Advances in Phycological Studies

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Author: Nadja Ognjanova-rumenova | Kalina Manoylov

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her current and former students, colleagues, collaborators, and friends over the years. This volume, in celebration of her 70th birthday, is a collection of 25 peer-reviewed research articles on different aspects of phycology and is intended to reflect her diverse academic interests and areas of expertise. The topics covered in this volume include: taxonomy, paleoecology, physiology, and ecology of different algal groups. Much of the work focuses on diatom research including new taxonomic descriptions (a new genus Hyalosigma, fourteen new diatom species, and one new combination), discussion of evolutionary patterns, ecology of freshwater species and the use of diatoms in bioassessment. However, the systematics and physiology of some other algal groups are also presented. This collection, a representation of the many ways algae contribute to our understanding of nature, will be of value to any true “phycologist” in the field as well as to any biological library.

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ADVANCES IN PHYCOLOGICAL STUDIES

Dobrina Temniskova-Topalova includes articles by some of

Festschrift in honour of Prof. Dobrina Temniskova-Topalova

Advances in Phycological Studies: Festshrift in honour of Prof.

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ADVANCES IN PHYCOLOGICAL STUDIES Festschrift in honour of Prof. Dobrina Temniskova-Topalova

Editors Nadja Ognjanova-Rumenova & Kalina Manoylov

St. Kl. Ohridski University Publishing House

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ADVANCES IN PHYCOLOGICAL STUDIES Festschrift in Honour of Prof. Dobrina Temniskova-Topalova Editors Nadja Ognjanova-Rumenova & Kalina Manoylov

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ADVANCES IN PHYCOLOGICAL STUDIES Festschrift in Honour of Prof. Dobrina Temniskova-Topalova Editors Nadja Ognjanova-Rumenova & Kalina Manoylov

Pensoft Publishers St. Kliment Ohridski University Press Sofia–Moscow 2006 3

ADVANCES IN PHYCOLOGICAL STUDIES Festschrift in Honour of Prof. Dobrina Temniskova-Topalova

Editors Nadja Ognjanova-Rumenova & Kalina Manoylov

First published 2006 ISBN-10: 954-642-260-6 (Pensoft Publishers) ISBN-13: 978-954-642-260-6 (Pensoft Publishers) ISBN-10: 954-07-2354-X (St. Kliment Ohridski University Press) ISBN-13: 978-954-07-2354-9 (St. Kliment Ohridski University Press)

Cover photo: Pontodiscus baldjickianus Temniskova et Kozyrenko. Photo: Prof. D. Temniskova-Topalova.

©

PENSOFT Publishers

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.

Pensoft Publishers Geo Milev Str. 13a, 1111 Sofia, Bulgaria E-mail: [email protected] www.pensoft.net

Printed in Bulgaria, April 2006 4

CONTENTS

Preface

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Prof. Dr. Biol. Sc. Dobrina N. Temniskova-Topalova (A life dedicated to the students) Dimitar Vodenicharov, Kalina Manoylov & Nadja Ognjanova

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Two new sigmoid, naviculoid diatoms from the Eocene of Kazakhstan N. I. Strelnikova, J. P. Kociolek & E. Fourtanier

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Species of the genus Terpsinoë Ehrenberg (Bacillariophyta) from the Miocene of Middle Russia, Moldova, and Ukraine Tatyana F. Kozyrenko

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Comparison of seven species of Navicula sensu stricto. Six species described as new to science from Miocene lacustrine deposits in Bulgaria and Romania Horst Lange-Bertalot & Ditmar Metzeltin

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Evolution of the extinct genera belonged to the family Stephanodiscaceae (Bacillariophyta) during the last eight million years in Lake Baikal Galina Khursevich

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A new Cymbella from the Neogene in Bulgaria and its stratigraphic significance Nadja Ognjanova-Rumenova & Ditmar Metzeltin

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Nitzschia toskalensis sp. nov. a new diatom (Bacillariophyceae) from the sediments of Toskaljavri, northwestern Finland Paul B. Hamilton & Jan Weckström

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Distribution of resting spores of Eunotia soleirolii and Meridion circulare var. constrictum (Bacillariophyta) in sediments of peat bogs from Mt. Central Sredna Gora, Bulgaria Rosalina Stancheva 5

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Taxonomic Status of Detonia Frenguelli and the Establishment of Detonia dobrinae sp. nov. (Bacillariophyceae) Ioanna Louvrou, Daniel Danielidis & Athena Economou-Amilli Staurosira incerta (Bacillariophyceae) a new fragilarioid taxon from freshwater systems in the United States with comments on the structure of girdle bands in Staurosira Ehrenberg and Staurosirella Williams et Round Eduardo A. Morales Achnanthidium temniskovae sp. nov., a new diatom from the Mesta River, Bulgaria Plamen Ivanov & Luc Ector A new Gyrosigma species from lakes Prespa and Ohrid Levkov Zlatko, Svetislav Krstic & Teofil Nakov

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Diatom species composition from the river Iskar in the Sofia region, Bulgaria Plamen Ivanov, Emilia Kirilova & Luc Ector

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Chara corfuensis J. Gr. ex Fil. 1937 (Characeae) – an endemic species of Balkan Peninsula, rare and globally endangered Jelena Blaženčiæ

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First record of the tropical invasive alga Compsopogon coeruleus (Balbis) Montagne (Rhodophyta) in Flanders (Belgium) Maya P. Stoyneva, Koenraad Vanhoutte & Wim Vyverman

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Initial observations on uniparental auxosporulation in Muelleria (Frenguelli) Frenguelli and Scoliopleura Grunow (Bacillariophyceae) Mark B. Edlund & Sarah A. Spaulding

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Effect of abscisic acid on host susceptibility during different ontogenetic phases of the host alga in the pathosystem Scenedesmus acutus – Phlyctidium scenenedesmi Irina D. Pouneva & Christo Christov

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Diatom succession in the Ferdynandovian Interglacial lacustrine deposits of Poland Barbara Marciniak

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Environment variation in the Black Sea region during the Late Quaternary based on fossil diatoms A. P. Olshtynskaya

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Late Pleistocene/Holocene dinoflagellate cyst assemblages from the Southwestern Black Sea shelf Mariana Filipova-Marinova

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Diatoms as indicators of the influence of the Vistula river inflow on the Gulf of Gdańsk during the Holocene Katarzyna Stachura-Suchoples

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Diatom flora diversity in the strongly eutrophicated and α-mesosaprobic waters of the Szczecin Lagoon, NW Poland, southern Baltic Sea Ma³gorzata B¹k, Andrzej Witkowski & Horst Lange-Bertalot

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Use of nonparametric multiplicative regression for modeling diatom habitat: a case study of three Geissleria species from North America Marina G. Potapova & Diane M. Winter

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Density-dependent algal growth along N and P nutrient gradients in artificial streams Kalina M. Manoylov & R. Jan Stevenson

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Atypical Tabularia in Coastal Lake Erie, USA E. F. Stoermer & N. A. Andresen Refining diatom indicators for valued ecological attributes and development of water quality criteria R. Jan Stevenson

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List of Referees ALVERSON, Andrew 1 University Station (A6700), 311 Biological Laboratory, University of Texas at Austin, Austin, Texas 78712 ANDRESEN, Norman Andresen Consulting, LLC, 5742 Princeton Place, Ypsilanti, MI 48197 BERKMAN, Julie US Geological Survey, 6480 Doubletree Ave., Columbus, OH 43229 HEINLEIN, Juliane Department of Zoology, Michigan State University, 203 Natural Sciences Bldg. East Lansing, MI 48824 KHURSEVICH, Galina Institute of Geochemistry and Geophysics, National Academy of Sciences of Belarus, Kuprevich street 7, Minsk 220141, Republic of Belarus LEROY, Suzanne Department of Geography and Earth Sciences, Brunel University, Uxbridge, Middlesex UB8 3PH, (West London), UK LANGE-BERTALOT, Horst Botanical Institute, J.-W. Goethe-University, Senckenberganlage 31–33, 60054 Frankfurt am Main, Germany LOWE, Rex Department of Biological Sciences, Bowling Green State University Bowling Green, OH 43403 MANOYLOV, Kalina Department of Zoology, Michigan State University, 203 Natural Sciences Bldg. East Lansing, MI 48824 MORALES, Eduardo Phycology Patrick Center for Environmental Research, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195 U.S.A. OGNJANOVA-RUMENOVA, Nadja Institute of Geology, Bulgarian Academy of Sciences, Acad. G. Bonchev str. 24, 1113 Sofia, Bulgaria

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PANAYOTOFF, Lara Kentucky Division of Water, 14 Reilly Road, Frankfort, KY 40601 POTAPOVA, Marina Phycology Patrick Center for Environmental Research, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195 U.S.A. ROLLINS, Scott Department of Zoology, Michigan State University, 203 Natural Sciences Bldg. East Lansing, MI 48824 SLAVCHOVA, Nadezda Environmental Sciences and Resources Portland State University PO Box 751 Portland, OR 97207 STACHURA-SUCHOPLES, Katarzyna Alfred Wegener Institute for Polar and Marine Research, Telegrafenberg A43, D-14473 Potsdam, Germany STANCHEVA, Rozalina Department of Botany, Faculty of Biology, St. Kliment Ohridski University of Sofia, D. Tsankov Blvd 8, 1164 Sofia, Bulgaria Current: Environmental Sciences and Resources Portland State University PO Box 751 Portland, OR 97207 STERRENBURG, Frithjof Stationsweg 158, 1852 LN Heiloo, The Netherlands STRELNIKOVA, Nina Department of Botany, Biological Faculty, St.Petersburg State University, Universitetskaya Emb. 7/ 9, St. Petersburg, 199034, Russia SWEETS, Roger P. University of Indianapolis, Biology Department, Indianapolis, IN 46227 TONCHEVA, Tonka Institute of Plant Physiology, Bulgarian Academy of Sciences, Sofia 1113, Bulgaria, Georgi Bonchev Str., Block 21 URBANC-BERCIC, Olga Department of Aquatic and Terrestrial ecology, National Institute of Biology, Vecna pot 111, 1000 Ljubljana Slovenia WITKOWSKI, Andrzej Institute of Marine Sciences, University of Szczecin, Waska 13, PL-71-415 Szczecin, Poland WACHNICKA, Ania Southeast Environmental Research Center & Earth Sciences Department Florida International University, Miami, FL 33199

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Preface The Festschrift to honor Prof. Dobrina Temniskova-Topalova’s 70th birthday includes articles by some of her current and former students, colleagues, collaborators, and friends over the years. The volume is a collection of 25 research articles on different aspects of Phycology and is intended to reflect her versatile academic interests and areas of expertise. The topics covered by the papers include: taxonomy, palaeoecology, physiology, and ecology of different algal groups. Articles describe new taxa from fossil and recent materials of Europe, Asia and North America: a new genus Hyalosigma, fourteen new diatom species, and one new combination. Important patterns of the morphological evolution in extinct diatom genera of family Stephanodiscaceae in Lake Baikal are discussed. New floristic and taxonomic information for the distribution of the endemic Chara corfuensis J. Gr. ex Fil. and the tropical Compsopogon coeruleus (Balbis) Montagne is provided. Research on various biostratigraphical and palaeoecological marine and lacustrine basins, based on diatoms and dinoflagellate cysts, is presented. More specialized topics like uniparental auxosporulation are presented also. Careful analyses of the infection process in the green microalga pathosystem and an unicellular fungal parasite. A substantial group of articles focuses on the ecology of freshwater diatoms – redefinition of their importance in bioassessment, density and nutrient dependence of diatom, and the use of new analytical methods like Nonparametric multiplicative regression modeling. The editors would like to thank all colleagues that participated in the thorough review of every paper. We would also like to extend our gratitude to the people we were in contact in PENSOFT and St. Kliment Ohridski University Press, especially Dr. Lyubomir Penev (Managing Director of Pensoft) and Teodor Georgiev (Chief of Preprint Department) for meticulously working with the authors and editors in putting together the final touches of the book. Nadja Ognjanova-Rumenova & Kalina Manoylov

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Prof. Dr. Biol. Sc. Dobrina N. Temniskova-Topalova

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Nadja Ognjanova-Rumenova & Kalina Manoylov (eds.) 2006 ADVANCES IN PHYCOLOGICAL STUDIES Festschrift in Honour of Prof. Dobrina Temniskova-Topalova (pp. 13-24) © PENSOFT Publishers & University Publishing House Sofia–Moscow

Prof. Dr. Biol. Sc. DOBRINA N. TEMNISKOVA-TOPALOVA (A life dedicated to the students) Dimitar Vodenicharov1, Kalina Manoylov2 & Nadja Ognjanova3 Han Asparuh str. 88, Plovdiv, Bulgaria Michigan State University, Department of Zoology, 203 Natural Sciences Bldg. East Lansing, MI 48824, USA 3 Institute of Geology, Bulgarian Academy of Sciences, Acad. G. Bonchev str. 24, 1113 Sofia, Bulgaria 1 2

Unexpectedly for her active mentors and her students Prof. Temniskova jumped over a life long barrier that allowed her to celebrate her 70th Birthday anniversary. She managed to preserve the charm and temperament of her student years for most of her life. Maybe the biggest reasons for her energy are the beautiful algae she has studied for 40 years and still finds extremely exciting, the mornings around the pristine lakes of Pirin and Rila Mountains, and evenings around a camp fire full with intellectual conversations, endless funny stories and new project ideas for tomorrow. Happy anniversary Prof. Temniskova! We wish you health, continuous endless energy, drive and success in the future! Dobrina Nikolova Temniskova was born on November, 12 1934 in the historic and cultural center Veliko Turnovo. Veliko Turnovo brings pride in every Bulgarian as the capital of the Second Bulgarian Kingdom (1187–1396), and an energetic university town today. Prof. Temniskova still loves her birth town and in her academic speech in front of the Academic Board of the Sofia University. (21 January 2004) she said:… ‘for all my life, the beauty of my birthplace, its energizing atmosphere, together with my family, my friends and mentors gave me strength and inner vigor’. She completed her elementary and secondary education in V. Turnovo. Her favorite subjects were biology, science and Bulgarian language and literature. During her high school years she played violin in the school orchestra, participated in theatre and literature clubs. When she was in 11th grade she won second place for literature from the Ministry of Education of 13

Dimitar Vodenicharov, Kalina Manoylov & Nadja Ognjanova

Bulgaria. In 1952 D. Temniskova graduated from High School and started her undergraduate studies in the Biology-Geology- Geography Faculty of the Sofia University ‘St. Kliment Ohridski’, Biology major. She specialized in the department of Plant Systematics and Plant Geography (today department of Botany) and in 1957 she graduated with honors. Prof. Temniskova is proud that she specialized (magistrate) in one of the oldest departments of Sofia University and the oldest in the Biology Faculty. The Botany department was established in 1891, three years after the establishment if the first school of Higher Education in Bulgaria, named Bulgarian University in 1904. This is the first University in Bulgaria after the 500 years under Turkish rule (1396–1878). In the Botany Department the beginning of Phycological studies began with Prof. St. Petkov (Doctor of Sciences from the University of Gent, Belgium) and chair of the Botany Department for many years. The University mentors of D. Temniskova were Prof. D. Vodenicharov, acad. Daki Yordanov, coresp.member BAS A. Vulkanov, Prof. K. Popov etc, most of whom graduated with Doctoral degrees from Western European Universities prior to Word War II. Prof. Temniskova many times had expressed her gratitude to her teachers and she didn’t forget them in her Academic speech when she was awarded the prestigious Blue Ribbon from Sofia University:…’I am immeasurably grateful to my mentors. They showed me what a university professor has to be – not only a great researcher, but an excellent and engaging teacher, with great professionalism, wide scientific and cultural knowledge, a person with decency, work ethics and energy.’ As a graduate student Prof. Temniskova showed great interest in algae. She participated in the Botany club, student scientific forums and in 1957 received Second place in science at the First Republican festival of student sciences in Bulgaria. After her graduation D. Temniskova was sent by the Government to work for 3 years at the town of Samokov (DIP“ Rilski Len”, 1957-1959), she worked for 2 years as a lab manager of the central factory lab and then for 2 years in DIP ’Uchtechprom’ department of Microscope slides. She organized the first factory production of permanent microscope slides with educational purposes in botany, zoology, and human anatomy (1960–1961).

Doing her dissertation research.

