberger, 1893 (continued)

3.2 Principles of Taxonomic Classification

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Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Massilina inaequalis (d’Orbigny), 1839 Massilina secans (d’Orbigny), 1826 Genus Lachlanella Vella, 1957 Lachlanella sp. Genus Pateoris Loeblich and Tappan, 1953 Pateoris dilatatus (d’Orbigny), 1838 Pateoris sp., in Yanko, 1989 Genus Pyrgo Defrance, 1824 Pyrgo elongata (d’Orbigny), 1826 Pyrgo fisheri (Schlumberger), 1891 Subfamily Miliolinellinae Vella, 1957 Genus Miliolinella Wiesner, 1931 Miliolinella circularis (Bornemann), 1865 Miliolinella elongata Kruit, 1955 ¼ Miliolinella sp., in Yanko, 1989 Miliolinella risilla Mayer, 1972* Miliolinella selene (Karrer), 1868 Miliolinella subrotunda (Montagu), 1803 Subfamily Tubinelinae Rhumbler, 1906 Genus Articulina (d’Orbigny), 1826 Articulina tubulosa (Seguenza, 1862) Articulina ex gr. tenella Eichwald, 1830 Articulina sp., in Yanko, 1989 Subfamily Triloculininae Bogdanowicz, 1981 Genus Triloculina d’Orbigny, 1826 Triloculina (?) angustioris (Bogdanowicz), 1952 Triloculina marioni Schlumberger, 1883 Triloculina sp. 2, in Yanko, 1989 Genus Sigmella Azbel & Mikhalevich, 1983 Sigmella distorta (Phleger and Parker), 1951 Sigmella tenuis (Czjzek), 1848 Sigmella sp., in Yanko, 1989 Genus Cycloforina Łuczkowska, 1972 Cycloforina? sp. Order Lagenida Fursenko, 1958 Superfamily Nodosariidae Ehrenberg, 1838 Family Nodosariidae Ehrenberg, 1838 Subfamily Nodosariinae Ehrenberg, 1838 Genus Orthomorphina Stainforth, 1952 Orthomorphina calomorpha (Reuss), 1866 Orthomorphina drammenensis Feyling-Hanssen, 1964 Orthomorphina filiformis (d’Orbigny), 1826 Subfamily Lageninae Brady, 1881 Genus Lagena Walker and Jacob in Kanmacher, 1798 Lagena quadrilatera quadrilatera Earland, 1934 Lagena semistriata (Williamson), 1858 Lagena striata (d’Orbigny), 1839 Lagena vulgaris Williamson, 1858 Lagena sp., in Yanko, 1989 Superfamily Glandulinoidea Yanko, 1989 Family Glandulinidae Reuss, 1860 Subfamily Glandulininae Reuss, 1860 (continued)

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3

Taxonomic Classification of Foraminifera

Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Genus Glandulina d’Orbigny, in de la Sagra, 1839 Glandulina sp., in Yanko and Troitskaya, 1987 Family Polymorphinidae d’Orbigny, 1839 Genus Guttulina d’Orbigny, 1839 Guttulina lactea (Walker and Jacob), 1798 Subfamily Laryngosigminae Saidova, 1981 Genus Esosyrinx Loeblich and Tappan, 1953 Esosyrinx jatzkoi Yanko, 1974 Esosyrinx praelongus (Terquem), 1878 Esosyrinx undulosus (Terquem), 1878 Esosyrinx sp., in Yanko, 1989 Genus Laryngosigma Loeblich and Tappan, 1953 Laryngosigma williamsoni (Terquem), 1878 Subfamily Entolingulinae Saidova, 1981 Genus Entolingulina Loeblich and Tappan, 1961 Entolingulina deplanata Yanko, 1974 Subfamily Oolininae Loeblich and Tappan, 1961 Genus Oolina d’Orbigny, 1839 Oolina squamosa (Montagu), 1803 Oolina sp., in Yanko, 1989 Genus Fissurina Reuss, 1850 Fissurina fabaria Troitskaya, 1987 Fissurina fragilis Troitskaya, 1987 Fissurina lineata (Williamson), 1858 Fissurina lucida (Williamson), 1858 Fissurina nummiformis (Büchner), 1940 Fissurina porrecta Troitskaya, 1987 Fissurina solida Seguenza, 1862 Fissurina tamanica Yanko, 1989 Fissurina sp., in Yanko, 1989 Genus Parafissurina Parr, 1947 Parafissurina aventricosa McCulloch, 1968 Parafissurina dzemetinica Yanko, 1974 Parafissurina ex gr. lateralis Cushman, in Yanko and Troitskaya, 1987 Order Rotaliida Delage and Herouard, 1896 Superfamily Discorboidea Ehrenberg, 1838 Family Discorbiidae Ehrenberg, 1838 Subfamily Discorbiinae Ehrenberg, 1838 Genus Discorbis Lamarck, 1804 Discorbis bertheloti (d’Orbigny), 1839 Discorbis vilardeboana (d’Orbigny), 1839 Discorbis sp. Genus Gavelinopsis Hofker, 1951 Gavelinopsis sp., in Yanko, 1989 Subfamily Rosaliinae Reuss, 1963 Genus Rosalina d’Orbigny, 1826 Rosalina catesbyana d’Orbigny, 1839 Rosalina sp., in Yanko, 1989 Family Glabratellidae Loeblich and Tappan, 1964 Genus Heronallenia Chapman and Parr, 1931 Heronallenia chasteri (Heron-Allen and Earland), 1913 (continued)

3.2 Principles of Taxonomic Classification

27

Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Superfamily Anomalinoidea Cushman, 1927 Family Cibicidae Cushman, 1927 Genus Cibicides Montfort, 1808 Cibicides dispars (d’Orbigny), 1838 Cibicides lobatulus (Walker and Jacob), 1798 Cibicides mckannai Galloway and Wiesler, 1927 Family Planorbulinidae Schwager, 1877 Genus Planorbulina d’Orbigny, 1826 Planorbulinella mediterranensis d’Orbigny, 1826 Family Acervulinidae Schultze, 1854 Genus Acervulina Schultze, 1854 Acervulina inhaerens Schulze, 1854 Superfamily Nonionoidae Schultze, 1854 Family Nonionidae Schultze, 1854 Genus Nonion Montfort, 1808 Nonion matagordanus Kornfeld, in Cushman, 1939 Nonion pauciloculum Cushman, 1944 Genus Florilus Montfort, 1808 Florilus trochospiralis Mayer, 1968 Florilus cf. atlanticum (Cushman, 1947) Family Trichochyalidae Saidova, 1981 Genus Trichochyalus Loeblich and Tappan, 1953 Trichochyalus aguajoi (Bermudez), 1935 Superfamily Rotalioidea Ehrenberg, 1839 Family Rotaliidae Ehrenberg, 1839 Genus Rotalia Lamarck, 1804 Rotalia calcar (d’Orbigny in Deshayes, 1830)* Family Ammoniidae Saidova, 1981 Genus Ammonia Brünnich, 1772 Ammonia agoiensis Yanko, 1990 Ammonia ammoniformis (d’Orbigny), 1826 Ammonia beccarii (Linne), 1758 Ammonia caspica Stschedrina, 1975 Ammonia caucasica Yanko, 1990 Ammonia compacta Hofker, 1969 Ammonia novoeuxinica Yanko, 1979 Ammonia parasovica Stchedrina and Mayer, 1975 Ammonia parkinsoniana (d’Orbigny), 1839 Ammonia tepida (Cushman), 1928 Family Canaliferidae Krasheninnikov, 1960 Genus Canalifera Krasheninnikov, 1960 Canalifera earlandi (Cushman), 1936 Canalifera nigarensis (Cushman), 1939 Canalifera oweniana (d’Orbigny), 1839 Canalifera parkerae Yanko, 1974 Canalifera punctata (Terquem), 1878 Canalifera ex gr. verriculata (Brady), in Yanko and Troitskaya, 1987 Family Elphidiidae Galloway, 1933 Subfamily Haynesininae Yanko, subfam.n. Genus Porosononion Putrja, 1958 Porosononion martkobi martkobi Bogdanowicz, 1947 (continued)