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Prof. Dr. Biol. Sc. Dobrina N. Temniskova-Topalova (A life dedicated to the students)

In 1962, D. Temniskova started as an adjunct assistant (free aspirant) in the Botany Department, Faculty of Biology, Sofia University ‘St. Kliment Ohridski’, where in 1964 she continued as a regular assistant. In 1972, as a free aspirant she defended her dissertation and became ‘Doctor of Sciences’, later in 1994 she completed a second (habilitation) dissertation and was awarded ‘Doctor of Biological Sciences’. In 1966, Dr. D. Temniskova had an opportunity to continue her studies in Germany with a Humbold stipend, but the political circumstances in Communist Bulgaria during this period were hard to overcome. Regardless of the political barriers and thanks to her love to her research, graduate students and teaching she continued her work at Sofia University. Luckily, later D. Temniskova had the opportunity to specialize at the St. Petersburg diatom lab at the Saint Petersburg University, the Botanical Institute of Academy of Sciences of USSR. She studied with Prof. D.Sc. V. S. Sheshukova-Poretzkaja, Prof. D.Sc. N. I. Strelnikova, Prof. Dr. T. F. Kozyrenko, Prof. D.Sc. I. V. Makarova, Prof. D.Sc. A. I. Moiseeva and many others (1968-1969). D. Temniskova specialized in the Harkov University ‘T. Shevchenko’ with Prof. D.Sc. A. M. Matvyenko, student of the great phycologist from Ukraine, Prof. Korshikov. From all her studies abroad and especially as gifts from her colleagues from St. Petersburg University and Athens University, Greece (Prof. Dr. K. Anagnostidis and Prof. Dr. A. Economou-Amilli) she collected extensive literature on diatoms for her research and her student’s research. D.

International Symposium: Biology and Taxonomy of blue-green algae, Smolenize Castle, Republic of Czechoslovakia (1987).

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Temniskova had mentioned in many occasions that without the help and the scientific exchange with colleagues from abroad the development of Diatom analyses and Phycology in Bulgaria was never going to be successful. All teaching and research years of Prof. Temniskova were at the Sofia University, Biology Faculty, Department of Botany; 1962 – adjunct assistant, 1964 - regular assistant, 1982 - docent (Associate professor) and from 1995 – professor. She established and was a leader of the Laboratory of Diatom Analyses (1982–2002). From 1996-2002 she was Department Head of the Botany Department. After her retirement (2004) she remained engaged with the university as adjunct professor and mentor of doctoral students. She has been a member, for many years, of the Faculty Board of the Biology Faculty (1987–2002) and a member of the Academic board of Sofia University (1999–2003). She has been active on the Academic boards and her opinions and expertise were sought when decisions were made on the Botanical Garden of the Sofia University and other University events. She is tremendously respected among her academic peers and was deservingly elected as the first ombudsmen in the History of Sofia University (2004). Prof. Temniskova is an excellent teacher, lecturer, and a favorite professor. Her classes have been required classes in the Botany department and some specialized classes. She had laboratory and field practices in morphology and systematics of lower plants, phytogeography, and she lectured on systematics of algae and fungi; phycology with diatom analyses, taxonomy and evolution of algae, etc. Prof. Temniskova created a diverse collection of permanent diatom slides preserved algal and fungal specimens. Her extensive collection has been used in the educational laboratory practices, scientific observations and learning in the

International Diatom Symposium, Budapest, Hungary (1980)- from left to right Dr.G.Khursevich, Dr. B. Marciniak, Dr. D.Temniskova, Dr. R.Ross, Dr. I. Makarova, and Dr. L. Loginova).

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Botany Department. She started a Herbarium collection of permanent diatom slide from hers and her student’s research. For the first time in Bulgaria she developed and taught a course in Diatom analyses (lectures and lab work). The slide collection, microscopes, and literature available in the lab contributed to the future studies and successes of her student working with recent and fossil diatoms (N. Ognjanova, M. Vuleva, S. Passy, K. Manoylova, R. Stancheva, P. Ivanov, R. Zidarova). She started, maintained and developed the only laboratory for Diatom Research in Bulgaria especially in the field of Paleodiatomology. Fossil studies with diatoms had been the most dynamic branch of studies in Bulgarian Phycology in the last decade. Many of Prof. Temniskova’s students, who graduated with M.Sc. degree, work in different fields of Botany (D. Ivanov – palinology, P. Robeva - embryology, K. Uzunova - anatomy, A. Uzunova – plant physiology, N. Misaleva – plant genetics etc) and many of her Masters students work in the pharmaceutical and food industries. As a mentor D. Temniskova left a shining trail of successful scientists, all touched and inspired by her scientific approach, enthusiasm, knowledge and personal conviction. D. Temniskova loved lecturing and teaching. She never considered working at other prestigious institutions like the Bulgarian Academy of Sciences, because of she couldn’t part with

Celebrating the Day of Slavic (24.May.2002) – from left to right Prof. B. Biolchev (Rector of Sofia University), prof. D. Temniskova, prof. V. Radeva and prof. O. Gerdzhikov (National Assembly Chairman).

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her students. She confesses this: …‘I was unable to imagine living without the magic of teaching. I loved looking at excited student eyes when they understood a process or relationship, discovered a new species or got excited assigning a correct scientific name. In my priorities after my family, my teaching and work, I have close to my heart all my masters and doctoral students. They will be part of me forever. They are my pride and joy that energizes me’. That is why Prof. Temniskova left a memorable part of herself in all her students, grateful of their knowledgeable and inspiring mentor, and not daring even to dream to measure as colleagues with her one day in the future. As a token of her students’ appreciation, of the respect of the academic body of the Sofia University and the scientific community of the University, Prof. Temniskova had been warmly congratulated for her 70th year anniversary and awarded with many prestigious awards. She was awarded the Honorary Sign of Sofia University level 1, Honorary Sign of Sofia University with blue ribbon (2003) celebrating 40 years of the establishment of the Faculty of Biology and awarded the Icon of Saint Kliment Ohridski from the Rectorate of Sofia University (2004). Prof. Temniskova started her research as an undergraduate student. This resulted in her first publication (1961): Algal flora of the rivers Iskur and Palakarya in the man-made lake Iskar. In her paper she discussed representatives of Chlorophyta, Zygnemophyta, Cyanobacteria, Chrysophyta and Xanthophyta. From that time on algae become her favorite and main tool of research. For most of her career Prof. Temniskova addressed different Phycological questions, and her research can be divided into two major periods. In the first period of her research she studied algae from different groups and different water bodies, with major emphases on temporary water bodies. As a result she reported 632 species, varieties and forms from seven algal divisions: Chlorophyta, Euglenophyta, Bacillariophyta, Chrysophyta, Zantophyta, Dinophyta and Award ceremony when the Honorary Golden Sign of Sofia University with blue ribbon was awarded (2003) - Prof. B. Biolchev (Rector of Sofia University) and prof. D. Temniskova.

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Cyanobacteria. From those, 239 taxa were reported for the first time in Bulgaria. She studied 748 temporary water bodies from 106 sites in Bulgaria, provided data on temperature, pH, chemical content of water, time period of existence, drying processes etc. She presented algal species composition, classification of the algal bodies and characteristics of the algal bodies first in Bulgaria. She presented her results with her first dissertation ‘Algal flora from temporary waters in Bulgaria’; she defended her work as a free aspirant and received a Ph.D. degree (1972). It is noteworthy to mention that for the Dissertation work of D. Temniskova there were official opinions from 14 European algologists: G. Deflandre, T. Hortobagyi, G. Jerkovic, M. M. Gollerbach, A.M. Matvyenko, V. S. Sheshukova-Poretzkaya, I. B. Makarova, Z. I. Asaul and others). They all gave high praise of her research as they were looking at it from different algal groups’ perspectives. We will site only Prof. G. Deflandre (honorary member of the French Academy of Sciences Paris, honorary member of the Austrian Academy of Sciences, member of the Royal Academy Belgium, fellow of the German Academy, Hale Germany) who in his opinion states:…’I have been studying algae for more than 10 years and I can attest for the enormous amount of work done by D. Temniskova to show this results. As far as the science in the work, many times I felt I am reading my own thoughts, taken from my unpublished manuscripts, results and interpretations I came as a result of shorter and way too shallow observation than those presented by D. Temniskova. With this review I want one more time to state that the presented work is in line with my own ideas and plans for Phycological research, what I have neglected in lately as I have been doing mostly micropaleontology. I agree with the presented classification of temporary waters presented by D. Temniskova, with the presentation of continuous historical perspective and a helpful discussion on the terminology of those waters’. During the first period of her research Prof. Temniskova did extensive research of floristic, taxonomic and horologic characteristics of some flagellate algal groups like Euglenophyta and Volvophyceae (Chlorophyta) from different waters. D. Temniskova was a coauthor of the first algal flora of Bulgaria –‘Flora of Bulgaria: algae’ (1971). During this time Prof. Temniskova promoted science for the general public with the publication of “Plantsdetectives’ (1964) and numerous educational papers in magazines and newspapers like ‘Plants-geologists’, ‘The extinct gentian’ etc. The second period of D. Temniskova’s scientific research has been dedicated to the diatoms (Bacillariophyceae) and the development of Diatom Analyses as a tool in Bulgaria. Today, the current accomplishments in the Bulgarian Diatom Analyses are determined by her research and the research of her students. In her diatom research there are several directions: As a result of taxonomic investigations D. Temniskova introduced a new genus, Pontodiscus, three species P. baldjickianus, Actinocyclus makarovae, Actinocyclus fungiformis and seven within species diatom taxa. There are 34 new taxonomic combinations. She uses biometric methodology for within species taxonomy of Aulacoseira granulata. She also studied variability in non-diatom populations like the variability within two species of Trachelomonas (Euglenophyta). She studied with SEM numerous recent and fossil diatoms, important in biostratigraphic and paleoecological research. The most prestigious and numerous are the contributions of Prof. Temniskova to the studies of fossil diatoms. Most of her work was summarized in the Dr.Sc. thesis 19


(Dissertation): ‘Miocene diatom floras from Bulgaria-community composition, structure, evolution, paleoecology and biostratigraphy’ (in 3 parts: 1. 318 pp text, 2. 120 plates with 1236 LM and SEM original micrographs of fossil diatoms and 3. 4 maps, 30 tables and 111 graphs, diagrams, and dendograms). The major contributions of Prof. Temniskova in Paleodiatomology are in several areas: paleofloristic and taxonomy, biostratigraphy, paleoecology, paleobiogeography and evolution of diatoms. In those directions she characterized many florotypes: Chokrakian, Karaganian, Konkian, Volhynian, Bessarabian, Chersonian, Pontian and Pont-Pliocene. This encompasses a broad paleosuccession cycle for the Chokrakian-Chersonian stages and the narrower Pont-Pliocene stages. From the research on diatoms from Neogene marine-brackish and continental sediments, paleoflora of Bulgaria was enriched with 845 taxa (535 species, 268 varieties and 42 forms). For the first time 1 genus with 3 species of Archeomonada, 2 species and 2 varieties of Silicoflagellatae, representatives of type Poryfera (3 species) and one Phytolitaria were identified. Research on late glacial and Holocene sediments was performed (over 200 species and varieties) as a result of the first studies of Durankulak Lake and Beloslav Lake, Varna District. As contributions to the European paleoflora: there are 182 species, varieties and forms of diatoms new for the Miocene diatom floras from the Euxinian Basin. Three genera and 86 species, varieties and forms, new for the Miocene diatom flora of the Whole Paratethys were reported. Also reported for the first time, were archeomonads in East Paratethys. Based on these rich data sets Prof. Temniskova for the first time in Bulgaria divided and characterized diatom floras from the Chokrakian, Karaganian, Konkian, Volhynian, Bessarabian and Chersonian age. She followed changes in diatom flora for 5.5 million years (16-10.5 million years BC) in the Northwest part of the Euxinian Basin. She uncovered the florogenesis during the Middle-Late Miocene and proved that the Miocene flora played a significant part in the formation of the modern floras of the Black, Azov and Caspian Seas. Her largest contribution to the modern communities was her work from Chersonian flora in the Black Sea - Caspian Sea basin. She described a repeated sequence in the evolution of diatoms for this time period. During this geological interval most of the modern diatom genera evolved. She determined the speed of macro evolutionary processes for the Middle and Late Miocene diatoms. The highest speed of evolvement of new genera was during the Karaganian stage (century) of the Middle Miocene, 2.26 genera/ 1 million years. Prof. Temniskova completed the first, for our country, biostratigraphic subdivision of the Miocene sediments (Chocrakian, Karaganian and Konkian stages and Volhynian, Bessarabian and Chersonian substages of the Sarmatian stage in Northeast Bulgaria) based on diatoms. She separated and characterized 5 taxon acro zones, 3 taxon acro subzones, one interval zone, and one barren interval zone for marine-brackish sediments. Those are the first diatom zones for the Miocene sediments of the Eastern Paratethys paleobasin. Very important is the parallel correlation with other biostratigraphic zones, based on different fossils groups: mollusks, foraminifers and ostracods. For the first time Prof. Temniskova compared the Late Miocene diatom floras from the Eastern and Central Paratethys and Tethys. She completed the biostratigraphic, chronostratigraphic and magnitostratigraphic units in the Tethys, Eastern and Central Paratethys. 20


Prof. Temniskova carefully researched the Neogene continental basins in South Bulgaria. She established two types of diatom floras during the Late Miocene (Pontian) and Early Pliocene (Dacian): type ‘Aulacoseira’ and type ‘Actinocyclus’. She underlined the specificity of the type ‘Actinocyclus’ and presented the hypothesis that during the Late Miocene, the SouthBulgarian continental basin was one of the world’s centers of evolution of the genus ‘Actinocyclus’. She introduced a novel hypothesis for the possible ways of distribution of diatoms during colonization of the Neogene nonmarine basins in Bulgaria. Based on the diatom analyses she established the age of South-Bulgarian Neogene sediments. With this she proved that in different basins, sediments were deposited at the same chronological interval Late Miocene (Pontian) - Early Pliocene (Dacian). For the nonmarine Upper Miocene - Lower Pliocene sediments of the Sofia Basin there were distinguished 4 diatom taxa acro zones. Her contributions to paleoecology continued with the establishment of the character, dynamics and sequences of paleoecological diatom spectra from different stratigraphic levels in the Middle and Upper Miocene in different paleobasins. She reported a well defined reconstruction of the paleoecological conditions in different stages and sub stages of the Middle-Late Miocene. Based on changes in the weighted mean value of every halobic diatom group she determined the cycles of increase and decrease of the salinity of the waters in the studied basins. The results of the paleoecological analyses of different types of floras are diverse, but they summarized the full ecological characteristics of biotopes in different consequent periods of development of the diatom paleofloras during the Miocene stage.

16th International Diatom Symposium, Greece 2000, from left to right Drs: Kalina M. Manoylova, Nadja Ognjanova-Rumenova, Prof. Dobrina Temniskova-Topalova and Sophia Passy.