28

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Taxonomic Classification of Foraminifera

Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Porosononion martkobi ponticus Yanko, 1989 Porosononion martkobi tschaudicus Yanko, 1989 Porosononion subgranosus mediterranicus Yanko, 1989 Porosononion subgranosus pshadicus Yanko, 1989 Porosononion submartkobi Yanko, 1989 Genus Aubignyna Margerel, 1970 Aubignyna mariei Margarel, 1970 Aubignyna perlucida (Herron-Allen and Earland), 1913 Aubignyna suckumiensis Yanko, 1989 Genus Haynesina Banner and Culver, 1978 Haynesina anglica (Murray), 1965 Haynesina eltigenica Yanko, 1989 Haynesina ex gr. germanica (Ehrenberg, 1840) Subfamily Elphidiinae Galloway, 1933 Genus Elphidium Montfort, 1808 Elphidium aculeatum (d’Orbigny), 1846 Elphidium caspicum azovicum Yanko, 1989 Elphidium caspicum caspicum Yanko, 1989 Elphidium caspicum karadenizum Yanko, 1989 Elphidium caspicum uzunlarum Yanko, 1989 Elphidium incertum tuberculatum (Williamson, 1858) Elphidium josephinum (d’Orbigny), 1846 Elphidium margaritaceum Cushman, 1939 Elphidium ponticum Dolgopolskaja and Pauli, 1931 Elphidium shochinae Mayer, 1968 Elphidium umbilicatulum (Williamson), 1858 Elphidium sp., in Yanko, 1989 Genus Cribroelphidium Cushman and Bronnimann, 1948 Cribroelphidium percursum Yanko, 1974 Cribroelphidium poeyanum (d’Orbigny), 1839 Cribroelphidium translucens (Natland), 1938 Cribroelphidium troitskayae Yanko, 1989 Genus Mayerella Yanko 1987 Mayerella aralica Yanko, 1987 Mayerella brotzkajae (Mayer), 1968 Mayerella kolkhidica Yanko, 1989 Superfamily Globogerinoidea Carpenter, 1862 Family Globigerinidae Carpenter, 1862 Genus Globigerina d’Orbigny, 1826 Globigerina bulloides d’Orbigny, 1826 Globigerina praebulloides Blow, 1959 Globigerina quinqueloba Natland, 1938 Genus Globoquadrina Finlay, 1947 Globoquadrina dutertrei (d’Orbigny, 1839) Globoquadrina hexagona (Natland, 1938) Genus Globorotalia Cushman, 1927 Globorotalia pumpilio Parker, 1962 Globorotalia crassaformis (Galloway and Wissler, 1927) Order Buliminida Fursenko, 1958 Family Buliminidae Jones, in Griffith et al., 1875 Subfamily Buliminidae Jones, 1875 (continued)

3.3 Agglutinated Taxa

29

Table 3.1 (continued) Kingdom Protoctista Class Foraminifera D’Orbigny, 1826 Subclass Sarcodina Eichwald, 1830 Genus Bulimina d’Orbigny, 1826 Bulimina aculeata d’Orbigny, 1826 Bulimina elongata d’Orbigny, 1826 Globobulimina affinis (d’Orbigny, 1839) Family Uvigerinidae Haeckel, 1894 Genus Trifarina Cushman, 1923 Trifarina angulosa (Williamson), 1858 Family Bolivinidae Cushman, 1927 Genus Bolivina d’Orbigny, 1839 Bolivina doniezi Cushman and Wickenden, 1929 Bolivina pseudoplicata Heron-Allen and Earland, 1930 Bolivina variabilis (Williamson), 1858 Bolivina ex gr. dilatata Reuss, 1850 Bolivina sp. Genus Brizalina Costa, 1856 Brizalina spathulata (Williamson), 1858 Brizalina striatula (Cushman), 1922 Brizalina ex gr. danvillensis (Howe and Wallace), 1932, in Yanko and Troiskaya, 1987 Note. Asterisks mark the species, images, and, in some cases, descriptions that are provided in earlier publications on the Black and Caspian Seas: * Velkanova 1981; ** Mayer 1968, 1972, 1976; *** Mikhalevich 1968; **** Tufescu 1973, 1974. These species were not found in the material described in this book and are not illustrated herein. Most of the others are supplemented by images and descriptions herein

position of the Foraminifera has varied since their recognition as Protozoa (Protista) by Schultze in 1854, who referred to the order Foraminiferida. Loeblich and Tappan (1992) reranked Foraminifera as a class as it is now commonly regarded by paleontologists. Tests of agglutinated foraminifera are represented by four main types that differ in their morphology and internal structure and are placed into four orders: Astrorhizida, Ammodiscida, Schlumbergerinida, and Ataxophragmiida. All their representatives are characterized by different foreign particles and cement as well as the ratio between them. Secretory calcium carbonate foraminifera also belong to four orders: (1) Miliolida (high Mg calcite, porcellanous, three layers, non-lamellar, imperforate, tangle-shaped tests). (2) Lagenida (low Mg calcite, monolamellar, hyaline radial, single or multiple chambers, uniserial or planispiral), (3) Rotaliida (low Mg calcite, bilamellar and bilamellar–rotaloid, hyaline, perforated, multichambered), and (4) Buliminida (low Mg calcite, bilamellar, perforated, multichambered, biserial, triserial).

3.3

Agglutinated Taxa

To classify agglutinated foraminifera from the Ponto-Caspian geographic region, special attention was paid to the composition and quantitative ratio of agglutinated particles and cement in their tests. In calcareous foraminifera special

attention was given to the microstructure (granular, radial), layer (monolamellar, bilamellar, bilamellar–rotaliid), porosity (size, shape, pore density), structure of the septal sutures, and canal system (e.g., in rotaliids). The agglutinated foraminiferal genera Miliammina and Siphonaperta with quinqueloculine chamber arrangement were moved from the family Rzehakinidae (Orlov 1959) or suborder Textulariina (Loeblich and Tappan 1964, 1981, 1984), where they were located based on siliceous content of their tests, to the family Shlumbergerinidae described by Mikhalevich (1980, 1983). The content of agglutinated particles and cement are not permanent characters of these species; they vary significantly on different substrates. For example, the tests for Eg. scaber and Mi. fusca collected from the quartz substrate are dominated by SiO2 and are not soluble in H2S. In contrast, those collected from the carbonate substrate have an elevated proportion of carbonate particles soluble in H2S. In our opinion, a false impression is created wherein the composition of cement is not a result of the functional characteristics of the cytoplasm, but depends on nutrition. Rather, apparently as Saidova (1981) noted, forms that build their shells from agglutinated material extracted from the soil are “soil eaters” and receive all the necessary biochemical elements by assimilating the soil. The chamber arrangement seems to be the most stable feature of agglutinated species, as was stated by Mikhalevich

30

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Taxonomic Classification of Foraminifera

(1980, 1983). In general, agglutinated foraminifera are rare among both living and fossil assemblages and have a sporadic distribution in geological sequences.