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A complete reconstruction was presented of the paleoecological conditions of the Durankulak and Beloslav Lakes during Late Glacial and Holocene times. She presented that the Durankulak Lake was part of the littoral zone of the Black Sea and went through three periods of salinity in its development. During those periods the habitat was brackish – marine (modern salinity of this Lake is 4‰). Similarly 3 periods in the salinity of the Beloslav Lake in northern Bulgaria were determined. The introduction of Diatom analyses in the geological practices in Bulgaria has been very important in the understanding of the genesis of depositional rocks, and for the practical answering of biostratigraphic, correlational and paleoecological questions. Biostratigraphic and paleoecological results of Prof. Temniskova have been used in the publication of the geologic map of Bulgaria (M 1:100000), also in the assessment of different Neogene basins in Bulgaria. Using diatom species composition there have been several assessments of diatomite and diatomite sediments in different regions in Bulgaria. Those assessments are important for future exploitation of different sediments. Her work also contributed to the use of electronic paramagnetic resonance in diatomite research. Most of the scientific research of Prof. Temniskova also has application in student education. Most of the published research of Prof. Temniskova can be used in the understanding of applied ecological problems. Many publications have been on the anthropogenic pollution of water bodies and the use of diatoms as bioindicators. Those publicaProf Temniskova with a group of students leading practicum “Ecology of algae”, Kaliakra Cape, the Black Sea (2003)

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tions are an important contribution to the application and use of biomonitoring in ecosystem management. She contributed tremendously in the taxonomic variability of diatoms in recent lakes and rivers from different regions in Bulgaria (Rhodope Mountains, Strandza Mountains, Ograzhden Mountain etc). For the classification of freshwater basins ecomorphs of Aulacoseira granulata have been used. Prof. Temniskova pioneered research on benthic diatoms from the Bulgarian Black Sea shelf: taxonomic composition (reported over 250 species, varieties and forms), changes in biomass, dominant species, and mean density. Research was done over a 5 year period (May 1986- December 1990) immediately after the Chernobil accident in Ukraine. She studied the first cores from the deep Black Sea shelf. She described 15 complexes (communities) of diatoms in the shallow and deep shelf. Especially she related anthropogenic influence (increase in heavy metals and radionucleotides) and established an indicator species (Podosira pellucida Pr-Lavr.). Her research has been a model as the diatom composition and structure of marine benthic communities can be used in understanding the eutrophicaion relationships and changing ecological condition in marine ecosystems. Temniskova studied for the first time silicoflagellates and representatives of Ebridae from recent sediments in the Bulgarian Black Sea Shelf. She reported 2 new genera, 3 new species and 1 variety new for Bulgaria. For the first time for the modern Black Sea community she reported the living fossil Dictyocha triacantha Ehr. (Dictyophyta = Silicoflagellatae). In conclusion, through her research on recent algal flora from Bulgaria ranging from diatoms, green algae, euglenophytes, blue-green algae, and silicoflagellates, she contributed to the describing of algal biodiversity in Bulgaria. She reported for the first time 7 genera and 610 species, varieties and forms of algae. Prof. Temniskova contributed to the algal studies in Irak, for the territory of ancient Babylon, together with Prof. E. Blaženčiæ (University of Belgrade, Serbia) she reported 5 Charophyta species. In recent years the collaboration continued with a project on Charophyta from Serbia, Montenegro and Bulgaria. In the last decade Prof. Temniskova has been a part of a team study of Antarctic flora and fauna on Livingston Island where Sofia University has built a research Base named after St. Kliment Ohridski. She works on materials collected from Bulgarian Antarctic expeditions, with participation of Biologists, including a student of hers. Diatoms from continental water and terrestrial biotopes have been studied and 190 taxa have been reported. The species in this community are mainly cosmopolitan, where arcto-antarctic and northern alpine species were rare. Diatom communities were dominated by the cosmopolition and, typical for the Antarctic communities, Luticola muticopsis (V. H.) D. G. Mann. Preliminary data has been published on blue green algae and green algae, distributed mainly in water and terrestrial habitats in the Bulgarian and Spanish bases on Livingston Island. The identified representatives are mainly cosmopolitan. Prof. Temniskova reported that there was no difference between the species living in aquatic and terrestrial habitats. This conclusion was similar to that reported in the literature, that clear boundaries of aquatic and terrestrial habitats in extreme environments are hard to distinguish and the same species live in both habitats. 23


In Antarctic communities green algae (reported 33 species from 21 genera) were represented with typical cryophytes and aerophytes, peat-loving and rheophilous mountain species, as well as terrestrial algae. Of the habitats studied, the snow and glacier ones have a specific flora. Lake habitats manifest a greater variety of representatives of class Zygnemophyceae, and the streams – Ulothrix spp., Zygnema spp. and Prasiola calophyta (Corm.) Meneghini. As expected, Prasiola crispa (Lightf.) Meneghini grew abundantly on rocks close to the penguin colonies. The research on different algal groups from Antarctic is important contribution to Phycology of the vastly unknown habitats from this continent. Prof. Temniskova was a principle investigator and participant in many national and international projects (projects 25, 158B and 329 UNESCO, project on complex research on Livingston Island, Antarctica, etc). Prof. Temniskova has participated in many national and international conferences, symposia and congresses. She presented more than 50 papers and posters. Prof. Temniskova is a member of many Bulgarian Scientific societies including the Bulgarian Botanical Society since 1964, Union of the Sciences of Bulgaria from 1966, and the Society of the Bulgarian Ecologists since 1999. Since 1999 she has been the president of the Bulgarian Botanical Societies, where she has an active and supportive role bringing the Society to new and fruitful scientific levels. Dr. Temniskova is a member of the International Phycological Society, and the International Society for Diatom Research. Prof. Temniskova has been a member of many scientific boards and expert working groups in Bulgaria: Committee of Medico-biological sciences with the High Attestation Committee (VAK) at the Ministry Council of Bulgaria, Specialized Council in botany and Mycology at VAK (1995-present), Scientific council of the Central laboratory in General Ecology at the Bulgarian Academy of Sciences (1999-present), Science-coordination center for Global projects at the Bulgarian Academy of Sciences (2004-present). Prof. Temniskova is the editor of the Yearly Scientific Journal of Sofia University ‘St. Kliment Ohridski’. All scientific research publications of Prof. Temniskova are significant and essential contributions to the development of Phycology and especially Paleodiatomology in Bulgaria. She introduced the beginning of systematic and modern studies of diatoms in Bulgaria with recent and relict aquatic habitats. Prof. Temniskova has presented many original and applied scientific contributions. Results from her studies have nation, European and World importance, especially the original classification of temporary water bodies; the new taxa described; the biostratigraphic subdivision and correlation, based on diatoms from the Miocene sediments from the Northwest parts of the Eastearn Paratethys and the correlations of the biostratigraphic zones of the whole Eastearn and Central Paratethys and Tethys.

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Two new sigmoid, naviculoid diatoms from the Eocene of Kazakhstan N. I. Strelnikova1, J. P. Kociolek2 and E. Fourtanier2 Department of Botany, Biological Faculty, St. Petersburg State University, Universitetskaya Emb. 7/9, St. Petersburg, 199034, Russia, e-mail: [email protected] 2 Diatom Collection, California Academy of Sciences, 875 Howard St., San Francisco, CA 94103-3098, U.S.A. 1

ABSTRACT A new genus of sigmoid, naviculoid diatoms Hyalosigma gen. nov. and two new species H. temniskovae-topalovae sp.nov. and Gyrosigma kazachstanicum sp.nov. were studied with light and scanning electron microscopy and described from Lower Eocene-Early Middle Eocene diatomite quarry Kirgizskoe, Emba River basin, Kazakhstan. Gyrosigma kazachstanicum differs from other members of the genus by its convex valve and round external areaolar openings. Hyalosigma differs from Gyrosigma and Pleurosigma by having ornamentation of the striae interrupted by large hyaline areas on the valve face and acute angle of marginal and mantle striae. These records are the oldest known for the sigmoid group of naviculoid diatoms. Key words: Sigmoid, naviculoid diatoms, Hyalosigma gen. nov., Hyalosigma temniskovaetopalovae sp. nov., Gyrosigma kazachstanicum sp. nov., Paleogene sediments, Kazakhstan

INTRODUCTION Very little is known about pennate diatoms from the Cretaceous and Paleogene Epochs. Scattered descriptions of pennate diatoms from these geological epoch can be found in the literature (Weisse 1854, Witt 1886, Grove and Sturt 1886-1887a-c, Pantocsek 1903-1905, Schrader and Fenner 1976, Aphanassyeva and Gleser 1986, Harwood 1988, Desikachary and Sreelatha 1989, Fenner and Mikkelsen 1990, Edwards 1991, Fenner 1991, 1994, Strelnikova 1992). The most extensive study is that of H.J.Schrader (1969) who described pennate diatoms 25

N. I. Strelnikova, J. P. Kociolek and E. Fourtanier

from the Upper Eocene/Lower Oligocene deposits of Oamaru, New Zealand. A total of 313 species from 52 genera of pennate diatoms are known from the Cretaceous and Paleogene sediments. Of those forms reported, araphid genera represented by the most species include Sceptroneis (21). Rhaphoneis (17), Synedra s. l. (11), while in the raphid group the genus Navicula s. l. is represented by the most taxa (69 species), followed by Amphora (26), Cocconeis (24), Diploneis (18), and Nitzschia (16). Of the raphid forms reported from Oamaru, only a few have been investigated with SEM (e.g.Novitski et al, in press). Of members of the sigmoid naviculoid, only two taxa have been reported: Donkinia antiqua Grove & Sturt (1887c) from the Oamaru (Eocene-Oligocene) and Gyrosigma ? sp. (Gleser et al. 1965) from the Late Eocene of Ukraina. In this report we describe two new sigmoid naviculoid diatoms and document their morphologies with light and scanning electron microscopy. MATERIALS AND METHODS Material observed herein is from a diatomite quarry in Kirghizskoe, Kazakhstan, from the Emba River basin. Samples were received by the senior author from the Geological Institute of Russian Academy of Scienses, Moscow, from Drs V.N.Beniamovski and E.P.Radionova. Samples examined have been accessioned in the Diatom Collection of the California Academy of Sciences as follows: Sample Number 19/422 22/425 23/434 29/440

CAS Diatom Accession Number 624811 624813 624814 624815

The diatomite belongs to the Ypresian stage and is dated as Lower Eocene, determined by stratigraphic position ( dates by V.N. Beniamovski) and confirmed by diatom analysis. Samples 19, 22 and 23 are from diatomite alternating with diatomitic clay and glauconitic sand and diatomite with sand lenses, belongs to the Pyxilla gracilis zone and sample 29 from silted clays with an admixture of sand, and belong to the Pyxilla oligocaenica var. tenuis zone. The sequence of Kirghizskoe was established by Gleser (1979) as a stratotype of Pyxilla gracilis and P. oligoceanica var. tenuis zones dated as Late Eocene. Later Gleser et al. (1997) dated these zones as Late Early Eocene - Middle Eocene. The age of these zones are under disscussion. Strelnikova (1992) believed the age of P. gracilis and P. oligocaenica var. tenuis zones to be Early Eocene. According to new dates (Radionova et al. 2003, Akhmetiev and Beniamovski 2004), the age of P. gracilis zone is Early Eocene and corresponds to the nannoplankton zone NP12 (CP10) while the P. oligocaenica var. tenuis zone is Early EoceneEarly Middle Eocene and corresponds NP13-NP14 (CP11-CP12) nannoplankton zones. Light microscope observations were made with a Leitz DMRB (CAS). For SEM observations, individual specimens were selected according to the procedures described by Nikolaev (1982). SEM observations were made on a Leo-1450VP (CAS). 26

Two new sigmoid, naviculoid diatoms from the Eocene of Kazakhstan

RESULTS AND DISCUSSION Pennate diatom assemblages. The samples investigated here have a rich, well-preserved flora of marine planktonic centric diatoms and several pennate forms. Well represented in the samples are Oestrupia powellii (Lewis) Heiden, Clavicula polymorpha Pantocsek, Navicula hennedy f. granulata Grun., Sceptroneis caducea Ehrenb. Amphora sp., Amphiprora sp., Nitzschia sp. and Rhaphoneis sp. The presence of these pennate diatoms and sand lenses indicate a near shore, shallow depositional environment. In addition to the forms listed above, we encountered two sigmoid, naviculoid diatoms that appear to be new to science, including a new genus. Descriptions of these two new forms are presented below. Class Bacillariophyceae Order Naviculales Family Naviculaceae Genus Gyrosigma Hassal Gyrosigma kazachstanicum, sp.nov. Strelnikova et Kociolek (Figs 1-25). Holotype: CAS slide number 581071 Type material: CAS Accession number 624815 Holotype is Figure 1. DESCRIPTION: Valves sigmoid, lanceolate, valve face slightly convex, valve margin with hyaline line. Length 144-270 µm, breadth 15-27 µm. Areolae in transverse rows 15-21/10 µm. Openings of the areolae are round both externally and internally. Raphe slightly sigmoid, excentric at the end but in the middle of the valve at the center. External raphe fissures are curved to opposite sides at the apices. External proximal raphe ends simple, straight, without deflections, located in a small central area. Internal proximal raphe ends are positioned on a hyaline ridge, which has a lunate form in the center. Internal distal raphe ends are helictoglossae. Girdle hyaline – structure not discerned. COMMENTS: The suite of features that assign this species to the genus Gyrosigma include sigmoid form of the valve, transverse striation pattern of the areolae, and striae pattern without a diagonal axis (Sterrenburg 1989, 1991, 1992, 1993, Stidolph 1988, 1992, 1994). It differs from other known species of the genus by the convex form of the valve and external openings of the areolae being round, instead of slit-like. The convex valve structure is reminiscient of Plagiotropis Pfitzer (Paddock 1988), but G. kazachstanicum differs from Plagiotropis species by the lack of a raised valve face with raphe and by its pronounced sigmoid outline of valve and raphe. This species was noted from all four samples examined in this study.

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2

1

3 Figs 1-3. Gyrosigma kazachstanicum Strelnikova & Kociolek, LM. Scale bars =10 µm. Fig. 1. Valve view, holotype. CAS 624815, slide 581071. Figs 2,3. Valve views showing structure of central area. CAS 624815, slide 581071 (Fig.2); CAS 624813, slide 581069 (Fig.3).

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5

4

6

7 Figs 4-7. Gyrosigma kazachstanicum Strelnikova & Kociolek, LM, Scale bars = 10 µm. Figs 4, 5. View of frustule from girdle. CAS 624811, slide 581066. Figs 6,7. Valve views showing structure of central area. CAS 624815, slide 581071.

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8

9

10

11

12

Figs 8-12. Gyrosigma kazachstanicum Strelnikova & Kociolek, SEM. Fig. 8. General aspect view. Figs 9, 10. External view of valve terminus. Fig. 11. External view of central area showing raphe. Fig. 12. Broken valve, showing locaular areolae and raphe structure. Figs 8-11. CAS 624814. Fig. 12. CAS 624815.

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14

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15

Figs 13-15. Gyrosigma kazachstanicum Strelnikova & Kociolek, SEM. View of valve from girdle view. Scale bars: 13 = 10 µm; 14, 15 = 3 µm. Fig. 13. General aspect view. Fig. 14. External valve margin with hyaline girdle element. Fig. 15. External view showing round areolae and curved distal raphe end. Internal view of valve end shows prominent helictoglossa. Figs 13-15. CAS 624813.

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16

17

18

19

21 20

Figs 16-21. Gyrosigma kazachstanicum Strelnikova & Kociolek, SEM. Figs 16-18. Internal views showing rounded areolae. Fig. 19. Internal view showing elevated central nodule. Fig. 20. Broken valve showing locaular arrangement of areolae. Fig. 21. External view of central area showing narrow proximal raphe ends and central hyaline rib bordered by shallow grooves. Figs 16,19, 20. CAS 624814. Fig. 17. CAS 624815. Figs 18, 21. CAS 624811.

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22

23

24

25

Figs 22-25. Gyrosigma kazachstanicum Strelnikova & Kociolek, SEM. Fig. 22. Whole frustule, general aspect. Fig. 23. External view of valve mantle and hyaline girdle element. Figs 24, 25. Terminus of frustule with rounded areolae and external distal raphe ends curving onto valve mantle. Figs 22, 24, 25. CAS 624811. Fig. 23. CAS 624814.

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Class Bacillariophyceae Order Naviculales Family Naviculaceae Genus Hyalosigma, gen. nov. Strelnikova et Kociolek DESCRIPTION: Valves flat, in their apical axis sigmoid, linear-lanceolate in shape, with sigmoid raphe system and striae interrupted by large hyaline areas on the valve face. Marginal and mantle striae at acute angles. GENERITYPE: Hyalosigma temniskovae-topalovae, sp.nov. Strelnikova et Kociolek Hyalosigma temniskovae-topalovae sp.nov. Strelnikova et Kociolek Figures 26-51 Holotype: CAS slide number 581070 Type Material: CAS Accession number 624814 Holotype: Figs 26-29 DESCRIPTION: Valves flat, valve face curved smoothly into a very shallow mantle which is scarcely distinguishable. Valve linear-sigmoid with subacutae ends. Length 196-280 µm, breadth 11-23 µm. Proximal raphe ends located in the center of the valve within a small hyaline central area. Raphe ends only slightly deflected in the same direction, raphe branches highly sigmoid, curved along valve margin and towards the center of each apex. Internally, the distal raphe ends are helictoglossae. The raphe is contained in narrow axial area. Striae along the axial area are parallel to convergent, and number 18-23/10 µm. On the side of the valve opposite the raphe branch, beyond the striae comprising the axial area, there is a large hyaline area that extends to almost the apex of the valve and tapers to a point past the central nodule. This hyaline area is bordered along the margin by another group of striae that run the entire length of the valve, equal in density to the striae along the axial area. In the SEM, areolae of these striae are arranged in acute angles to the length of valve. The naviculoid organization of valve, sigmoid shape of the valve and sigmoid raphe suggest a close affinity of this new genus and species with Pleurosigma W.Smith or Gyrosigma Hassal. The interrupted striae on the valve face and acute angle of the marginal and mantle striae separate this form from either previously known genus. This taxon occurred in all four samples examined in this study. We dedicate this new species of diatom to Dr. Dobrina Temniskova-Topalova, on the occasion of her 70th Birthday. These observations extend the known geological range for Gyrosigma, which was previously known from the Late Eocene epoch (Gleser et al. 1965) and suggest the sigmoid diatoms were morphologically more diverse than previously understood. These new taxa underscore we still have a tremendous amount of work to do to understand the diversity and history of pennate diatoms, especially the earliest known raphid diatoms. 34


26

27

29

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Figs 26-29. Hyalosigma temniskovae-topalovae Strelnikova & Kociolek, LM, Scale bars = 10 µm. Fig. 26-29. Holotype. Fig.26. Valve view, plane of focus on the center of the valve. Figs 27, 28. Focus on the ends of the valve showing wide hyaline areas. Fig. 29. Valve center showing small external proximal raphe ends and axial area bordered by jagged-edged hyaline areas on either side. Figs 26-29. CAS 624814, slide 581070.