3.4

Taxonomic Classification of the Order Miliolida

The order Miliolida in the Ponto-Caspian geographic region includes 42 lower taxa with a tangle-shaped structure of tests that have a porcelain-like, three-layer, non-lamellar-type wall with a thin organic matrix. At magnification of 5500 times in the SEM (Yanko 1989, volume II, Table IV, Fig. 11b), crystals composing the wall have an elongated habit and are arranged randomly, as a result of which there are gaps of different shapes and sizes between the crystals that apparently act as pores. Clear pore canals are absent. Crystal sizes include length from 0.3 to 1.4 μm and width of 0.01 μm. The test thickness decreases or increases in response to a decrease or increase of salinity, respectively. This dependence can be seen both on modern and fossil foraminifera and can be used for paleogeographic reconstructions. The internal structure of miliolid tests is a fundamental feature in generic diagnosis. However, it is often not expressed in test morphology, which produces errors in determining the genus, as was pointed out by Krasheninnikov (1959), Bogdanowicz (1969), and others. With plentiful and varied material, the author analyzed this feature in all morphological differences of the PontoCaspian Quaternary miliolids using thin cross sections. The following results were obtained: (1) Tests with four chambers on the multichamber and three on the small-chamber sides have quinqueloculine (fivefold) arrangement of chambers at all stages of ontogenesis, with angular distances between them of 72 (Fig. 3.1). Tests with three chambers on the multichamber and two on the small-chamber sides can have (a) a triloculine structure, in which the chambers are located at an angle of 120 (Fig. 3.2a, b), and (b) cryptoquinqueloculine (Bogdanowicz 1969) or pseudotriloculine (Krasheninnikov 1959) structure (Fig. 3.1b, e). In the latter case, the three outer chambers (I, II, III) of the last whorl in adult forms sharply increase in size and overlap the previous ones. The angular distance is 144 between chambers I and II and II and III and 72 between chambers III and I. Chambers IV and V are hidden by external ones; however, they are all adjacent and the angle between them is 72 (Fig. 3.2c, d). Such structure of miliolid tests was first described by Bogdanowicz (1969) and has been fully confirmed on our material. Representatives of Massilina and Pateoris (generation B) have a quinqueloculine arrangement of chambers in the initial whorl, cryptoquinqueloculine in the next one, and planospiral in the rest of the test (Fig. 3.3a–d). Tests of A1 generation have predominantly quinqueloculine arrangement of chambers in all stages of

Fig. 3.1 Inner structure of Quinqueloculina tests. Here and in Figs. 3.3 and 3.4: B, megageneration B; A1 and A2, microgenerations. (Drawn by Irena Motnenko)

Fig. 3.2 Inner structure of Triloloculina (a, b), Sigmella (c, d), and Cornuspira (e, f) tests. (Drawn by Irena Motnenko)

ontogenesis, while tests of A2 generation have all three types of chamber arrangement (Fig. 3.3b–d). A similar arrangement of chambers has a type of species of given genera Ms. rugosa Sidebottom (Loeblich and Таррап

3.5 Taxonomic Classification of Order Lagenida

Fig. 3.3 Inner structure of Massilina and Pateoris tests: (a) and (e) apertural view; (b. c, d) chamber arrangement. (Drawn by Irena Motnenko)

1964, Fig. 350–2с) and Pt. hauerinoides (Rhumbler) (Loeblich and Таррап 1964, Fig. 150–6b). This enables us to supplement and clarify diagnoses of given genera. There is no common understanding of miliolids with a lamellar tooth. Loeblich and Tappan (1964, 1981) distinguish the genera Miliolinella and Soutuloris with triloculine and quinqueloculine, respectively, in the early stages. Bogdanowicz (1969) indicates that, judging by the “descriptions and images of the type species Ml. subrotunda (Montagu), it may include tests with three and four to five external chambers, and there is no certainty that the triloculine type is inherent in the former” (Bogdanowicz 1969, 94). Representatives of this group of miliolids have both cryptoquinqueloculine and quinqueloculine types of chamber arrangements. We found no such forms with triloculine chamber arrangement. Therefore, we refrain from isolating the genus Scutuloris. A more precise diagnosis of the genus Miliolinella is given in the systematic section (see Chap. 8). Thus, we refer to the genus Quinqueloculina for species with quinqueloculine (Q. angulata, Q. ех gr. gracilis, Q. lamarckiana, Q. lata, Q. milletti, Q. seminulum,

31

Q. veпusta, Q. vulgaris) and cryptoquinqueloculine (Q. bicornis, Q. consobrina, Q. inflata, Q. laevigata, Q. oblonga, Q. curvula) types of chamber arrangement. For tests with triloculine chamber arrangement, we refer to the genera Triloculina and Sigmella. In the latter, only the late sections are sigmoiline, while the early chambers are characterized by the triloculine plane of coiling (Fig. 3.2d). With the exception of the genera Massilina, Miliolinella, Pateoris, and Sigmella, diagnoses of all other Miliolida are accepted without change in accordance with the classifications used. In addition to the abovementioned types of test arrangements, our material contains forms with a large proloculum and a spiral-coiled undivided second chamber (Fig. 3.2e, f), with a porcelainlike nonporous wall (genus Cornuspira). We support the opinion of Bogdanowicz (1969) about the need to preserve the name Cornuspira, although it is a junior synonym for Cyclogyra. This is because the former is firmly entered in the paleontological literature and gave rise to the terms derived from it (e.g., “cornuspira,” “cornuspiroid”), which are widely used for the characteristic of a certain type of test arrangement in foraminifera. Thus, in the order Miliolida, we distinguish two families: Cornuspiridae (the most primitive type of test arrangement) and Miliolidae (various modifications of the correctly ball-shaped type of test arrangement). The latter includes four subfamilies: (1) Quinqueloculinae with quinqueloquline and cryptoquinqueloquline chamber arrangements on all stages of ontogenesis (genus Quinqueloculina) (Fig. 3.1), or only at the beginning, later becoming planospiral (Massilina, Pateoris) (Fig. 3.3) or biloculine (Pyrgo) (Fig. 3.4), (2) Miliolinelinae (quinqueloquline type with laminated tooth, Miliolinella), (3) Tubinellinae (quinqueloquline in early stage, later becoming uniserial, Articulina), and (4) Triloculinae (triloculine in all stages of ontogenesis, Triloculina, or only in the early stage and later becoming sigmoline, Sigmella).

3.5

Taxonomic Classification of Order Lagenida

The order Lagenida in the Ponto-Caspian geographic region includes 31 lower taxa. Lagenida tests have a monolamellar, optically, radially radiant and thin perforated wall. The main characteristic features are the test arrangement and aperture that were studied in detail. As a result, seven types of test arrangement were established: secondary single-chambered; uniaxial multichambered; double row in the initial and single row in the final sections; close to quinqueloculine; single row flattened; two rows, laterally compressed; and three rows in the initial part, later turning into two rows. Regardless of the test type and aperture shape (plain or lucid), the

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Taxonomic Classification of Foraminifera

chambers, radial aperture with inner tube on all stages of ontogenesis), (7) Entolingulina (secondary uniserially flattened test, oval aperture with inner tube on all stages of ontogenesis), (8) Oolina (ball- or egg-shaped test, simple rounded or radial aperture with inner tube on all stages of ontogenesis), (9) Fissurina (oval- or ball-shaped, slit-like aperture with inner tube on all stages of ontogenesis), and (10) Parafisurina (oval-shaped, slit-like aperture with inner tube on all stages of ontogenesis) (Table 3.1).

3.6

Fig. 3.4 Inner structure of Pyrgo tests: (a) peripheral view, (b) apertural view, (c) quinqueloquline and cryptoquinqueloquline chamber arrangements at the beginning only, (d, e) biloculine chamber arrangement on the later stage. (Drawn by Irena Motnenko)

overwhelming majority of lagenids have an internal tube of different lengths and configurations as an element of the test internal structure at all stages of ontogenesis. This depends on the functional characteristics of the cytoplasm and acts as a controlling element in the most vulnerable part of the test, the aperture, making such forms better organized compared to those without internal tubes. Within the order Lagenida, the author distinguishes two superfamilies—Nodosarioidea Ehrenberg, 1838 (without internal tube) and Glandulinoidea Yanko, superfam. Nov. (with internal tube)—represented by 30 species from ten genera: (1) Orthomorphina (uniserial test shape, simple rounded aperture), (2) Lagena (single-chambered test, simple rounded aperture), (3) Guttulina (test asymmetrical with irregular arrangement close to the triloculine one at the early stage and uniserial latter, radial aperture with inner tube on all stages of ontogenesis), (4) Glandulina (test biserial at the beginning and uniserial at distal end, slit-like aperture with inner tube on all stages of ontogenesis), (5) Esosyrinx (biserial test with angle between chambers of 90 , radial aperture with inner tube on all stages of ontogenesis); (6) Laryngosigma (biserial test with 180 angle between

Taxonomic Classification of Order Rotaliida

The order Rotaliida is dominant in our material. It includes 65 benthic and 7 planktonic lower taxa. The taxonomic classification of Rotaliida is based on a combination of a significant number of criteria that includes arrangement of test chambers, wall layering and microstructure, presence (absence) of a system of canals, location of single or multiple apertures, and others. Different researchers evaluate differently the significance of individual features, especially among nonionids and elphidiids. Voloshinova and Dain (1952) consider both of them as belonging to one family Nonionoidea, while later Voloshinova (1958) separated them into different families. Loeblich and Tappan repeatedly changed their opinion on this matter. In the Treatise (Loeblich and Tappan 1964), they referred Elphidium to the superfamily Rotalioidea and Nonion to Cassidulinoidea on the basis of a radially radiant wall in the former and granular in the latter. In a later work, Loeblich and Tappan (1981) refer these genera to superfamilies Discorboidea and Nonionoidea, respectively. At the same time, they believed that the nonionids are more highly organized compared to elphidiids. In Loeblich and Tappan (1984), Nonion and Elphidium are included in superfamily Nonionidea and Rotaloidea, respectively, and here the latter are considered to have a higher level of development compared to the former. Mikhalevich (1983) had a similar point of view referring Nonion and Elphidium to independent orders— Nonionida and Elphidiida. Saidova (1981) assigned them to different suborders Nonionina and Elphidiina of the order Nonionida. Similar problems can be noted for other representatives of the order Rotaliida. The ambiguity of the position of individual taxa in the system of order Rotaliida that occupies a high taxonomic position in the evolutionary scheme of foraminifera is associated with a significant variability of signs of the external and internal structures as well as the layering and microstructure of the test wall. These challenges are well illustrated using examples of the PontoCaspian representatives.

3.6 Taxonomic Classification of Order Rotaliida

3.6.1

Test Morphology and Accepted Terminology of Rotaliids

Test Arrangement, Coiling, and Shape In all studied lower taxa, two types of coiling are observed in the course of ontogenesis: (a) without changing the coiling axis (28 lower taxa) and (b) changing the coiling axis at the late stage of the development (38 lower taxa). In both cases, the early whorl (excluding Acervulina and Nonion that have a planospiral test at all stages of ontogenesis) of the tests is either trochospiral at all stages of ontogenesis (e.g., Ammonia) or partially trochospiral with later planispiral (e.g., Haynesina), completely trochospiral, rather high (e.g., Discorbis) or low (e.g., Aubignyna), evolute on dorsal and involute on ventral side. Partially trochospiral tests are usually low-spiral, slightly asymmetric, semi-evolute on dorsal and involute on ventral side (e.g., Haynesina). The spire of the test can increase gradually or very quickly, but the first whorl is always very small and weakly trochoidal. Among trochospiral forms, tests with right- and left-handed direction of coiling are observed, while a change in the direction of coiling in planktonic foraminifera is associated with the alternation of generations (Vaĉiĉek 1953) or with the temperature regime of the sea basin (e.g., Bandy 1960; Shvemberger 1965). For benthic foraminifera from the order Rotaliida (genus Ammonia), an attempt of such analysis was undertaken by Kirienko (1979). According to her data, the decrease in the percentage of right-coiled specimens of Ammonia indicates colder conditions and vice versa. Our studies in this direction do not confirm the conclusions of Kirienko. In the second case, when spiral plane coiling appears at a late stage, semi-involute tests are formed on one side and involute tests on the other (complete involution is quite rare), producing slightly asymmetric tests. The dependence of degree of involution and asymmetricity on di- or trimorphism is observed. As a rule, these signs are most clearly expressed in megalospheric generations and the least in microspheric generations. The canal system, structure of the wall, and septal sutures are correlatively related to each other and therefore are considered together. Before proceeding to their characteristics, some remarks on terminology of the test wall should be made since in the literature, the terms “bilamellar, primary bilayered with primary bilayered septa, and secondary layers of growth” are not always unambiguously understood. Hansen and Lykke-Andersen (1976) combine these two terms and indicate that all rotaliids have a primary bilayered wall with a first bilayered septum and a septal flap (third layer) in some of them as well. The term “bilamellar” wall is used by us in its original sense, while the concept of “bilamellar with a septal flap” (Fig. 3.5b) is embedded in the content of the term “bilamellar–rotaliid” (Fig. 3.5a).