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Figs 30-33. Hyalosigma temniskovae-topalovae Strelnikova & Kociolek, LM, Scale bars = 10 µm. Fig. 30. Valve view showing sigmoid nature of valve and raphe. Fig. 31. Valve terminus showing raphe in axial area and wide hyaline area. Fig. 32. Central area of valve. Fig. 33. Internal linear-shaped central nodule. Figs 30-33. CAS 624814, slide 581070.

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35

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39 Figs 34-39. Hyalosigma temniskovae-topalovae Strelnikova & Kociolek, SEM. Fig. 34. General aspect view. Figs 35, 37. Ends of valve with raphe along margin. Figs 36, 39. External view of valve center with small proximal raphe ends positioned close together. Fig. 38. Broken part of valve, showing external and internal surfaces of valves with raphe groove. Figs 34-39. CAS 624814.

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Figs 40-46. Hyalosigma temniskovae-topalovae Strelnikova & Kociolek, SEM. Views showing whole frustule in different positions. Figs 40, 46. General aspect view. Figs 41, 43, 44, 45. Valve face, ends of valve with wide hyaline area. Acute angle of mantle striae is also visible. Fig. 42. View showing central area of valve and “shaped” nature of hyaline area. Figs 40-45. CAS 624813.

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47

48

49

50

51

Figs 47-51. Hyalosigma temniskovae-topalovae Strelnikova & Kociolek, SEM. Views showing whole frustule in different positions. Fig. 47. General aspect view. Fig. 48. External distal raphe end curved onto valve mantle. Figs 50, 51. View of center of valve with raphe becoming more highly arched. Note highly acute angle of mantle striae. Fig. 49. Valve terminus showing internal view with helictoglossa evident. Figs 47,48, 50, 51. CAS624814. Fig. 49. CAS 624813.

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ACKNOWLEDGEMENTS We thank Dr. N.Ognjanova-Rumenova for the invitation to take part on the edition on the honor of Dr. D.Temniskova-Topalova. We also thank Drs V.N.Beniamovski and E.P.Radionova for the interesting material, masters student A.Schvezova for the cleaning material and J.Rubinstein and S.Serrata for the assistance with SEM photomicrographs. The Russian part of work was supported by Grant #03-04-48711 of Russian Fund of Fundamental Research. REFERENCES AKHMETIEV, M. A. and V. N. BENIAMOVSKI 2004. Paleocene and Eocene of Western Eurasia (Russian sector) – stratigraphy, paleogeography, climate. – N.Jb. Geol. Paläont. Abh., 234(1–3): 137-181. APHANASSYEVA, N. I. and Z. I. GLESER 1984. Paleocenovie diatomovie so schvom is Srednego Povoljya. in: Actualjnie voprosi sovremennoy paleontologii, Kiev, Naukova Dumka: 73–76. (in Russian) CLEVE, P.T. 1894. Synopsis of the Naviculoid Diatoms. – Kongl. Svenska Vetenskapsakademiens Handlingar, part I, 26(2): 194 pp. DESIKACHARY, T. V. and P. M. SREELATHA 1989. Oamaru Diatoms. – Bibliotheca Diatomologica, 19: 330 p., 145 pl. J.Cramer, Berlin-Stuttgart. EDWARDS, A. R. 1991. The Oamaru Diatomite. – New Zealand Geological Survey paleontological bulletin, 64: 260 p. FENNER, J. AND N. MIKKELSEN N. 1990. Eocene-Oligocene diatoms in the Western Indian ocean: taxonomy, stratigraphy, and paleoecology. in: Proceeding of the Ocean Drilling Program, Scientific Results, 115: 433–463. FENNER, J. 1991. Taxonomy, stratigraphy, and paleoceanographic implications of Paleocene diatoms. in: Proceeding of the Ocean Drilling Program, Scientific Results, 114: 123–154. FENNER, J. 1994. Diatoms of the Fur Formation, their taxonomy and biostratigraphic interpretation. Results from the Harre borehole, Denmark. – Aarhus Geosciece, 1: 99–164. GLESER, Z. I. 1979. Zonal dismemberment paleogene sediments by the diatoms. – Soviet Geology, 116:19–30. (in Russian) GLESER, Z. I., V. U. ZOSIMOVICH and M. N. KLUSHNIKOV 1965. Diatoms of paleogene deposits in the North Donez basin and their stratigraphic position. – Paleontologichesky sbornik, 2:73–87. (in Russian) GLESER, Z. I., L. A. PANOVA, I. P. TABACHNIKOVA and S. G. VJALOVA. 1997. Correlation marine Eocene of North-West Eurasia according to microphytofossiliens (West Siberia, Povolje). – Stratigraphy, Stratigraphic correlation, 5(4): 35–45. (in Russian)

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GROVE, E. and G. STURT 1886. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Part I. – Journ. Queckett Microscop. Club, ser.II, 16: 321–330. GROVE, E. and G. STURT 1887a. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Part II. – Journ. Queckett Microscop. Club, ser.II, 17: 7–12. GROVE, E. and G. STURT. 1887b. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Part III. – Journ. Queckett Microscop. Club, ser.II, 18: 63–78. GROVE, E. AND G. STURT 1887c. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Appendix. – Journ. Queckett Microscop. Club, ser.II, 19: 131–148. HARWOOD, D. M. 1988. Upper Cretaceous and lower paleocene diatom and silicoflagellate biostratigraphy of Seymour Island, eastern Antarctic Peninsula. – Geological Society of America Memoir, 169: 55–129. NIKOLAEV, V. A. 1982. On the method of preparation of diatoms for studies by light and scanning electron microscopy. – Bot. Zurn. (Moscow&Leningrad), 67: 1677–1679. NOVITSKI, L., J. P. KOCIOLEK and E. FOURTANIER (in press) Preliminary light and scanning electron microscope observations of marine fossil Eunotia species with comments on the evolution of the genus Eunotia. – Diatom Research. PADDOCK, T. B. B. 1988. Plagiotropis Pfitzer and Tropidoneis Cleve, a summary account. – Bibliotheca Diatomologica, 16: 151 p. PANTOCSEK, J. 1903-1905. Beitrage zur Kenntinis der fossilen bacillarien Ungarns. – Berlin, Verlag von W. Junk (2 verbesserte Auflage). Teil 1 - 76 pp, 2 - 122 pp, 3 - 118 pp. RADIONOVA, E. P., V. N. BENIAMOVSKI, A. I. IAKOVLEVA, N. G. MIZYLÖV, V. ORESHKINA, E. A. SHERBININA and G. E. KOZLOVA 2003. Early Paleogene transgressions: stratigraphical and sedimentological evidence from the northern Peri-Tetis. – Geological Society of America. Special Paper, 369: 239–261. ROUND, F., R. CRAWFORD and D. G. MANN 1990. The Diatoms. Biology and morphology of the genera. – Cambridge Univ. Press, Cambridge: 747 pp. SCHRADER, H. J. 1969. Die pennaten Diatomeen aus dem obereozän von Oamaru Neuseeland. –Nova Hedwigia, Beih. 28:124 p. SCHRADER, H. J. and J. FENNER 1976. Norwegian Sea Cenozoic biostratigraphy and taxonomy. Part I. Norwegian Sea Cenozoic diatom biostratigraphy. – Initial reports of the Deep Sea Drilling Project, 38: 921-1099. STERRENBURG, F. A. S. 1989. Studies on tube-dwelling Gyrosigma . – Diatom Research, 4(1): 143-150. STERRENBURG, F. A. S. 1991. Studies on the genera Gyrosigma and Pleurosigma (Bacillariophyceae). Light microscopical critetia for taxonomy.– Diatom Research, 6(2): 367–389. STERRENBURG, F. A. S. 1992. Studies on the genera Gyrosigma and Pleurosigma (Bacillariophyceae). The type of the genus Gyrosigma and other attenuate sensu Perogallo. – Diatom Research, 7(1): 137–155. STERRENBURG, F. A. S. 1993. Studies on the genera Gyrosigma and Pleurosigma (Bacillariophyceae). Rules controlling raphe fissure morphogenesis in Gyrosigma. Diatom Research, 8(2): 457–463. 41


STIDOLPH, S. R. 1988. Observations and remarks on the morphology and taxonomy of the diatom genera Gyrosigma Hassal and Pleurosigma W.Sm. – Nova Hedwigia, 47 (3–4): 377–388. STIDOLPH, S. R. 1992. Observations and remarks on the morphology and taxonomy of the diatom genera Gyrosigma Hassal and Pleurosigma W.Smith. III.Gyrosigma sterrenburgii sp.nov., and Pleurosigma amara sp.nov. – Diatom Research, 7(2): 345–366. STIDOLPH, S. R. 1994. Observations and remarks on the morphology and taxonomy of the diatom genera Gyrosigma Hassal and Pleurosigma W. Smith. IV. Gyrosigma fogedii sp. nov., and some diatoms similar to G. fasciiola (Ehrenb.) Griffith & Henfrey. – Diatom Research, 9(1): 213–224. STRELNIKOVA, N. I. 1992. Paleogenovye Diatomovye Vodorosli. – St.Petersburg University Press, St.Petersburg. 309 p. (in Russian) WEISSE, J. F. 1854: Mikroskopische Analyse eines organischen Polirschiefers aus dem Gouvernement Simbirsk – Melanges bioigiques. Bulletin de l”Academie Imperiale des Sciences de St.Petersburg, 2: 237–250. WITT O. N. 1886. Uber den Polierschifer von Archangelsk-Kuroedovo im Gouv. Simbirsk. – Verhandlungen der Russisch-kaiserlichen mineralogischen Geselschaft zu St.Petersburg. Serie II, 2: 137–177.

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Species of the genus Terpsinoë Ehrenberg (Bacillariophyta) from the Miocene of Middle Russia, Moldova, and Ukraine Tatyana F. Kozyrenko Department of Botany, Biological Faculty, St. Petersburg State University, Universitetskaya Emb. 7/9, St. Petersburg 199034, Russia, e-mail: [email protected] ABSTRACT The frustular morphology of three Terpsinoë species from the Miocene deposits of Russia, Ukraine, and Moldova was studied with SEM. T. americana and T. solida both have poroid areolae, while T. intermedia has pseudolocular areolae. Heterovalvar frustules of T. intermedia (one valve with pseudolocular areolae and another with poroid areolae) have been observed in the past. The pseudolocular structure is might be absent in the earlier stages of cell cycle, in initial or post-initial cells. In this study such heterovalvy was not observed. The welldeveloped pseudolocular structure of the valve should be considered as a reason to maintain the species rank of T. intermedia as opposed to downgrading this taxon to a variety of T. musica, a species that does not have such a well-developed pseudolocular valve structure. Key words: Terpsinoë, pseudosepta, poroid areola, pseudolocula, Neogene, Russia INTRODUCTION The genus Terpsinoë and its type species T. misuca have been described by Ehrenberg (1843) from samples collected in Mexico at the seashore of Atotonilco El Grande and from marine macroalgae and sediments at the shore of Vera Cruz. Van Landingam (1978) listed seven species within this genus. Round et al. (1990) described this genus as an inhabitant of brackish and fresh waters. Luttenton et al. (1986) stated that Terpsinoë has a wide ecological range and lives in both marine and fresh waters.

43

Tatyana F. Kozyrenko

This paper describes the frustular morphology of three Terpsinoë species from the Miocene deposits from Russia, Ukraine, and Moldova. These species are T. americana (Bailey) Ralfs in Pritchard, T. intermedia Grunow, and T. solida Kozyrenko. MATERIALS AND METHODS The following Miocene deposits were studied (stratigraphic descriptions follow Iosifova 1977): 1. Russia, Oka-Don Plain a. Borehole 10 (Lamka Formation, Middle Miocene - Konkian stage) and borehole 92 (Gorelka Formation, Late Miocene - Sarmatian stage) near station Selezni, b. Boreholes 11 and 961 (Gorelka Formation, Late Miocene - Sarmatian stage) near town Komsomolets, c. Outcrop 15 near town Staroe Grayznoe (Gorelka Formation, Late Miocene -Sarmatian stage). 2. Russia, Volga-Khoper interfluvial a. Outcrop near town Gurovo (Ilovlinsko-Gurovskye layers, Middle MioceneKonkian stage). 3. Moldova a. Outcrop near town Naslavcha (Late Miocene - Early Sarmatian stage). 4. Ukraine, Lviv District a. Borehole near town Polyana (Late Miocene - Tortonian stage). Terminology follows Ross et al. (1979).

RESULTS Studied samples contained rich brackish-marine diatom flora where representatives of the genera Hyalodiscus Ehrenberg, Plagiogramma Grewille, Melosira Agardh, and Rhabdonema Kützing were especially abundant. These diatoms apparently inhabited a large shallow brackish lagoon. In Russian materials T. americana was most abundant among Terpsinoë species, in some samples T. solida was moderately abundant, and T. intermedia was rare. In materials from Moldova T. intermedia was found occasionally, and in Ukrainian materials T. solida was observed. Terpsinoë americana (Bailey) Ralfs in Pritchard (1861, p. 859). (Figs 1–27) The frustule is rectangular with rounded corners in girdle view, 20-24 µm high (Fig. 3). Valves are linear and linear-elliptical, triundulate, 35-90 µm long, 20-38 µm wide, with drawn-out ends and two pseudoseptae. Some valves have rounded-elliptical shape, 50-66 µm long, 40-50 µm wide. Pseudoseptae are straight, drop-shaped at free ends (Figs 1, 5, 11). Areolae are poroid, arranged 44

Species of the genus Terpsinoë Ehrenberg (Bacillariophyta) from the Miocene of Middle Russia

5

1 6

2 7 3

4

8 Figs 1-8. Terpsinoë americana, SEM. Fig. 1. External valve view. Scale bar = 10 µm. Fig. 2. external valve surface with granules. Scale bar = 1 µm. Fig. 3. Frustule in girdle view. Scale bar = 10 µm. Fig. 4. Detail of valve mantle, external view. Scale bar = 5 µm. Figs 5-8. Internal valve views. Fig. 5. Whole valve. Scale bar = 10 µm. Figs 6, 7. Central part of the valve. Fig. 6. Scale bar = 10 µm. Fig. 7. Scale bar = 5 µm. Fig. 8. View of a broken valve showing poroid areolae. Scale bar = 1.5 µm.

45


10

9 11

12

13

15

14

16

Figs 9-16. Terpsinoë americana, SEM. Fig. 9. Valve mantle and fragments of the girdle. Scale bar = 20 µm. Figs 9-16. Scale bars = 10 µm. Figs 9-12. External valve surface. Fig. 10. Valve mantle. Fig. 11. Whole frustule. Fig. 12. Apical view of the frustule. Figs 13, 14. Internal valve views. Fig. 13. Whole valve. Fig. 14. Valve with additional apically oriented pseudoseptae. Figs 15, 16. External views of the valve face.

46


17

19

18

20

24

21

22

25

27

23 26

Figs 17-27. Terpsinoë americana. Scale bars = 10 µm. Figs 17-23. LM. Valves of different size and shape. Figs 24-27. SEM. Fig. 24. Internal valve view. Fig. 25. External view of whole frustule. Fig. 26. Detail of the girdle. Fig. 27. External view of the apical part of frustule.