33

In the studied material, four types of septal sutures are observed: 1. simple one- or two-contour, superficial or slightly recessed sutures. Tests with such type of sutures are predominantly or completely trochoid, the wall is bilamellar and radially radiant, and the canal system is absent (Discorbis, Cibicides). The tests can be also flat spiral (Nonion) or low trochoidal (Trichochyalus) with bilamellar and granular walls. They do not have a true canal system; instead there are rudiments of vertical umbilical canals (protocanals) in the form of passages to the umbilical area. 2. Narrow, sunken septal sutures widening toward the umbilical area with development of cracks, so-called lateralumbilical supplementary apertures that open into the interseptal lacunas (e.g., Haynesina, Aubignyna). The latter can extend over the entire or over a part of the septum height. The test usually has a lobular peripheral edge, is either trochoid in the initial whorl, and subsequently becomes spiral planar (Haynesina) or completely low trochoid (Aubignyna). The wall is roughly radially radial (Krasheniknikov 1956), coarsely radial (e.g., Haynesina), or finely granular (e.g., Canalifera) with a bilamellar– rotaloid wall. The septal valve is either short, not significantly reaching the center of the umbilical area (Haynesina), or extending to the entire height of the septum, but in all cases not curved, as in Elphidium. There is no real canal system; however, there are analogues either in the form of interchamber lacunae and additional lateroumbilical apertures (Haynesina) or in the form of a hollow umbilical area covered by strongly elongated abdominal blades of chambers (Ammonia), or there are not always clearly defined two (Porosononion) or one (Aubignyna) spiral canal. 3. Straight or curved septal sutures, complicated by holes and sculpture in the form of septal (Cribroelphidium) or interseptal (Elphidium) bridges. The system of canals is well developed and consists of two spiral (on both sides of the test), meridional and umbilical (vertical) canals. The wall is hyaline, coarsely or indistinctly radially radiant (the vast majority of genera), granular (Nonion) or microgranular (Canalifera), bilamellar–rotaloid, or a septal flap of varying length and degree of bending. 4. Straight or slightly curved sutures, complicated by two rows of holes located in the depressions of the wall with wrinkled relief. The system of canals is similar to that of the genus Elphidium but differs by dichotomous branches of the meridional canals (see Yanko 1989, v. II, Table LXVI, Fig. Ia). The wall is radially radiant and bilamellar. The canal system is similar to that in Elphidium but differs by dichotomously branched meridional canals.

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Taxonomic Classification of Foraminifera

Fig. 3.5 The structure of the foraminiferal test and septal sutures: (a) bilamellar, (b) bilamellar–rotaliid, (c) Nonion, (d) Haynesina, (e) Elphidium, (f) Cribroelphidium, (g) Mayerella. (Drawn by Irena Motnenko)

The Structure of the Aperture and Foramen The study of test apertures in SEM shows that quite often they are indistinguishable due to their “sealing” by an additional skeleton in the form of abundant fine granulation in the lower part of the apertural surface. In an optical microscope, this granulation can be mistaken for holes, giving the false impression of a perforated aperture. This is especially typical for planispiral forms. Obviously, in this case, it is more correct to speak about the taxonomic significance of the foramen structure rather than the aperture. In general, on the studied tests,

two types of aperture (foramina) are observed: simple (slitlike, rounded), predominantly single, rarely multiple (Planorbulina), and complex, perforated. A number of forms have additional apertures: suture (Cibicides) or later umbilical (Aubignyna, Haynesina). Some taxa have an aperture with a lip (Ammonia). The overwhelming majority of completely trochoid and part of planospiral tests have a simple slit-like or rounded aperture. Only forms with a spiral-planar test with complicated septal sutures and a well-developed system of canals have a perforated aperture (foramen).

3.6 Taxonomic Classification of Order Rotaliida

Porosity All lower taxa, without exception, have perforated tests with inner membranes. The pore sizes, their density, and configuration vary widely, but for a certain species these parameters are quite stable and independent of di- or trimorphism. Thus, the common features for all studied rotaliids are trochoid test in the initial section (with the exception of Nonion), the porous bilamellar (with the bilamellar–rotaliid variant) wall. These characters seem to have a high taxonomic rank, that is, are characteristic of the order. The structure of the test in the later sections, the structure of the wall (granular or radially radiated), and the presence (absence) of a real system of canals or its analogues should be considered as characteristics of the superfamily. Family-level features should consider the structure of septal sutures as a whole (complicated or uncomplicated), the system of canals or its analogs, the microstructure of the wall, and the location and structure of the aperture. The nature of the septal sutures (the presence of septal, interseptal bridges, or suture openings on them), the structure of additional apertures, and the shape of the main aperture (perforated, slit-like, rounded) are used as the main features of the genus. As for the species, along with the usual features of this taxonomic category (peripheral edge, the nature of the additional skeleton, the aperture size, etc.), great importance is assigned to the nature of porosity (density of pores, their configuration, location, and size).

3.6.2

Systematics and Genetic Interrelations

In our material, Ponto-Caspian Rotaliida are represented by the superfamilies Discorboidea, Anomalinoidea, Nonionidea, and Rotalioidea. Within Discorboidea and Anomalinoidea, the steady transformation from solitary, intermarginal, slit-like aperture on ventral side of the family Discorboidea that can be short in the subfamily Discorbiinae (Discorbis, Gavelinopsis) or long in the subfamily Rotaliinae (Rosalina) to a centered round-shaped aperture in the family Glabratellidae (Heronallenia) and then to multiple apertures located on the dorsal side in families Cibicidae (Cibicides), Planorbulinidae (Planorbulina), and Acervulinidae (Acervulina) has been noticed. The superfamily Nonionidae is represented by the families Nonionidae (Nonion, Florilus) and Trichochyalidae (Trichochyalus). The former has an unclear aperture, slitlike foramen, and granulated umbilical area, which exits as protocanals. The latter has intermarginal slit-like and slightly obliquely truncated toward the ventral side aperture covered with the cork of skeletal additional material perforated by exits as protocanals. The superfamily Rotalioidea includes the family Ammoniidae, as well as the Canaliferidae and Elphidiidae.

35

The former includes taxa with trochoid tests of different heights. The real canal system is absent. The hollow umbilical area is covered by long lobes with an umbilical– extraumbilical aperture under the last lobe. Foramens are transformed in suture or umbilical slits and partially covered with supplementary skeleton on early chambers. The family is represented by the genus Ammonia only, which plays a dominant role (up to 100%) in foraminiferal assemblages. The taxonomic classification of Ammonia is very complicated due to the high variability, which is based on the combination of test morphology, diameter, thickness, number of chambers in the final coil, ratio between diameter and thickness, size of proloculum, and density, distribution, configuration, and size of the pores distinguishing the ten groups of Ammonia. The ecological, geographic, and stratigraphic characteristics of each group indicate that it is a separate species (Yanko 1990a). A. caspica lives in the Caspian and Aral Seas. This oligohaline species is widely distributed in the Neopleistocene of the Ponto-Caspian region as well as the Holocene of the Caspian Sea. A. novoeuxinica is a holeuryhaline species known in the Ponto-Caspian region from the Pliocene. Its maximal distribution coincides with regressional phases (e.g., Kolkhidian, Pontian). Ammonia tepida is also holeuryhaline species, which appears in geological sequences in the middle of Early Neopleistocene. It is widely distributed today in the Black Sea, Sea of Azov, Mediterranean Sea, and World Ocean. A. caucasica is a stricteuryhaline species. It appears at the beginning and disappears at the end of the Uzunlarian period. Then it appears again in the middle Holocene with the most extensive distribution in transgressive phases. Today it is widely distributed mainly on the Caucasian shelf. Polyhaline A. agoiensis, A. beccarii, and A. parkinsoniana are known from the Karangatian layers. The two latter species are widely distributed in the Mediterranean Sea but absent in the Black Sea even near the Bosphorus. Polyhaline A. compacta first appeared in the Karangatian layers and then again in the middle Holocene with the highest abundance at transgressive phases. In the Black Sea today this species occurs most commonly in waters with salinity higher than 18‰. It does not live in the Sea of Azov but is widely distributed in the Mediterranean Sea (Hofker 1969). The strictoeuryhaline species, A. ammoniformis, appears in the Black Sea in the middle of the Holocene, with maximum distribution in transgressive phases. Today, this species occurs most commonly in western and southwestern parts of the Black Sea and is common in the Mediterranean Sea and Atlantic Ocean (Colom 1974). A. caspica and A. novoeuxinica may have a common ancestor, Ammonia galiciana (Putrja) (Putrja 1964). The latter is also probably the ancestor of A. parasovica, while A. caucasica could have diverged from A. tepida. The family Canaliferidae includes only one genus, Canalifera. Based upon its granular wall structure, this