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in radial rows originating from the small round central area, which sometimes has irregular areolae (Figs 15, 16, 23). There are 11-15 rows of areolae in 10 µm, 13-16 areolae in 10 µm in a row. Pseudocelli are large, situated at the valve ends; they are usually bordered by narrow hyaline stripes. Pseudocelli have apically arranged rows of small pores, 18-20 rows in 10 µm, and 20-24 pores in 10 µm in a row (Figs 12, 15). The valve margin bears a row of irregularly spaced elongated granules, 13-20 in 10 µm (Figs 1, 2). Poroid areolae on the valve mantle are arranged in vertical rows, 14 rows in 10 µm (Fig. 4). The lower portion of the mantle is hyaline. No rimoportulae were found. The girdle has several copulae. T. americana was found in Russian materials, where it was usually quite abundant, but not in materials from Ukraine or Moldova. In studied samples this species was extremely variable in shape and size, but not in frustule structure. Besides frustules of typical size and shape, some rounded valves with one pseudoseptum circling middle part of the valve (Figs 19, 20) and valves with additional apically oriented pseudoseptae (Figs 14, 18) were found. Terpsinoë solida Kozyrenko in Kozyrenko et al. (1977, p. 130, pl. 30, figs 6 a, b). (Figs 28-43) The frustule is thick-walled, rectangular with rounded corners in girdle view, 24-36 µm high (Fig. 28). Valves are linear-elliptical, rarely elliptical, with rounded ends, 36-67 µm long, 18-20 µm wide. Valves have two robust straight or slightly bent pseudoseptae, thickened at their free ends (Fig. 28). Areolae are poroid with sunken volae which are more conspicuous from the internal surface of the valve (Figs 38, 40). Valve face has radial rows of areolae originating from a small round central area (Figs 29-31). At the valve ends rows of areolae become less distinct, there are some irregular pores amongst areolae (Figs 34, 43). In the middle part of the valve there are 6-8 rows of areolae in 10 µm, 6-9 areolae in 10 µm in a row. Pseudocelli are small, 7-9 µm in diameter, they are vaguely distinct at the valve ends and have pores in irregular apically oriented rows at the valve ends, 14-16 rows in 10 µm, 14-15 areolae in 10 µm in a row (Figs 34, 35). Valve mantle is 12-20 µm high with areolae in parallel vertical rows, 68 rows of areolae in 10 µm, 5-7 areolae in 10 µm in a row. Low portion of the valve mantle is hyaline (Figs 30, 41). Copulae have vertical rows of small areolae, 18-20 rows of areolae in 10 µm, 13-14 areolae in 10 µm in a row (Figs 42, 43). On the internal valve surface near pseudoseptum a rimoportula opening was found (Fig. 39). T. solida was sometimes quite abundant in Miocene deposits from the Oka-Don Plain, Russia and from Ukraine. Terpsinoë intermedia Grunow (1884, p. 59). (Figs 44-52). The frustule is thick-walled, rectangular with rounded corners in girdle view, 24-30 µm high (Fig. 44). Valves are linear-elliptical, triundulate, with drawn-out attenuate ends, 75-130 µm long, 20-25 µm wide (Fig. 45). Pseudoseptae are straight, with thickened ends which are curved towards valve center. Usually there are four well developed pseudoseptae per valve, 48


28

32

29

33

34

30

35

31

Figs 28-35. Terpsinoë solida. Figs 28, 29. LM. Scale bars = 10 µm. Fig. 28. Girdle view. Fig. 29. Valve view. Figs 30-35. SEM. External valve views. Figs. 30, 31. Whole valves. Scale bars = 10 µm. Figs 32, 33. Central part of the valve. Fig. 32. Scale bar = 10 µm. Fig. 33. Scale bar = 1 µm. Figs 34, 35. Apical part of the valve with pseudocellus. Small pores between regular-sized areolae. Fig. 34. Scale bar = 1 µm. Fig. 35. Scale bar = 10 µm.

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36

37

38

39 40

41 42

43 Figs 36-43. Terpsinoë solida. Figs 36-40. Internal valve surface. Fig. 36. Whole valve. Scale bar = 20 µm. Fig. 37. Central area. Scale bar = 2 µm. Figs 38-40. Scale bars = 1 µm. Fig. 38. Poroid areolae with sunken volae and two small pores. Fig. 39. Rimoportula. Fig. 40. Poroid areolae. Figs 41-43. External valve surface. Fig. 41. Valve mantle with vertical rows of areolae and hyaline edge. Scale bar = 10 µm. Figs. 42, 43. Valve mantle with remains of the valvocopula. Scale bars = 5 µm.

50


48

44 45

49

46

50

51 47 52

Figs 44-52. Terpsinoë intermedia. Figs 44, 45, 48. LM. Scale bar = 10 µm. Fig. 44. Whole frustule in the girdle view. Fig. 45. Whole frustule in valve view. Fig. 48. Detail of the valve in girdle view showing pseudlocular structure of the surface. Figs 46, 47, 49-52. SEM. Figs 46, 47. Scale bars = 10 µm. Fig. 46. External view of the apical part of the valve showing pseudocellus and pseudoloculi. Figs 49, 50. External valve views showing pseudoloculi. Scale bars = 10 µm. Figs 51, 52. Internal valve views. Fig. 51. Detail of internal valve surface. Scale bar = 1 µm. Fig. 52. Rimoportula. Scale bar = 10 µm.

51


sometimes there four additional reduced pseudoseptae. Valve face and mantle bear a network of irregularly spaced pseudoloculi of variable size, 2.5-3, sometimes up to 5 in 10 µm (Figs 44-50). The bottoms of pseudoloculi are perforated by areolae (Fig. 49), areolae openings are clearly seen on the internal valve surface (Fig. 51); there are 10-15 areolae in 10 µm. Central area is absent. Pseudocelli situated at the valve ends are distinct, with large pores in parallel rows, 24-26 pores in 10 µm (Fig. 46). Valve mantle has a hyaline edge (Fig. 49). Copulae were not observed. An internal rimoportula opening was observed in the middle portion of the valve, slightly off the valve center (Fig. 52). T. intermedia was sometimes observed in studied materials from Russia and Moldova. DISCUSSION The pseudoseptae are the most stable character of the genus Terpsinoë, while the structure of the valve surface varies among species. The species studied here have either simple structure of poroid areolae (T. americana, T. solida) or have pseudoloculi in the basal layer (T. intermedia). The term “pseudoloculus” used here following Ross et al. (1979) corresponds to the term “poroid canals in depressions formed by anastomosing costae” of Nikolaev and Harwood (2002, p. 108). These two types of valve structure were illustrated by Round et al. (1990). Their figures a, i, and h (pp. 256, 257) show poroid areolae, while figures b, c, d, and e show pseudoloculi, but pseudoloculi were not mentioned in the genus description. Ehrlich (1995, p. 39, pl. 5, figs 6-9) describes the valve of T. musica as ”radially striate-punctate in its central part, apically striate-punctate towards the extremities, but her fig. 8 clearly illustrates pseudoloculi. Old authors, such as Ralfs (1861) regarded the structure of valve surface in Terpsinoë as an important diagnostic character, and Grunow (1884) used the presence of reticulate (“unregelmässigen Maschen”) structure to distinguish his new species T. intermedia. The taxonomic rank of Terpsinoë intermedia has been the subject of a discussion for a long time. Grunow (1884, p. 59) did not leave a formal text description of this species, but in the note to Biddulphia flos (Ehrenberg) Grunow discussed the genus Terpsinoë and noted that unlike T. musica and T. americana, T. intermedia has a reticulate valve surface structure. Pantocsek (1903), Peragallo and Peragallo (1987-1908), Boyer (1926), and Schmidt (1895) recognized T. intermedia as separate species, but later Hustedt (1930) downgraded it to a variety of T. musica. Hustedt argued that some cells have one valve with reticulate structure, but another valve does not have such a structure. This heterovalvy was documented by Schmidt (1895, Plate 199, Figs 1-8) who illustrated various stages of life cycle of T. intermedia. His fig. 4 shows a regular cell with reticulate structure of both valves and his fig. 1 shows an initial cell with one reticulate valve, and another with a simple structure of poroid areolae. Schmidt (1895) also wrote that T. musica has thin distinct rows of areolae, while T. intermedia in most cases has reticulate structure, but mentioned that Cleve did not distinguish these two species. In studied populations of T. intermedia from Miocene deposits from Russia and Moldova heterovalvar frustules were absent. It appears that such heterovalvar frustules 52


have been only observed in initial or post-initial cells, and therefore, the well-developed reticulate or pseudolocular structure is a character that allows distinguishing T. intermedia from T. musica. The pseudocellus clearly separated in T. intermedia is also a distinctive character of this species. ACKNOWLEDGEMENTS This work was supported by the grant 03-04-48711 of the Russian Foundation for Fundamental Research. Many thanks to N. Ognjanova for the invitation to participate in the volume honoring Prof. D. Temniskova-Topalova. I also thank A. Ulanova for several SEM photos; and N. I. Strelnikova and M. G. Potapova for their help in preparing this manuscript. REFERENCES BOYER, C. S. 1926. Synopsis of North American Diatomaceae. Part I. Proceedings of the Academy of Natural Sciences of Philadelphia, 78: 228 p. EHRENBERG, C. G. 1843. Verbreitung und Einfluß des mikroskopischen Lebens in Süd- und Nord-Amerika. Abhandl. der Königl. Academie der Wissenschaften zu Berlin: 291-446. EHRLICH, A. 1995. Atlas of the inland-water diatom flora of Israel. Flora Palestina. Jerusalem, The Israel Academy of Sciences and Humanities. GRUNOW, A. 1884. Die Diatomeen von Franz Josefs-Land. Denkschriften der matematisch-naturwissenschaftlichen Classe der kaiserlichen Akademie der Wissenschften, 48: 53-112. HUSTEDT, F. 1930. Die Kieselalgen Deutschlands, Österreichs und der Schweiz. Mit Berücksichtigung der übrigen Länder Europas sowie der angrenzenden Meeresgebiete. in: Rabenhorst’s Kryptogamen-Flora von Deutschland, Österreich und der Schweiz. Leipzig, Akadem. Verlag., 7(1): 1-920. IOSIFOVA, J. I. 1977. Geology of Miocene deposits of the Oka-Don Plain. Pages 6-53 in: Miocene of the Oka-Don Plain. Moscow, Nedra (in Russian). KOZYRENKO, T. F., A.P. JOUSÉ and KOSLOVA O. G. 1977. Diatom flora of Miocene deposits of the Oka-Don Plain. Pages 113-131 in: Miocene of the Oka-Don Plain. Moscow, Nedra. LUTTENTON, M. R., L. PFIESTER and P. TIMPANO 1986. Morphology and growth habit of Terpsino¸ musica Ehr. (Bacillariophyceae). – Castanea, 51: 175-182. NIKOLAEV, V. A. and D.M. HARWOOD 2002. Morphology, taxonomy and system classification of centric diatoms. St.Petersburg, Nauka (in Russian). PANTOCSEK, J. 1903. Beitrage zur Kentnis der fossilen Bacillarien Ungarns. I Teil. Marine Bacillarien. 2. verbesserte Auflage. Verlag von W.Junk, Berlin. PERAGALLO, M. H. and M. PERAGALLO 1897-1908. Diatomées marines de France et des districts maritimes voisins. Tempère, Grez-sur-Loing. 53


RALFS, J. 1861. Sub-group Diatomeae or Diatomaceae. Pages 756-940 in Pritchard A. History of Infusoria including the Desmidiaceae and Diatomaceae, British and Foreign. Wittaker and Co., London. ROSS, R., E.J. COX , N.I. KARAYEVA , D.G. MANN, T.B.B. PADDOCK, R. SIMONSEN and P.A. SIMS 1979. An amended terminology for the siliceous components of the diatom cell. Nova Hedwigia, Beih. 64: 513-533. ROUND, F. E., R.M. CRAWFORD and D.G. MANN 1990. The diatoms. Biology and morphology of the genera. Cambridge University Press. Cambridge. SCHMIDT, A. 1895. Atlas der Diatomaceenkunde. R. Reisland, Leipzig. VAN LANDINGHAM, S. L. 1978. Catalogue of the fossil and recent genera and species of diatoms and their synonyms. Part VII. Rhoicosphenia through Zygoceros. J. Cramer, Vaduz.

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Comparison of seven species of Navicula sensu stricto Six species described as new to science from Miocene lacustrine deposits in Bulgaria and Romania Horst Lange-Bertalot1 and Ditmar Metzeltin2 Botanical Institute, J.-W. Goethe-University, Senckenberganlage 31–33, 60054 Frankfurt am Main, Germany 2 Am Stegskreuz 3b, D-65719 Hofheim, Germany, E-mail: [email protected] 1

ABSTRACT Navicula bulgariarum, N. nadjae, N. peripontica, N. serdicensis, N. temniskovae are described as fossil freshwater species, new to science, from material of a core 1.50-50.00 m depth, located at 18 km North of Sofia, Bulgaria. N. superhasta is described from comparable Neogene material found near Köpecz in Transylvania, Romania. Both materials contain likewise Navicula hasta Pantocsek. The common but not quite clear species concept of this prominent taxon is critically discussed. Name of the new taxa is known to occur extant. Key words: Bacillariophyta, Navicula, new species, Sofia Neogene Basin, Bulgaria INTRODUCTION Late Alpine tectonic events in the central parts of the Balkan Peninsula differ from coeval tectonic events in the other parts of the Alpine orogens in Eurasia (Zagorchev 1996). The geodynamics of the realm had been changed by an intense compressional phase in earliest Neogene times. The long lasting peneplanation led to the formation of a large swells, separating graben complexes and rift valleys, surrounded and insulated by marine basins (Pannonian, Peri-Carpathian, Euxinian, Aegean, Adriatic). Neogene Sofia Basin is situated in South Bulgaria and its deposits fill the Sofia graben. Four lithostratigraphic units are described in the neogene from the basin: 1. Varriagated 55

Horst Lange-Bertalot and Ditmar Metzeltin

terrigenous formations (probably Meotian age); 2. Gniljane Formation (Pontian inf.); 3. Novi-Iskar Formation (Pontian, partly Dacian inf.) and 4. Lozenec Formation (DacianRomanian age, Kamenov and Kojumdgieva 1983, Angelova and Yaneva 1998). With the exception for Navicula superhasta, from Transylvania, all other new taxa and also the taxon here identified as Navicula hasta Pantocsek 1897 originate from the locality Goljanovci near Sofia. Diatomologically both materials have been investigated by several authors in the past. Generally they contain taxa that have survived from the Pliocene, Pleistocene and Holocene periods until recently (Temniskova-Topalova and Ognjanova-Rumenova 1997), whereas various other taxa are known exclusively as fossil records from the Neogene period, and from several locations in SE– or E– Europe or eastern central Europe. For a major part they can be found documented and depicted in form of line drawings or photographs, particularly by Pantocsek (1892-1905) and more recently e.g. by Ognjanova-Rumenova (1991) or Temniskova-Topalova and Ognjanova-Rumenova (1997). However, a critical study and comparison show that more than a few populations can be proved to be not identifiable as established taxa. Concerning Navicula s. str. Several obviously independent populations have been overlooked, neglected or lumped together under few prominent names like Navicula hasta or Navicula lacusbaicali. Symptomatically older species concepts turn out to appear much too broad. MATERIAL AND METHODS The sediments studied were collected from borehole C-1, village of Goljanovci, 18 km North of Sofia, 23o13’48’’ East and 42o51’00’’ North. The borehole penetrated through Novi Iskar Formation and is represented by irregular alternation of diatom clays (0-50.00 m), lignite and lignite clays (50.0-53.0 m), gray clays (53.0-68.0 m), lignite clays (68.0-76.70 m) and gray limestones (76.7-89.30 m). Diatom bearing sediments from borehole C-1 (1.50-48.80 m) were investigated. The other investigated material (containing Navicula superhasta) originates from Köpecz Neogene deposits, southern Carpathians, Transylvania, Romania. Slides containing that material are housed in the Hustedt Collection, Bremerhaven and were probably mounted by Pantocsek from the material collected during the last decades of the 19th century. The samples for the current study were prepared following standard laboratory procedures (Ognjanova-Rumenova 1991). RESULTS AND DISCUSSION Navicula hasta Pantocsek 1892, Icon. Beitrg. III, pl. 5, fig. 74 (pl. 1: Figs 1-3, pl. 2: Figs 1-4) Description: Pantocsek 1905, p. 69 To exclude from the diagnosis is: “Navicula hasta” Pantocsek 1892, Icon Beitrg. III, pl. 14, fig. 213. Pantocsek’s diagnosis, translated from Latin, reads:

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Comparison of seven species of Navicula sensu stricto

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1 Plate 1, Figs 1-3 Navicula hasta Pantocsek 1892. Scale bar 10 µm.

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4 3 2 1 10 µm Plate 2, Figs 1-4 Navicula hasta Pantocsek 1892. Scale bar 10 µm.

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Valves elongated lanceolate, 63-76 µm long, 14-18 µm broad, with produced obtuse poles. Raphe straight, encircled by a rather dilated axial area. Striae coarse, radiate, lineolated, 8-10 in 10 µm, in the centre longer and shorter. In Tertiary freshwater deposits of Bibarczfalva, Bodos and Köpecz in Transylvania. Icon: Beitrg. III, tab. 5, fig. 74, tab. 14, fig. 213.