36

3

genus was earlier placed in the Nonionidae (Saidova 1981). On its bilamellar–rotaloid wall texture, well-developed canal system, and sutures with septal bridges, this genus was placed in Elphidiidae (Loeblich and Tappan 1984). However, those test characters are equal in importance and therefore neither placement is valid. The Canaliferidae seems to be an independent family in the superfamily Rotalioidea. The family Elphidiidae includes the subfamilies Haynesininae and Elphidiinae. The former includes the genera Porosononion, Aubignyna, and Haynesina, which are known starting from the Palaeocene, Lower Pliocene, and Upper Pliocene, respectively. Similarity in the wall structure and texture, septal sutures, aperture, and foramen characteristics suggests their relationship. Aubygnina seems to be an ancestor of Haynesina (Banner and Culver 1978; Levchuk 1983; Yanko 1989), while Porosononion possibly diverged from Protelphidum. Porosononion and Protelphidum are not synonyms, as suggested by Loeblich and Tappan (1964, 1981). Their type species, Protelphidium hofkeri Haynes and Po. subgranosus Putrja, have different test structures (granular and radial, respectively). Moreover, the former does not have canals. The subfamily Elphidiinae includes the genera Elphidium, Cribroelphidium, and Mayerella. In their early stages, these genera have slightly trochoid tests that rapidly develop into planispiral forms. Other common features include welldeveloped canal systems represented by spiral, umbilical, and meridional canals and the presence of septal and interseptal bridges on septal sutures with a specific architecture for each genus (Yanko and Troitskaya 1987; Yanko 1989). Mayerella was first discovered in the Lower Holocene of the Pontic region. Later it was found in the Pliocene and Eopleistocene of the Caspian region and Lower Neopleistocene of the Ponto-Caspian region. Today members of the genus live in the Black Sea (Ma. kolchidica) only in shallow deltaic areas with very low salinity, as well as in the Caspian (Ma. brotzkajae) and Aral (Ma. aralica) Seas. Possible phyletic lineages between Elphidium and Porosononion species and subspecies are shown in Yanko (1990b). Some representatives of these phyletic lineages correspond to LAGENIDA 17% MILIOLIDA 24%

ROTALIIDA 37%

Taxonomic Classification of Foraminifera

certain stratigraphic levels, making them especially valuable for stratigraphic purposes.

3.7

Taxonomic Classification of Order Buliminida

The order Buliminida includes 12 species. All studied species have primary bilamellar (Hansen and Reiss 1972; Afanasyeva 1982), spiral, helical, finely perforated tests with radial wall and inner membrane. The order includes three families: Buliminidae (triserial tests: Bulimina), Uvigerinidae (primarily triserial and later from biserial to uniserial chamber arrangement: Trifarina), and Bolivinidae (biserial on all stages of ontogenesis). The Buliminida never play significant roles in the assemblages and are represented by accessory forms only. Based on the above morphological concepts of the author, the following classification of the Ponto-Caspian Quaternary foraminifera is proposed (Table 3.1).

3.8

Systematics and Comparison with Other Basins

The Ponto-Caspian Quaternary benthic foraminifera are represented by eight orders, 35 families, 66 genera, and 180 species (some in open nomenclature) and subspecies (i.e., lower taxa). Eighteen species are absent in our material (marked by stars in Table 3.1) but are still included in the classification. Of the 180 species, 123 lower taxa inhabit the Black and Caspian Seas today (see Chap. 4). A majority of lower taxa (150) belong to calcareous orders: Rotaliida (65), Miliolida (42), Lagenida (31), and Buliminida (12). The other 30 species belong to the agglutinated orders Ammodiscida, Astrorhizida, Ataxophragmiida, and Schlumbergerinida (Fig. 3.6). The depleted concentration of agglutinated foraminifera may be partially associated with postmortem breakdown of their tests. BULIMINIDA 12%

AMMODISCIDA 7% BULIMINIDA 6% ASTRORHIZIDA 4% ATAXOPHRAGMIDA 3% SCHLUMBERGERINIDA 2%

a

MILIOLIDA 21%

ROTALIDA 45%

Fig. 3.6 Cyclogram of the percentage of orders of Quaternary foraminifera the Ponto-Caspian (a) and Russian North basins (b)

LAGENIDA 22%

b

3.8 Systematics and Comparison with Other Basins

The dominant role of the order Rotaliida among the lower taxa is maintained throughout the entire Neopleistocene and Holocene. In the Eopleistocene, only Rotaliida compose foraminiferal assemblages. The order Miliolida appears at

37

the beginning of the Early Neopleistocene, while Lagenida at its end (Fig. 3.7). Possible reasons for the uniqueness of the systematic composition at different stratigraphic levels are discussed in Chap. 6.

Fig. 3.7 Vertical distribution of foraminiferal orders in the Quaternary sediments of the Pont

38

Compared with other basins/regions, only 132 lower taxa are found among the Ponto-Caspian foraminifera; 19 of them are absent among life foraminifera. The most common occurrences (72 taxa) are between the Black and Mediterranean Seas (Todd 1958; Hofker 1960; Parisi 1981; Cimerman and Langer 1991; Yanko et al. 1994, 1998; Yanko 1995; Basso and Spezzaferri 2000). The Black Sea and Russian North have only 8 species in common (Gudina 1976). The number of species in common with the southern and the tropical Atlantic is 41, North Atlantic is 24, North Sea is 32, and Antarctica is 14. All these unambiguously indicate the main source of the Ponto-Caspian Quaternary foraminiferal fauna. According to Mikhalevich (1983), 306 species of benthic foraminifera belonging to 131 genera and 12 orders live on the shelf of the tropical Atlantic, while in the Pleistocene of the Russian North, 152 species and subspecies from 74 genera and 8 orders were found. Thus, the foraminiferal fauna is equally depleted in the Ponto-Caspian region and the Russian North compared with the tropical Atlantic. Only one order (Cassidulinida) is absent in the Ponto-Caspian region but present in the Russian North (Fig. 3.6b). The other seven orders (Fig. 3.6) are common to both regions, revealing the tendency of forming foraminiferal fauna from orders that occur in multiple kinds of facies. The absence of the order Cassidulinida in the Ponto-Caspian region, although it is well represented in the Mediterranean, can be explained by its depth preferences. They typically live at depths that in the Black Sea are contaminated with H2S. Of the seven orders found in both the south and north, the most widespread are representatives of order Rotaliida. However, the family Elphidiidae dominates in the north, while Ammoniidae prevails in the south (Ponto-Caspian). This is the result of the more severe thermal conditions of the northern basins, in which the warm-dwelling genus Ammonia does not have favorable conditions. Apparently for analogous environmental reasons, the families Canaliferidae, Planorbulinidae, and Acervulinidae are not found in the Russian north, though they are widespread in the Mediterranean and the southern and tropical Atlantic and are found quite often in the Black Sea. Instead, the cold-water genus Buccella is abundantly represented in the north. Higher taxonomic categories of the Ponto-Caspian foraminifera have high similarity with those from the Northern basins (e.g., Barents Sea, North Sea) and low similarity with those of the Mediterranean and southern and tropical Atlantic. This enables us to conclude about the cold-water character of the Ponto-Caspian foraminifera. It also mirrors the general tendency of the formation of foraminiferal faunas in hydrologically anomalous basins (e.g., low salinity of the Ponto-Caspian, or low temperature of the northern seas) from a limited number of orders: Astrorhizida, Ammodiscida,

3

Taxonomic Classification of Foraminifera

Ataxophragmiida, Lagenida, Rotaliida, and Buliminida occurring in multiple kinds of facies.

References Afanasyeva MS (1982) Strukturnye elementy stenki rakovin buliminid (foraminifery) (Structural elements of the wall of the tests of buliminids (foraminifera)). Paleontologichkiy zhurnal 4:14–20 (in Russian) Bandy OL (1960) General correlation of foraminiferal structure with environment. Int. Geol. Congress. 21st Ses. Rep. 1960. Proc. sect. 22, pp 7–19 Banner FT, Culver S (1978) Quaternary Haynesina, n.gen. and Paleogene Protelphidium Haynes; their morphology, affinities and distribution. J Foram Res 8(3):177–207 Basso D, Spezzaferri S (2000) Distribution of living (stained) benthic foraminifera in the Iskenderun Bay (Eastern Turkey): a statistical approach. Boll Soc Paleontol Ital 39(3):359–379 Bogdanowicz AK (1969) Meoticheskie Miliolida zapadnogo Predkavkazya (Meotic Miliolida of western Ciscaucasia). In: Yergoyan VL (ed) Geol. i neftegaz. Zap. Kavkaza i Zap. Predkavkazya. Trudy KF VNIINeft I9, Moscow, pp 64–114 (in Russian) Cimerman F, Langer MR (1991) Mediterranean Foraminifera: Slovenska Academia Znanosti, Umetnosti, Ljublana, 119 pp Colom G (1974) Foraminiferos ibericos. Introduccion al estudio de las especies bentonicas recientes. Investigacion Pesquera 38:1–245 Fursenko AV (1978) Vvedenie v izuchenie foraminifer (Introduction to the study of foraminifera). Trudy Instituta Geologii i Geofiziki 391. Nauka, Novosibirsk, 242 pp (in Russian) Gudina VI (1976) Foraminifery, stratigrafiya i paleoeoogeo-grafiya morskogo pleystotsena severa SSSR (Foraminifera, stratigraphy and paleoeoogeo-graphy of the marine Pleistocene of the north of the USSR). Novosibirsk: Trudy IGIG SO AN SSSR, vyp. 314, 126 pp (in Russian) Hansen HJ, Lykke-Andersen AL (1976) Wall structure and classification of fossil and recent elphidiid and nonionid foraminifera. Fossils and Strata, no 10, 60 pp Hansen HJ, Reiss Z (1972) Scanning electron microscopy of -wall structure in some benthonic and planctonic Foraminiferida. Rev. Esp. Micropal. 4(2):169–181 Hofker J (1960) Foraminiferen aus dem Golf von Neapel. Palaeontolog. Zeitshrift 34:233–262 Hofker J (1969) Recent foraminifera from Barbados. Studies on the Fauna of Curaçao and other Caribbean Islands 31:1–158 Holzmann M (2000) Species concept in foraminifera: ammonia as a case study: micropaleontology, v. 46, Supplement 1: advances in the biology of foraminifera, pp 21–37 Kirienko EA (1979) Napravleniye navivaniya i krupnost’ rakoviny Ammoniz neobeecarii subspec. i ikh znacheniye dlya stratigrafii (Winding direction and shell size of Ammonia neobeecarii subspec. and their significance for stratigraphy). Vestnik Leningr. un-ta. 6: 113–116 (in Russian) Krasheninnikov VA (1956) Mikrostruktura stenki nekotorykh kaynozoyskikh foraminifer i metodika ee izucheniya v polyarizovan-nom svete (Microstructure of the wall of some Cenozoic foraminifera and methods of its study in polarized light). Voprosy mikropaleontologii 1:37–48 (in Russian) Krasheninnikov VA (1959) Foraminifery (Foraminifera). In: Atlas srednemiotsenovoy fauny Severnogo Kavkaza i Kryma (Foraminifers. Atlas of the middle Miocene fauna of the North Caucasus and Crimea). Moscow, pp 15–103 (in Russian)

References Levchuk LK (1983) Rod Haynesina Banner Culver, 1978 (foraminifera) v pleystotsene severa Sibiri (Genus Haynesina Banner Culver, 1978 (foraminifera) in the Pleistocene of northern Siberia). In: Dagis AS, Dubatolova VN (eds) Morphology and systematics of Phanerozoic invertebrates. Moscow, Nauka, pp 96-IO2. (Tr. IGiG SB AS USSR; Issue 538) (un Russian) Linnaeus C (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata [10th revised edition], Laurentius Salvius, Holmiae, vol 1, 824 pp Loeblich AR, Tappan H (1964) Treatise on invertebrate paleontology, Part C: Protista 2, Sarcodina, chiefly “Thecamoebians” and Foraminiferida. Geological Society of America and University of Kanzas Press, Lawrence Loeblich AR, Tappan H (1981) Suprageneric revizions of some calcareous Foraminiferida. J Foraminifer Res 11:159–164 Loeblich AR, Tappan H (1984) Some new proteinaceous and agglutinated genera of foraminiferida. J Paleontol 58:1158–1163 Loeblich AR, Tappan H (1988) Foraminiferal genera and their classification. Van Nostrand Reinhold Company, New York Loeblich AR, Tappan H (1992) Present status of foraminiferal classification. In: Takayanagi Y, Saito T (eds) Studies in Benthic foraminifera. Proceedings of the fourth symposium on benthic foraminifera, Sendai, 1990, pp 93–102 Mayer EM (1968) Podklass foraminifery (Subclass foraminifera). In: Birshteyn YAA et al (eds) Atlas bespozvonochnykh Kaspiyskogo morya. Pischevaya promyshlennost, Moscow, pp 11–34 (in Russian) Mayer EM (1972) Sovremennye foraminifery Priapsheronskogo payona Kaspiyskogo morya (Recent forminifera of Near Apsheron region of the Caspian Sea). In: Leont’ev OK, Maev EG (eds) Kompleksnye issledovaniya Kaspiyskogo morya 3, pp 25–36 (in Russian) Mayer EM (1976) Zametka o sakkaminidakh (Foraminiferida) Kaspiyskogo morya. In: Maev YeG (ed) Kompleksnye issledovaniya Kaspiyskogo morya 5, pp 257–260 (in Russian) Mayr E (1971) Printsipy zoologicheskoy sistematiki (Principles of zoological systematic). Mir, Moscow, 454 pp (in Russian) Mayr E, Linsli E, Yüzinger R (1956) Metody i printsipy zoologicheskoy sistematiki (Methods and principles of zoological systematic). Izd-vo insstr.lit., Moscow, 49 pp (in Russian) Mikhalevich VI (1968) Otryad foraminifery (Order foraminifera). In: Vodyanitskiy VA (ed) Opredelitel fauny Chernogo i Azovskogo morey, pp 9–21 (in Russian) Mikhalevich VI (1980) Sistematika i evolyutsiya foraminifer v svete novykh dannykh po ikh tsitologii i ultrastrukture (Systematics and evolution of foraminifera in the light of new data on their cytology and ultrastructure). In: Krylov MV (ed) Printsipy postroeniya makrosistemy odnokletotsnykh zhivotnykh. Tr. Zool.in-ta AN SSSR, vol 94, pp 42–61( in Russian) Mikhalevich VI (1983) Donnye foraminifery shelfov tropicheskoy Atlantiki (The bottom foraminifera from the shelves of the tropical Atlantic). Izd-vo Zool. in-ta AN SSSR, Leningrad, 247 pp Mikhalevich VI (2013) New insight into the systematics and evolution of the foraminifera. Micropaleontology 59(6):493–527 Orlov YUA (ed) (1959) Osnovy paleontologii (Basics of paleontology). Volume 1, general part, protozoans. Russian Academy of Sciences, Moscow, 519 pp (in Russian) Parisi E (1981) Distribuzione dei foraminiferi bentonici nelle zone batiali del Tirreno e del Canale di Sicilia. Riv. Ital. Paleontol. 87(2):293–328 Pawlowski J, Holzmann M, Tyszka J (2013) New supraordinal classification of foraminifera: molecules meet morphology. Mar Micropaleontol 100:1–10