From the two icons only fig. 5: 75 matches the diagnosis, whereas the specimen in fig. 14: 213 is approximately 108 µm long and 25 µm broad. This exceeds the maximum data considerably. The two depicted specimens originate from Köpecz. Recent observations on the type material displayed several “size classes” of Navicula hasta – like specimens (cf. Lange-Bertalot 2001, plates 58-60). Whilst the specimens in plates 58 and 59 – although not all matching the diagnosis – may be identified as N. hasta, the population represented in plate 60 can hardly belong to this taxon in a strict sense (see under N. superhasta). In the deposits of the Sofia region N. superhasta was not observed. However – like in Köpecz – occurrence of Navicula hasta sensu lato is represented by a smaller and a larger sized deme. The first one with specimens about 80 µm long and 14-15 µm broad, striae 77.5/10 µm (plate 2: Figs 2-4); the second one (plate 1: Figs1-3, plate 2: Fig.1) 95-120 µm long, 17-20 µm broad, 6-6.5 striae/10 µm proximally and 8-9/10 µm distally. Other features e.g. position of the striae, lineola density, axial and central area, raphe with position of the central pores are coincident. Therefore a broader concept of N. hasta than given in Pantocsek’s diagnosis seems to be more appropriate. However, a “catch–all” concept encom-passing all N. hasta–like populations may hardly reflect the taxonomic reality. Navicula superhasta nov. spec. (pl. 3: Figs 1-3) Navicula hasta sensu Pantocsek 1892, pl. 14, fig. 213 (non pl. 5, fig. 74 nec description 1905, p. 69) Navicula hasta sensu Hustedt in A. Schmidt Atlas partim (figs 395: 10-12) Navicula (?nov.) spec. Lange-Bertalot 2001, pl. 60: 1-4 Diagnosis differens versus Navicula hasta Pantocsek et Navicula temniskovae nov. spec. Valvae rhombico-lanceolatae ad apices plus minusve leniter protractae. Longitudo 105-160 µm, latitude 21-25 µm (id est conspicue plus quam N. hasta quoad diagnosem). Raphe lateralis poris centralibus conspicue sitis (quam N. temniskovae). Area axialis fere lata, sterno lato raphis distincte conturato (ita differ tab alteris speciebus). Area centralis aliquid indistincta vel variabiliter formata quia striae hic irregulariter abbreviatae. Striae transapicales radiantes fortiter omnino usque ad apices, 6 proximaliter, 8 distaliter (nec 8-10 in mediis partibus quam in N. hasta). Lineolae 23-25 (nec 16-18) in 10 µm (ita differt N. temniskovae).

Typus: Praep. 400/116 in Coll. Hustedt, Bremerhaven Locus typicus: Köpecz Neogene Deposits Etymology: “super–hasta” means dimensions larger and structures coarser than in N. hasta. 59

10 µm

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Plate 3, Figs 1-3 Navicula superhasta nov. spec. Scale bar 10 µm.

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Differential diagnosis compared to Navicula hasta Pantocsek and Navicula temniskovae nov. spec. Valves rhombic-lanceolate tapering to the more or less distinctly protracted ends. Length 105-160 µm, breadth 21-25 µm (this is conspicuously more than in N. hasta where less than 100 and 14-20 respectively are indicated by Pantocsek). Raphe lateral, lying in a comparatively broad, strongly contoured sternum (different from both taxa in comparison). Central pores comparatively close, deflected to the secondary side. Axial area rather broad, central area somewhat indistinct or with variable shape, confined by some irregularly shortened striae. Other striae strongly radiate throughout (not subparallel at the ends) about 6 in 10 µm and 8 only at the ends (not 8-10 in all proximal parts). Lineolae 23-25 in 10 µm (not 16-18 as in N. temniskovae). Remarks: The complex of characters is much closer related to N. hasta than to N. temniskovae due to the arrangement of the striae, lineolae and the narrower standing central pores of the raphe. Nevertheless, the numerical dimensions, given in the protologue of N. hasta, differ so significantly that assumption of heterospecificity appears more likely than conspecificity in this case. Infraspecific offspring is not to exclude, but remains hypothetical just like supposing monophyletic sister species. Even the assumed conspecificity of various morphodemes in N. hasta with closer related numerical dimensions of cell sizes and striae densities is not out of question yet (see morphodeme 1 and 2 from Sofia and morphodeme with broader, more rhombic valve outlines from Dubravica shown in Lange-Bertalot 2001, figs 59: 1-7). Navicula serdicensis nov. spec. (pl. 4: Figs 1-6) Diagnosis differens specimem fossilem Navicula radians Héribaud 1903. Valvae simpliciter lanceolatae apicibus non protractis aliquid minus obtuse rotundatis. Longitudo 145-180 µm (nec 120-150), latitudo 23-30 µm (nec 20). Fissurae raphis distincte laterales sitae inter se poris centralibus comparate parvis ad latus secundum deflexis nec conspicue distanter nec dense sitis inter se. Area axialis fere lata irregulariter dilatata ad mediam versus (parum sed non conspicue latius formata unilateraliter). Area centralis variabilis ad instar (tamen numquam circularis) quia striae mediae irregulariter alternantes longae curtaeque (ita ut in N. radians). Striae transapicales radiantes omnino usque ad apices, constanter 6.5-7 in 10 µm (ut N. radians). Lineolae 23-26 in 10 µm (incognitae in N. radians).

Typus: Praep. Core 1/20, 30.0 m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia. Locus typicus: 18 km North of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; Borhole is 23o13’48’’ East and 42o51’00’’ North. Etymology: Serdica is the ancient name of the recent city of Sofia. Differential diagnosis compared to the fossil taxon Navicula radians Héribaud 1903. Valves simply lanceolate tapering to moderately acute (rather than somewhat more obtusely) rounded ends. Length 145-180 µm (not 120-150). Breadth 23-30 µm (not 20 as given in Héribaud’s protologue). Raphe fissures distinctly lateral with comparatively small 61


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Plate 4, Figs 1-6 Navicula serdicensis nov. spec. Scale bar 10 µm.

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central pores deflected to the secondary side, neither remarkably far nor close standing. Axial area moderately broad, gradually expanded towards the centre (not con-speciously broader unilaterally). Central area variable in shape (but never markedly circular) by irregularly alternately longer and shorter striae. Striae radiate throughout, very consistently 6.5-7 in 10 µm (as in N. radians). Lineolae 23-26 in 10 µm (unknown density in N. radians). Remarks: N. radians Héribaud 1903 non N. radians (Østrup 1899) Cleve-Euler 1922 is an almost forgotten taxon from the Neogene deposits of Ceyssac, Auvergne, France. Later on it was synonymized with Navicula radiosa var. acuta (W. Smith) Grunow 1860 by other authors. However, that fossil taxon appears by no means closely related to N. radiosa which is characterized by clearly convergent striae towards the ends instead of distinctly radiate striae throughout. Héribaud compares his taxon with N. vulpina Kützing which, however, resembles very little the two fossil taxa in question here. The other Navicula species from Neogene deposits are distinguished by other distinct complexes of characters. Comparable is also Navicula ajajensis Skabitschevsky 1936, recent from Lake Bajkal with a length of 95108 µm, breadth 14.4-16.2 µm, and 7.8-8.8 striae in 10 µm, only one of the central striae is shortened on either side. Navicula temniskovae nov. spec. (pl. 5: Figs 1-3) Diagnosis differens versus Navicula hasta Pantocsek. Valvae plus minusve rhombico-lanceolatae cuneatim attenuatae ad apices nec acute nec obtuse rotundatis omnino non protractis. Longitudo 70-140 µm, latitudo 22-31 µm (non 60-90 nec 1519 µm). Raphe lateralis poris centralibus distincte dilatatis, conspicue distanter sitis inter se et deflexis ad latus secundum. Area axialis modice lata ad mediam dilatata. Area centralis fere lata, variabilis ad instar plerumque plus minusve rhombice expansa. Striae transapicales radiantes in mediis partibus hic 5-6 in 10 µm tam usque ad apices paulatim subparallelae circiter 7 in 10 µm (nullo loco 8-10). Circum aream centralem complures striae abbreviatae irregulariter intermixtae (non una singula stria vel nulla). Item striae non conspicue distanter inter se. Lineolae 16-18 (nec circiter 26) in 10 µm. Alterae taxa similissimae adhuc non descriptae sed vide sub Navicula superhasta. Navicula peregrina sensu auct. nonnull satis differt.

Typus: Praep. Core 1/12, 18.0 m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia. Locus typicus: 18 km North of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; 23o13’48’’ East and 42o51’00’’ North. This species is dedicated to our colleague Prof. Dr. Dobrina Temniskova on the occasion of her 70 th birthday. Differential diagnosis compared to Navicula hasta Pantocsek. Valves more or less rhombic-lanceolate tapering to the non-protracted cuneately rounded ends. Length 70-140 µm, breadth 22-31 µm (not 60-90 and 15-19 µm respectively). 63


3 2 10 µm

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Plate 5, Figs 1-3 Navicula temniskovae nov. spec. Scale bar 10 µm.

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Raphe fissures narrow-lateral, central pores expanded conspicuously wide-standing and deflected to the secondary side. Axial area moderately broad expanding to the centre. Central area variable in outlines, mostly more or less rhombically expanded. Striae radiate proximally, 5-6/10 µm becoming less radiate distally and finally subparallel, c. 7 in 10 µm (nowhere 8-10). Around the central area several striae shortened irregularly (not only a single pair and not distinctly wider spaced here). Lineolae 16-18 (not c. 26) in 10 µm. Other large-sized Navicula taxa will be hardly confused with the new taxon because the complexes of characters like e.g. Navicula peregrina (see also under N. superhasta) differ significantly. Navicula nadjae nov. spec. (pl. 6: Figs 1-6, ?7) Diagnosis differens versus Navicula hasta Pantocsek. Valvae anguste-lanceolatae ad apices fere acute rotundatos paulatim attenuatae apices non vel vix protracti. Longitudo (?70)115-165 µm, latitudo 12-18 µm (nec rhombico-lanceolatae). Raphe lateralis poris centralibus potius aliquid distanter quam dense sitis inter se deflexis ad latus secundum. Area axialis anguste-lanceolata. Area centralis aliquid indistincta et variabiliter formata. Striae transapicales fortiter radiantes usque sub apices sed una singula stria utrimque parallelae vel parum convergentes, 8-8.5 in 10 µm in mediis partibus, 9-10 sub polos. Lineolae circiter 24-25 in 10 µm. Similissima Navicula rakowskae Lange-Bertalot differt lineolis densius sitis inter se enim 30-31 in 10 µm etiam mediis striis distantius sitis inter se.

Typus: Praep. Core 1/19, 28.5m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia. Locus typicus: 18 km North of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; Borehole is 23o13’48’’ East and 42o51’00’’ North. Etymology: The new species is dedicated to Dr. Nadja Ognjanova-Rumenova, Sofia. Differential diagnosis compared to Navicula hasta Pantocsek. Valves narrowly lanceolate tapering to fairly acutely rounded apices that are not or barely protracted. Length (?70)115-165 µm, breadth 12-18 µm (not rhombic-lanceolate). Raphe fissures lateral, central pores moderately distant rather than close-standing, deflected to the secondary side. Axial area narrow-lanceolate. Central area somewhat indistinct or variable in shape. Striae strongly radiate up to the ends, only the apical pair of striae parallel to slightly convergent, 8-8.5/10 µm, except for the most distal ones becoming 9-10/10 µm. Lineolae 24-25/10 µm. Similar large-celled taxa e.g. the fossil Navicula radiosa var. dubravicensis Grunow or Navicula costei Héribaud differ by the numerous distinctly convergent stria–pairs distally and the broader valves. Most similar to the new taxon is the recent Navicula rakowskae Lange-Bertalot, it is distinguished by significantly more densely spaced lineolae, 30-31/10 µm. Moreover, in N. rakowskae the central striae are much more spaced than the others. Navicula lucida Pantocsek (syn. N. lucifica Pantocsek) 1882 fig. 18: 264, fossil from Bodos and Köpecz (non Navicula lucida O’Meara 1876) is longer, broader and with denser striae (15-16 striae/10 µm). 65


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Plate 6, Figs 1-6, ? 7 Navicula nadjae nov. spec. Scale bar 10 µm.

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Navicula peripontica nov. spec. (pl. 7: Figs 1-3) Diagnosis difference versus Navicula lacusbaicali Skvortzow & Meyer 1928 et Navicula perturbata Jurilj 1954 Species subsectionis Navisantiqua Lange-Bertalot 2001. Valvae simpliciter lanceolatae apicibus non protractis sed cuneatim attenuatis denique rotundatis. Longitudo 80-90 µm, latitudo 18.5-20 µm (nec 24-27). Fissurae raphis laterales poris centralibus (parvius dilatatis quam in N. perturbata) ad latus secundum deflexis. Area axialis fere angusta, dilatata ad mediam versus. Area centralis fere late rectangulata striis 5-6 abbreviatis formata. Striae transapicales, 7.5 in 10 µm, radiantes sed 5-7 striae sub apices utrimque distinctissime convergentes parum densius sitae inter se enim 8-8.5 in 10 µm (numquam radiantes omnino usque ad apices ut in N. lacusbaicali et N. perturbata). Lineolae circiter 26 in 10 µm (non circiter 40 nec minus quam 26).

Typus: Praep. Core 1/20, 30.0 m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia. Locus typicus: 18 km North of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; Borhole is 23o13’48’’ East and 42o51’00’’ North. Etymology: “peri–pontica” means the area around the Black Sea in a wide sense. Differential diagnosis compared to Navicula lacusbaicali Skvortzow & Meyer 1928 and to Navicula perturbata Jurilj 1954. Valves lanceolate with simply wedge-shaped ends (neither protracted nor markedly obtusely or acutely rounded). Length 80-90 µm, breadth 18.5-20 µm. Raphe fissures lateral, central pores (smaller than in N. perturbata) deflected to the secondary side. Axial area rather narrow, expanded only in their proximal parts. Central area broadly rectangular or tie-bowshaped, confined by 5-6 shortened striae on either side. Other striae, 7.5 in 10 µm, proximally radiate, but the distal 5-7 pairs of striae becoming progressively more strongly convergent (not radiate throughout). The subpolar striae are only slightly more densely spaced, c. 8 in 10 µm (not up to 10 as in N. perturbata). Lineolae around. 26 in 10 µm (not about 40 as in N. perturbata or less than 25 as in all varieties of N. lacusbaicali). Ecology: See under N. bulgariarum. Remarks: This species is one among fairly few known taxa belonging to the subsection Navisantiqua of the section Navicula in Navicula s. str. (see Lange-Bertalot 2001). Navicula haueri Grunow from Dubravica is another fossil species. It possesses also convergent striae distally - in contrast to the recent taxa - and very bluntly rounded ends. Probably more than a single taxon is included in the range of forms identified uniformly as N. haueri by different authors. Thus, two specimens of N. haueri sensu P.T. Cleve in A. Schmidt Atlas, figs 212: 32a, b from Jastraba deposits possess narrower, slightly protracted ends (i.e. more similar to N. peripontica) and a narrower, transversely rectangular central area with 2-3 shortened striae instead of the almost circular one of N. haueri from Dubravica.

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Plate 7, Figs 1-3 Navicula peripontica nov. spec. Scale bar 10 µm.

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Navicula bulgariarum nov. spec. (pl. 8: Figs 1-3) Diagnosis differens versus Navicula peripontica nov. spec. (vide supra). Valvae potius paene rhombico-lanceolatae (quam simpliciter lanceolatae) quia constanter fortius attenuatae ad apices versus. Longitudo 60-70 µm quoad pauca specimina adhuc cognita. Latitudo 15-20 µm. Altera signa generaliter aequalia speciebus ambabus tamen striae transapicales densius sitae inter se enim proximaliter 9 (non 7-8) in 10 µm, sub apices 10-11 (non 8-9) in 10 µm.

Typus: Praep. Core 1/20, 30.0 m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia. Locus typicus: 18 km North of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; Borhole is 23o13’48’’ East and 42o51’00’’ North. Etymology: The species is dedicated to our colleagues (so far ladies are concerned) working at the fossil diatom flora of Bulgaria. Differential diagnosis compared to Navicula peripontica (see above). Valves consistently rhombic-lanceolate (not simply lanceolate) since more strongly attenuated from the middle towards the slightly protracted ends. Poles almost acutely (rather than obtusely) rounded. Length 60-70 µm, obviously representing only a small part of variability in the cell cycle; width 15-20 µm. The peculiar pattern of the other characters generally conforming in both taxa. However, transapical striae spaced more densely, 9 (not 7-8) in 10 µm proximally, becoming 10-11 (not 8-9) near the ends. Ecology: The results of the palaeoecological analyses of the sediment deposition of the Novi Iskar Formation, from which Navicula bulgariarum and N. peripontica were determined, demonstrated that during the Late Middle Miocene period there was an increase of the percentage of the mesohalobous diatoms. In the beginning of the sediment formation the salt content ranged between 0.2-0.3‰, and later went up to 5‰. Therefore the brackish influence during the sediment genesis was established. However most of the species in taxonomical composition are non-marine – including numerous species of Aulacoseira, Diploneis, Surirella, and Campylodiscus. (Ognjanova-Rumenova 1997) ACKNOWLEDGEMENTS We gratefully acknowledge the help rendered to us by Dr. Nadja Ognjanova-Rumenova yielding the fossil material that contained five of the new taxa. Moreover she made translations of data from her Doctoral Thesis (written in Bulgarian language).

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REFERENCES ANGELOVA, D. and M. YANEVA 1998. New lithostratigraphical data about Neogene of Sofia Basin. – Review of the Bulgarian Geological Society, 59(2): 37-40 (in Bulgarian). HÉRIBAUD, J. 1893-1908. Les diatomées d’Auvergne, Libr. des Sci. Nat. Paris. p. 1-233, 6 pl. (1893); premier mémoire, p. 1-79, 2 pl. (1902); deuxième memoire, p. 1-166, 4 pl. (1903); troisième memoire, p. 1-70, 2 pl. (1908). JURILJ, A. 1954. Flora I Vegetacija Dijatomeja Ohridskog Jezera. – Prirodosl. Istraz., 26: 99-110. KAMENOV, B. and E. KOJUMDGIEVA 1983. Stratigraphy of the Neogene in Sofia Basin. Palaeontology, stratigraphy and lithology. – Bulgarian Academy of Sciences, 18: 6985 (in Bulgarian). LANGE-BERTALOT, H. 2001. Diatoms of Europe. Diatoms of the European Inland Waters and Comparable Habitats. (ed. Lange-Bertalot, H.) Vol. 2. Navicula sensu stricto - 10 Genera Separated from Navicula sensu lato - Frustulia. 526 p., 140 pl. – A.R.G. Gantner Verlag K.G. FL 9491 Ruggell. Distributed by Koeltz Scientific Books, Koenigstein. OGNJANOVA-RUMENOVA, N. 1991. Neogene diatoms from sediments of Sofia Valley and their stratigraphic significance. – Ph.D. Thesis, Geological Institute, Bulgarian Academy of Sciences, 330 pp. (in Bulgarian). OGNJANOVA-RUMENOVA, N. 1997. Lake trophic evolution determined by fossil diatoms, Chrysophycean stomatocysts and mollusca. - Annales Geologiques des Pays Helleniques, Athenes, premiere serie, 37, 1996-1997: 97-117. O’MEARA, E. 1876. Report on the Irish diatomaceae.- Proc. R. Irish Acad., second. Ser., 2: 235-425, 9 pl. ØSTRUP, E. 1899. Diatoméerne. in: Hartz, N. and E. Østrup, editors. Danske Diatoméjord-Aflejringer og deres Diatoméer. Danmarks geol. Undersogelse II. 9: 35-81, 2 pl. – Kjoebnhavn. PANTOCSEK, J. 1892. Beiträge zur Kenntnis der fossilen Bacillarien Ungarns. Teil 3. Süßwasser Bacillarien. Anhang: Analysen 15 neuer Depots von Bulgarien, Japan, Mähren, Rußlands und Ungarn, 42 pl. Nagy-Tapolcsany. PANTOCSEK, J. 1905. Beschreibung neuer Bacillarien, welche in der Pars III der “Beiträge zur Kenntnis der fossilen Bacillarien Ungarns” abgebildet wurden. 118 pp. – Pozsony. SCHMIDT, A. et al. 1874-1959. Atlas der Diatomaceen-Kunde. Heft 1-120, Tafeln 1-480 (Tafeln 1-216 A. Schmidt; 213-216 M. Schmidt; 217-240 F. Fricke; 241-244 H. Heiden; 245, 246 O. Müller; 247-256 F. Fricke; 257-264 H. Heiden; 265-268 F. Fricke; 269-472 F. Hustedt). – Aschersleben & Leipzig. SKABITSCHEVSKY, A.P. 1936. Neue und interessante Diatomeen aus dem nördlichen Baikalsee. – Bot. Zhurnal, 21: 705-719, 3 pl. (Russian).

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SKVORTZOW, B.W. and C.I. MEYER 1928. A contribution to the diatoms of Baikal Lake. – Proc. Sungare River Biol. Stat., 1/5: 1-55, 3 pl. TEMNISKOVA-TOPALOVA, D. and N. OGNJANOVA-RUMENOVA 1997. Description, comparison and biastratigraphy of the nonmarine Neogene diatom floras from South Bulgaria. – Geol. Balcan., 27 (1-2): 57-81. ZAGORCHEV, I. 1996. Late Alpine (Palaeogene-Early Miocene) tectonics and neotectonics in the central parts of the Balkan Peninsula. – Z. geol. Wiss., 24, 1/2: 91-112.

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Evolution of the extinct genera belonged to the family Stephanodiscaceae (Bacillariophyta) during the last eight million years in Lake Baikal Galina Khursevich Institute of Geochemistry and Geophysics, National Academy of Sciences of Belarus, Kuprevich street 7, Minsk 220141, Republic of Belarus, email: [email protected] ABSTRACT In the Upper Cenozoic (Upper Miocene – Holocene) sediments of Lake Baikal, recovered by the long continuous drill cores BDP-98 (600 m long), BDP-96-1 (200 m long) and BDP96-2 (100 m long) on the top of the underwater Academic Ridge, revealed a rich and diverse composition of planktonic centric diatoms from the family Stephanodiscaceae. The latter is represented by 60 species and 12 intraspecific taxa belonging to 9 genera. Among them, 6 genera (3 endemic for Lake Baikal and the Transbaikal area), 42 species (41 baikalian endemics) and 10 intraspecific taxa (8 baikalian endemics) are extinct. The extinct genera from the family Stephanodiscaceae (Concentrodiscus, Mesodictyon, Mesodictyopsis, Tertiariopsis, Stephanopsis and Tertiarius) replace each other from bottom to top along the studied sections of the BDP-98 and BDP-96-1 cores that allows to trace the evolution of some morphological characters in time. The pattern of morphological evolution in these genera in the ancient Baikal ecosystem can be described as follows: a) change of the cribrum position in areola; b) change of the stria morphology; c) change of the structure of marginal fultoportulae on the valve mantle; and d) change of the number of rimoportulae. Key words: diatoms, Stephanodiscaceae, extinct genera, evolution, Lake Baikal. INTRODUCTION Lake Baikal (situated in the southeastern part of Siberia, Russian Federation), is the world’s deepest (1637 m), largest (about 600 km long) and oldest (25–30 million years) extant 73

Galina Khursevich

freshwater lake. It is a natural laboratory for studing a wide variety of modern and ancient global change processes. Lake Baikal is very sensitive to orbitally forced climatic changes because it is located at high latitude (52º- 56º N) in the middle of a continent and surrounded by mountain ridges (Baikal Drilling Project, BDP-Members 1997). This lake occupies the deepest portion of the Baikal Rift Zone (BRZ), one of the most active continental rift zones of the world. This makes it possible to use Baikal deposits to study the development of sedimentary basins and the evolution of geographically isolated populations including freshwater diatoms. The high-resolution continuous sedimentary records from drill cores BDP-98 (600 m long), BDP-96-1 (200 m long) and BDP-96-2 (100 m long) spanning the last 8 Ma (BDPMembers 2000, Sapota et al. 2004), 5 Ma and 2.5 Ma (BDP-Members 1997), respectively are unique for the whole Eurasian continent. The profile of diatom abundance, generic and species composition change allowed to distinguished 58 biostratigraphic diatom zones in Lake Baikal deposits over the Late Miocene – Holocene interval from 8 Ma to 0 Ka BP. Among them, 9 diatom zones were indentified in the Upper Miocene sediments, 8 diatom zones in the Pliocene deposits, and 41 diatom zones in the Quaternary sediments (Khursevich et al. 2001a, b, c, 2003c, 2005). Dramatic changes in insolation during the Pleistocene (1.79– 0.01 Ma) produced surprisingly rapid diatom speciations and extinctions in the ancient basin. The orbitally-tuned age model allowed diatom assemblages to be compared with individual marine isotopic stages and substages, especially of the Brunhes chron (Khursevich et al. 2001a,c). Diatom succession studied in the Brunhes section of the BDP-96-2 core showed that most of the interglacial periods are characterized by explosive speciation and appearances of new pelagic planktonic species, especially within the Stephanodiscus genus many taxa of which are limited to narrow age ranges. In contrast, most of the glacial periods are distinguished by spectacular extinctions (Khursevich et al. 2001c). At least 30 new centric species and intraspecific taxa appeared in the Pleistocene record of Lake Baikal. Among them, 26 members of centric diatoms belong to extinct endemics (see Table 1). The evolution of planktonic centric diatoms within the ancient Baikal ecosystem during the Late Miocene and Pliocene yielded both new species and new genera. Main trends in the evolution of extinct genera from the family Stephanodiscaceae in the Lake Baikal for the last eight million years are described in this paper. MATERIALS AND METHODS Numerous samples collected from the long continuous drill cores BDP-96-1, BDP-96-2 and BDP-98 were analyzed for diatoms. Two parallel cores BDP-96-1 and BDP-96-2 were taken on the top of the underwater Academician Ridge of Lake Baikal at 53°412483 N and 108°212 063 E in water depth of 321 m (BDP-Members 1997). The drilling of the BDP-98 core was performed on this ridge at 53°442483 N and 108°242343 E in water depth of 333 m (BDP-Members 2000, 2001). The total core recovery was about 95%. Sediments consist of alternating biogenic diatomaceous ooze and terrigenous clay intervals. The BDP-96-1 and BDP-96-2 cores were sampled each 2 cm, the BDP-98 core was sampled each 10 cm. The age 74

Evolution of the extinct genera belonged to the family Stephanodiscaceae (Bacillariophyta)

Taxon

Age range

Endemic (+) or extinct endemic (++)

Extinct species

Extant species

Table 1. Systematic composition of fossil diatoms of the family Stephanodiscaceae from the Upper Cenozoic sediments of Lake Baikal.

1

2

3

4

5

Division Bacillariophyta Class Coscinodiscophyceae Round & Crawford Subclass Archaegladiopsophycidae Nikolaev & Harwood Order Thalassiosirales Gleser & Makarova Family Stephanodiscaceae Makarova Genus Concentrodiscus Khursevich, Moisseeva & Sukhova C. indigenus Khursevich & Fedenya C. kuzminii Khursevich & Fedenya C. proteus Khursevich & Fedenya C. subabnormis Khursevich & Fedenya Genus Mesodictyon Theriot & Bradbury M. nativus Khursevich & Fedenya Genus Mesodictyopsis Khursevich, Iwashita, Kociolek & Fedenya M. academicus Khursevich, Iwashita, Kociolek & Fedenya M. baicalensis Khursevich, Iwashita, Kociolek & Fedenya M. insolitus Khursevich & Fedenya M. medius Khursevich & Iwashita M. peculiaris Khursevich, Kociolek & Fedenya M. similis Khursevich & Fedenya M. singularis Khursevich, Iwashita & Fedenya Genus Tertiariopsis Khursevich & Kociolek T. imperseptus Khursevich, Fedenya & Kociolek T. makarovae Khursevich & Kociolek T. sibericus Khursevich, Fedenya & Kociolek Genus Stephanopsis Khursevich & Fedenya S. costatus Khursevich & Fedenya Genus Stephanodiscus Ehrenberg S. asteroides var. baicalensis Khursevich & Fedenya S. baicalensis Likhoshway & Pomazkina S. baicalensis var. concinnis Pomazkina & Likhoshway S. binderanoides Khursevich & Fedenya S. binderanus (Kützing) Krieger var. binderanus S. binderanus var. hyalinus Khursevich & Fedenya

75

lMi lMi lMi lMi

++ ++ ++ ++

lMi

++

lMi lMi - ePl lMi lMi lMi LMi – ePl lMi

++ ++ ++ ++ ++ ++ ++

ePl ePl ePl

++ ++ ++

ePl

++

mPlei lPlei mPlei – lPlei mPlei lMi – R lPl

++ ++ ++ ++ + ++

Galina Khursevich

Table 1. Continued 1

2

S. carconeiformis Khursevich & Loginova S. compactus Khursevich & Fedenya S. dissimilis Khursevich & Fedenya S. distinctus Khursevich var. distinctus S. distinctus var. excentricoides Khursevich S. exiguus Khursevich S. flabellatus Khursevich & Loginova S. formosus Khursevich & Loginova S. grandis Khursevich & Loginova var. grandis S. grandis var. entis Khursevich S. hantzschii Grunow S. imperpetuus Khursevich & Fedenya S. inconspicuus Makarova & Pomazkina S. invisitatus Hohn & Hellerman S. jucundus Khursevich & Fedenya S. khurseviczae Likhoshway S. majusculus Khursevich & Fedenya S. meyerii Genkal & Popovskaya S. minutulus (Kützing) Cleve & Möller S. cf. niagarae Ehrenberg S. notabilis Khursevich & Fedenya S. parvus Stoermer & Håkansson S. princeps Khursevich S. veneris Khursevich & Fedenya S. williamsii Khursevich S. yukonensis var. antiquus Khursevich Genus Tertiarius Håkansson & Khursevich T. baicalensis Khursevich & Fedenya Genus Cyclotella (Kützing) Brébisson C. andancensis Ehrlich C. baicalensis Skvortzow C. comtaeformica Khursevich var. comtaeformica C. comtaeformica var. spinata Khursevich C. distincta Khursevich C. distinguenda Hustedt C. foveolata Khursevich & Fedenya C. gracilis Nikiteeva & Likhoshway C. iris Brun & Héribaud var. iris C. iris var. charetoni (Héribaud) Serieyssol

mPlei – lPlei mPlei ePlei mPlei mPlei lPl – mPlei lPlei – Hol mPlei – lPlei mPlei – lPlei mPlei – lPlei Pl – R ePlei lPlei – R Hol – R ePlei lPlei ePlei Hol – R lMi – R Pl – R ePlei Plei – R mPlei mPlei ePlei ePlei

++ ++ ++ ++ ++ ++ ++ ++ ++ ++

lPl

++

lMi – Pl? lPlei – R ePlei ePlei lPl Pl – R lMi lPlei lMi – R lMi – R

76

3

4

5

+ ++ + + ++ ++ ++ + + + ++ + ++ ++ ++ ++

+ + ++ ++ ++ + ++ ++ + +


Table 1. Continued 1

2

C. iris var. combierensis Serieyssol C. iris var. insueta Khursevich C. iris var. integra Peragallo & Héribaud C. iris var. ovalis Brun & Héribaud C. krammeri Håkansson C. meneghiniana Kützing C. minuta (Skvortzow) Antipova C. ocellata Pantocsek C. ornata (Skvortzow) Flower C. praeminuta Khursevich C. pseudostelligera Hustedt C. tempereiformica Khursevich Genus Cyclostephanos Round C. dubius (Fricke) Round

lMi lMi - ePl lMi – Pl lMi – R Pl – R Pl – R mPlei – R Pl – R mPlei – R mPlei Pl – R lPl lPl – R

3

4

5

+ ++ + + + + + + + ++ + ++ +

Legend: lMi – late Miocene, Pl – Pliocene, ePl – early Pliocene, lPl – late Pliocene, Plei – Pleistocene, ePlei – early Pleistocene, mPlei – middle Pleistocene, lPlei – late Pleistocene, Hol – Holocene, R – Recent.

model for these cores is based on the identification of paleomagnetic event/reversal boundaries and the correlation with marine oxygen isotope curve (Williams et al. 1997), as well as on 10Be isotope geochronology for the BDP-98 drill core (Sapota et al. 2004). Permanent diatom slides were prepared with identical volumes of material according to the method described in Grachev et al. (1997). Diatom abundance (mln valves per gram of dry sediment) was calculated in each slide using quantitative method. Specimens were examined with Ergaval light microscope with an oil immersion objective (100x, NA=1.25). In order to precisely define taxonomic belonging of diatoms, the ultrastructure of their valves was studied using JEOL (JSM–35C) scanning electron microscope. The terminology is recommended by Anonymous (1975) and Ross et al. (1979). A system of diatoms proposed by Round et al. (1990) and refined for centric diatoms by Nikolaev and Harwood (2001) has been used in the paper. The information about the age range of species and intraspecific taxa of the family Stephanodiscaceae found in the Upper Cenozoic sediments of Lake Baikal was taken from numerous publications (Ehrlich 1966, Serieyssol 1981, 1984, Fourtanier and Gasse 1988, Khursevich 1989, 1999, Serieyssol and Gasse 1991, Gleser et al. 1992, Nikiteeva and Likhoshway 1994, Likhoshway 1996, Ognjanova-Rumenova 1996, 2001, Julius et al. 1997, Temniskova-Topalova and OgnjanovaRumenova 1997, Edlund and Stoermer 2000, Khursevich et al. 2000, 2001a, b, c, 2002a, b, 2003a, b, c, 2004, 2005, Rasskazov et al. 2001, Khursevich and Fedenya 2005).