39 Putrja FS (1964) O nekotorykh novykh vidakh miotsenovykh foramiØnifer Vostochnogo Predkavkaz’ya (On some new species of Miocene foraminifers of the Eastern Ciscaucasia). Paleont. zhurn. 3:127–131 (in Russian) Saidova KHM (1981) O sovremennom sostoyanii sistemy nadvidovykh taksonov kaynozoyskikh bentosnykh foraminifer (On the current state of the system of supraspecific taxa of Cenozoic benthic foraminifers). AN SSSR, Moscow, 720 pp (in Russian) Schultze MS (1854) Über den Organismus der Polythalamien Foraminiferen, nebst Bemerkungen über die Rhizopoden im allgemeinen. Ingelmann, Leipzig, pp 1–68 Shvemberger YuN (1965) O znachenii napravleniya navivaniya u rannepaleogenovykh globorotaliy Severnogo Kavkaza (On the significance of the winding direction in the early paleogene globorotalians of the North Caucasus). In: Rauzer-Chernousova DM (ed) Voprosy mikropaleontologii vyp. 9, pp 189–197 (in Russian) Todd R (1958) Foraminifera from Western Mediterranean Deep-Sea cores. Rep. Swedish Deep-Sea exiled. 1947–1948 VIII(4):167–215 Tufescu M (1973) Les associations de foraminiferes du Nord-Quest de la Mer Noire. Rev Espanola de Micropaleontol 5(1):15–32 Tufescu M (1974) Populatiile de foraminifere din apele litorale romanesti. Edit. Acad. Rep. Soc. Romania. Bucuresti, 75 pp Vaĉiĉek M (1953) Zweny vsaiemneho pomeru ľavák a pravák jedinou foraminifery Globorotalia scitula (Brady) a jedinou vyzizi ve stratigrafii. Ustrêd. uszavu geol. Sbornik Odd Paleontol 20:1–76 Voloshinova NA (1958) O novoy sistematike nonionid (On the new taxonomy of nonionids). In: Subbotina NN (ed) Mikrofauna SSSR. Trudy VNIGRI vyp. 115, pp 117–224 (in Russian) Voloshinova NA, Dain LG (1952) Nonionidy, kassidulinidy i khillostomelidy (Nonionids, cassidulinids, and chyllostomelids). In: Iskopayemyye foraminifery SSSR. Trudy VNIGRI vyp. 63, 152 pp (in Russian) WoRMS Editorial Board (2021). World register of marine species. Available from http://www.marinespecies.org at VLIZ. Accessed 25 June 2021. https://doi.org/10.14284/170 Yanko V (1989) Chetvertichnie foraminiferi Ponto-Kaspiya (Chernoe, Azovskoe, Kaspiiskoe i Aral’skoe morya): taxonomiya, biostratigrafiya, istoriya razvitiya, ecologiya (Quaternary foraminifera of the Pontic-Caspian Region (the Black Sea, Sea of Azov, Caspian Sea, and Aral Sea): taxonomy, biostratigraphy, history, ecology). DSci thesis, Moscow State University, 1000 pp (in Russian) Yanko V (1990a) Chetvertichnye foraminifery roda Ammonia PontoKaspiya (Quaternary foraminifera of genus Ammonia of the PontoCaspian). Paleontol J 1:18–26. (in Russian) Yanko V (1990b) Stratigraphy and paleogeography of marine Pleistocene and Holocene deposits of the southern seas of the USSR. Mem Soc Geol Ital 44:167–187 Yanko V (ed) (1995) Benthic foraminifera as indicators of heavy metals pollution – a new kind of biological monitoring for the Mediterranean Sea. European Commission, Program Avicenne, Annual Report AVI CT92-0007, 270 pp Yanko V, Troitskaya TS (1987) Pozdnechetvertichye foraminifery Chernogo morya (Late Quaternary foraminifera of the Black Sea). Nauka, Moscow, 111 pp Yanko V, Kronfeld A, Flexer A (1994) The response of benthic foraminifera to various pollution sources: implications for pollution monitoring. J Foraminifer Res 24:1–17 Yanko V, Ahmad M, Kaminski M (1998) Morphological deformities of benthic foraminiferal tests in response to pollution by heavy metals: implications for pollution monitoring. J Foraminifer Res 28(3): 177–200

4

Modern Foraminifera

Abstract

Distributional patterns of modern foraminifera across a wide range of bionomic zones in the Ponto-Caspian basins, along with depth and salinity gradients, provide the information required for the Quaternary ecostratigraphy and paleogeographic reconstructions. The chapter is divided regionally: the Black Sea shelf is subdivided as northwestern (deltas, limans, lagoons), western, and eastern Crimean; Kerch Strait; Sea of Azov; Caucasian; Bulgarian, southwestern shelf, and Bosphorus outlet; and Caspian Sea (north, middle, south, bays, and straits). In the northwestern shelf, modern foraminifera were studied from the Danube Delta, open limans with major, minor, and without river input and semiclosed and closed limans and lagoons. The comparison of foraminiferal assemblages demonstrates that composition and structure vary with environmental parameters, most importantly, salinity. All assemblages are dominated by Ammonia species, including Ammonia novoeuxinica in river deltas and open limans with free connection to the Black Sea and intensive river discharge, while in limans that are influenced by seawater input through a narrow channel and without river discharge, Ammonia tepida dominates. Mayerella species are characteristic of the river delta and low-salinity limans; they sharply decrease and finally disappear in the limans connected to the sea. In most foraminiferal assemblages, Haynesina anglica is a dominant and characteristic species. Miliammina fusca is characteristic of open, brackish limans with sandy substrates. Trichochyalus aguajoi is found in all types of limans confined to floodplains, with abundance increasing with increasing salinity. Closed limans with salinity