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RESULTS AND DISCUSSION Upper Cenozoic (Upper Miocene – Holocene) sediments of Lake Baikal contain the diverse composition of centric diatoms from the family Stephanodiscaceae. The latter includes 60 species and 12 intraspecific taxa belonging to 9 genera. Among them, 6 genera (3 endemic for Lake Baikal and the Transbaikal area), 42 species (41 baikalian endemics) and 10 intraspecific taxa (8 baikalian endemics) are extinct (Table 1). Morphological peculiarities of some members of extinct genera from the family Stephanodiscaceae found in the Upper Miocene and Pliocene deposits from Lake Baikal have been illustrated in Plates 1 – 3 (Figs 1– 26). The extinct genus Concentrodiscus is known at present from the Paleo-Baikal and the ancient basins of the Transbaikal area. This genus and its type species Concentrodiscus abnormis Khursevich, Moisseeva and Sukhova were first described from the Middle-Upper Miocene diatomaceous rocks of the Upper-Sulbanian depression within the Transbaikal area (Khursevich et al. 1989). Later a new member of this genus, Concentrodiscus variabilis Khursevich & Chernyaeva, was defined in the Middle Miocene deposits within the Amalat plateau of the Transbaikal area (Khursevich and Chernyaeva 1994, Rasskazov et al. 2001). Four new representatives of Concentrodiscus were found in the Upper Miocene stratum from the BDP-98 drill core (Khursevich et al. 2002a, Khursevich and Fedenya 2005). Hence, the age range of this genus is from the Middle to Late Miocene. The genus Concentrodiscus is characterized by the following distinctive features: a) the presence of valves with one-two concentric convex and concave zones; b) the availability of internal flat cribra in areolae; c) the presence of marginal fultoportulae with 4 satellite pores; d) the various position of a single rimoportula on the valve surface (near the centre, in the submarginal zone of the valve face, at the valve face/mantle junction, on the mantle); e) the presence of valve face fultoportulae with 4 satellite pores in both isotopic and heterotopic position, rarely the absence of them. The age range of this genus in the ancient Baikal ecosystem is Late Miocene (ca. 8–7 Ma), according to the age model for the BDP-98 core (Sapota et al. 2004). However, this geological age may be extended because for the present the complete Miocene diatom record from Lake Baikal is absent. The genus Mesodictyon (Theriot and Bradbury 1987) is restricted mainly to the Late Miocene. It is widespread and diverse in lacustrine deposits of that time in the western regions of USA (Krebs et al. 1987, Theriot 1990, Krebs 1994, Khursevich and VanLandingham 1995), Ethiópia (Fourtanier and Gasse 1988), France (Serieyssol and Gasse 1991), northern Peru (Fortanier et al. 1993), southern Bulgaria (Temniskova-Topalova and Ognjanova-Rumenova 1997), Belarus (Rylova et al. 1999). The presence of the new fossil species Mesodictyon nativus in the Upper Miocene deposits of Lake Baikal (Khursevich et al. 2002a) is the first record of this genus within Asia. Thus, the genus Mesodictyon belongs to one of the important biochronological markers and can be used for cross-continental correlation of lacustrine sedimentary sections. The genus Mesodictyon is distinguished by the following morphological peculiarities: a) the presence of flat or concentrically undulate valves; b) the location of the areola cribrum inside of the loculus; c) the presence of marginal fultoportulae with 2 satellite pores; d) the position 78


2

1

3

4 6

5

7

8

Figs 1, 2, 5, 8. Concentrodiscus kuzminii Khursevich & Fedenya, SEM. Scale bars = 10 µm (Figs 1, 5) or 1 µm (Figs 2, 8). Fig. 1. External view of the valve surface with tubular extensions of marginal fultoportulae (arrows). Figs 2, 5. Fragments of the internal valve surface with flat cribra in areolae, a ring of marginal fultoportulae with four satellite pores (arrow in Fig. 2) and a single rimoportula on the valve face/mantle junction (arrow in Fig. 5). Fig. 8. Detail showing round external aperture of rimoportula (arrow). Figs 3, 4, 6, 7. Concentrodiscus indigenus Khursevich & Fedenya, SEM. Scale bars = 1 µm. Figs 3, 6. External view of the valve surface with a ring of small openings of valve face fultoportulae near the centre (arrow in Fig. 6). Fig. 4. Fragment of the internal valve surface with a single rimoportula on the mantle (black arrow) and a ring of marginal fultoportulae with four satellite pores (white arrows). Fig. 7. Detail showing a distinct opening of rimoportula on the mantle (arrow).

79

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of a single rimoportula on the valve mantle; e) the absence of valve face fultoportulae. The age range of this genus in the ancient Lake Baikal: Late Miocene (ca. 7.7–6.3 Ma). The genus Mesodictyopsis belongs to extinct endemics of the ancient lacustrine ecosystems of Lake Baikal and the Transbaikal area (Khursevich et al. 2004). At present it contains 8 species. Among them, 7 members of Mesodictyopsis are known from the Late Miocene–the beginning of Pliocene of Lake Baikal, and 1 taxon apparently from the Late Miocene of the Dzhilinda hollow (the Transbaikal area). The genus Mesodictyopsis is diagnosed by a combination of the following morphological features: a) the presence of medial velum (cribrum?) in areolae; b) the presence of marginal fultoportulae with 3 satellite pores; c) the various location of rimoportula(e) on the valve surface (near the centre, in the submarginal zone of the valve face, at the valve face/mantle junction, on the mantle); d) the presence of valve face fultoportulae mainly with 3 ( rarely 2 or 4) satellite pores. The age range of this genus in the ancient Lake Baikal is ca. 7–5.1 Ma. The extinct genus Tertiariopsis and its three species were described from the Lower Pliocene deposits of Lake Baikal (Khursevich et al. 2002b). This genus may include also two species described by Serieyssol et al. (1998) from the Upper Miocene sediments of France and Mexico. These species, originally placed in Thalassiosira, lack alveolae and appear to have a valve mantle dissected by hyaline strips. The genus Tertiariopsis possesses by the following characteristic peculiarities: a) the presence of internal raised cribra in areolae; b) the availability of a distinct break between the valve face areolae and those of the mantle; c) the breaking up of the valve mantle onto distinct areolar sections separated by hyaline strips; d) the presence of marginal fultoportulae with 3 satellite pores covered by a marginal lamina internally; e) the position of one-two rimoportulae mainly on the valve mantle; f) the presence of valve face fultoportulae with usually 3 (rarely 2) satellite pores. The age range of this genus in the ancient Lake Baikal: ca. 5.1- 4.6 Ma. The genus Stephanopsis has been found only in the Pliocene sediments from the BDP98 and BDP-96-1 drill cores and belongs to one of the extinct endemic genera of the ancient Lake Baikal (Khursevich et al. 2000). The type species of this genus, Stephanopsis costatus, is characterized by a great morphological variability (at least 5 morphotypes were revealed in the studied material). The genus Stephanopsis is distinguished by the following characters: a) the presence of internal raised cribra in areolae; b) the presence of thin radial costae crossing the valve mantle internally; c) the location of marginal fultoportulae with 3 satellite pores at the proximal ends of thin radial internal costae; e) the position of rimoportula(e) on the valve surface mainly at the proximal end of one or several thin radial internal costae, sometimes elsewhere on the valve face, but usually near the centre; f) the presence of valve face fultoportulae with 2 or 3 satellite pores. The age range of this genus in the ancient Lake Baikal is ca. 4.84 – 2.6 Ma, according to the age model for the BDP-96-1 and BDP-98 cores (Williams et al. 1997, Sapota et al. 2004). The genus Tertiarius was first proposed for three members of the genus Cyclotella occurred in the Pliocene material from Köpecz (Transylvania, Romania) in Pantocsek’s Collection and studied in SEM (Håkansson and Khursevich 1997). Later new species of 80


9

11

10

12

13

14

15

16

Figs 9 -11. Mesodictyon nativus Khursevich & Fedenya, SEM. Scale bars = 1 µm. Fig. 9. External view of the valve surface. Figs 10, 11. Internal view of the valve surface with a single sessile rimoportula on the mantle (arrow in Fig. 10) and a ring of marginal fultoportulae with two satellite pores (arrow in Fig. 11). Figs 12, 13. Mesodictyopsis singularis Khursevich, Iwashita & Frdenya, SEM. Scale bars= 1 µm. Fig. 12. Internal view of the valve surface with a single rimoportula on the valve face/mantle junction (arrow). Fig. 13. Detail showing medial velum within areolae (arrow). Fig. 14 – 16. Mesodictyopsis medius Khursevuch & Iwashita, SEM. Scale bars = 1 µm. Fig. 14. External view of the valve surface. Fig. 15. Internal view of the valve surface with a single rimoportula in the centre (arrow) and a ring of marginal fultoportulae with three satellite pores. Fig. 16. Detail showing medial velum within areolae (arrow).

81

Galina Khursevich

17

20

18

19

21

22

23

24

25

26

Figs 17-18. Tertiariopsis sibericus Khursevich, Fedenya & Kociolek, SEM. Scale bars = 1 µm. Fig. 17. External view of the valve surface with a ring of marginal fultoportulae openings (arrow). Fig. 18. Internal view of the valve surface with a single rimoportula on the mantle (arrow) and a ring of valve face fultoportulae with three satellite pores near the centre. Fig. 19. Tertiariopsis imperseptus Khursevich, Fedenya & Kociolek, SEM. Scale bar = 1 µm. Internal view of the valve surface with a single rimoportula on the mantle (arrow). Figs 20 – 22. Stephanopsis costatus Khursevich & Fedenya, SEM. Scale bars = 10 µm. Fig. 20. External view of the valve surface. Figs 21, 22. Internal view of the valve surface with the presence of thin radial costae crossing the mantle, three rimoportulae located near the centre (arrow in Fig. 21) or rimoportula positioned at the inner end of one of thin costae (white arrow in Fig. 22) and a ring of marginal fultoportulae with three satellite pores (black arrow in Fig. 22). Figs 23 – 26. Tertiarius baicalensis Khursevich & Fedenya, SEM. Scale bars = 1 µm. Figs 23, 25. External view of the valve surface. Figs 24, 26. Internal view of the valve surface with the marginal ring of alveolae and a single rimoportula (arrows).

82


Tertiarius have been identified in the Pliocene lacustrine sediments from the Former Yugoslavian Republic of Macedonia (Ognjanova-Rumenova 2001) and the Upper Miocene and Pliocene deposits from western USA (Khursevich and Kociolek 2002). The occurrence of Tertiarius baicalensis in the Upper Pliocene sediments from the ancient Lake Baikal is the first report of this genus in Asia (Khursevich et al. 2003a). Thus, the age range of this genus is restricted to the Late Miocene–Pliocene. The genus Tertiarius is diagnosed by a combination of the following morphological features: a) the presence of more or less flat valves; b) the availability of internal domed cribra in areolae on the valve face; c) the presence of alveolae on the valve mantle; d) the location of marginal fultoportulae with 2 or 3 satellite pores on the thick costae internally; d) the various position of rimoportula(e) on the valve surface (in the submarginal zone of the valve face, at the base of internal costa(e) or within the alveolar chamber(s); e) the presence of valve face fultoportulae with 2 or 3 satellite pores. The age range of this genus in the ancient Lake Baikal is ca. 2.7–2.5 Ma. The extinct genera from the family Stephanodiscaceae characterized above replace each other from the bottom to top along the long continuous sections of the BDP-98 (Fig. 1) and BDP-96-1 cores that allows to trace the evolution of some morphological characters in time. Pattern of morphological evolution in these genera in the ancient Baikal ecosystem can be described in the following ways: a) change of the cribrum position in areola from the internal flat cribrum (the genus Concentrodiscus) to the medial cribrum located within loculus (the genera of Mesodictyon and Mesodictyopsis), later again to the internal but slightly raised cribrum (the genera of Tertiariopsis and Stephanopsis), and lastly to the internal domed cribrum (the Tertiarius genus); b) change of the stria morphology from areolar stria (the genera of Concentrodiscus, Mesodictyon, Mesodictyopsis, Tertiariopsis, Stephanopsis) to areolar-alveolar stria (the genus Tertiarius); c) change of the structure of marginal fultoportulae on the valve mantle (the species of Concentrodiscus have marginal fultoportulae only with 4 satellite pores, whereas the members of Mesodictyon, Mesodictyopsis, Tertiariopsis, Stephanopsis and Tertiarius possess marginal fultoportulae with 2 or 3 satellite pores); d) change of the number of rimoportulae (if the more ancient genus Concentrodiscus has a single rimoportula on the valve surface, then the younger genera Mesodictyopsis, Tertiariopsis, Stephanopsis and Tertiarius have from one to several, sometimes up to seven rimopotulae forming a marginal ring). At the same time the extinct endemic genera of Concentrodiscus, Mesodictyopsis and Stephanopsis developed in the ancient Lake Baikal have some common inherited morphological features (such as the various placement of rimoportula on the valve surface including certain taxa with the location of rimoportula in or near the valve centre). Besides the inherited position of rimoportula in or near the centre of the valve is characteristic also of some ancient members of Stephanodiscus (S. dissimilis Khursevich and Fedenya, S. majusculus Khursevich and Fedenya and S. yukonensis var. antiquus Khursevich) which appeared and developed in Lake Baikal in early Pleistocene (Khursevich et al. 2001b, 2003a). Thus, the evolution of centric diatoms in the ancient Baikal ecosystem during the last 8 Ma is associated with the processes of extinction and neogeneration both of some genera and many species. Percentage of baikalian extinct endemics in the family Stephanodiscaceae makes up 70%. This diatom group has evolved in situ rapidly throughout the late 83

AGE

L.

M.

Early

Late

Early

Ma

9

8

7

6

5

4

3

2

1

0

MAG. POL.

8

7

6

5

C4A 9

C4

C3B

C3A

C3

C2A

C2

C1

CHRON Brunhes Matuyama Gauss Gilbert

Pleistocene

Pliocene

Miocene

84

Late

BDP96 depth, m BDP98 depth, m 600

550

500

450

400

350

300

250

200

150

100

50

0

600 1200 0

u A 500 1000 0

30

100 200 0

5

10 0

400 800 0

200 400 0

is ps n yo yo lla ct ct i i te od od lo s s c e e M Cy M

us sc

mln valves per gram of dry sediment

60 0

us di cl ro cy nt o e in nc ct Co A

300 600 0

o ol ve Al

a or ph

BDP96-1, BDP96-2 and BDP98 cores

0

s om at Di

ira se co la ti

is

400 800 0

no ha

us sc di

Concentrodiscus kuzminiiC. indigenusbenthic

Mesodictyon nativus

C.comtaeformica et var. spinata C.tempereiformicaC.distincta Tertiarius baicalensis Stephanopsis costatus Stephanopsis costatusAulacoseira sp. aff. A. islandica Tertiariopsis sibericusT. imperseptus Cyclotella iris group Cyclotella iris group -Mesodictyopsis similis Aulacoseira praegranulata -M. similis Mesodictyopsis baicalensisM.similis Cyclotella iris group -Mesodictyopsis baicalensis Mesodictyopsis singularis Mesodictyopsis academicus M. peculiaris Mesodictyopsis academicusMesodictyon nativus Mesodictyopsis insolitusM. medius Alveolophora baicalensisConcentrodiscus subabnormis -Actinocyclus immemoratus

* ** *** ****

Brunhes diatom zones

Diatom zones

Correction: 15 March 2005

600 1200

ep St

500 1000 0

s si s op an riu h ia rt ep St Te

500 1000 0

r Te

ps io ar

Galina Khursevich

Fig. 1. Diatom biostratigraphy of Lake Baikal sediments in BDP-96 and BDP-98 cores. The correlation of BDP-96 and BDP-98 cores with geomagnetic polarity time scale of Cande and Kent (1995) is given according to Williams et al. (1997), BDP-Members (2000) and Sapota et al. (2004). Diatom zones: * Stephanodiscus notabilis–S. binderanus–Synedra acus var. radians, ** Stephanodiscus jucundus– S. williamsii–Synedra ulna var. danica, *** Aulacoseira subarctica–Cyclotella ocellata, ****Stephanodiscus majusculus– Aulacoseira sp. aff. A. islandica


Miocene–Pleistocene, producing a number of short-ranging species. The ancient faultblock basin of Lake Baikal contains the endemic lineages within the centric genera of Concentrodiscus, Mesodictyopsis, Stephanodiscus and Cyclotella. Key for determination of the extinct genera belonged to the family Stephanodiscaceae from the Upper Cenozoic sediments of Lake Baikal Presence of loculate areolae with foramina on the external valve surface and cribra on the internal valve surface or within loculae. Striae areolate, areolate-alveolate or only alveolate. Marginal fultoportulae with two or three, rarely with four satellite pores. Rimoportulae from one to several have various position on the valve .............................. Stephanodiscaceae I. Striae areolate. 1. Areolae in no clear radial rows. Marginal fultoportulae with four satelliteores .......... ........................................................................................................ Concentrodiscus 2. Areolae in distinct radial rows. Marginal fultoportulae with two or three satellite pores. A. Presence of medial cribra in areolae. a. Marginal fultoportulae with two satellite pores. Valve face fultoportulae absent. The constant position of a single rimoportula on the mantle .... .............................................................................................................. Mesodictyon b. Marginal fultoportulae with three satellite pores. Valve face fultoportulae present. The various location of rimoportula(e) on the valve surface .... .......................................................................................................... Mesodictyopsis B. Presence of internal raised cribra in areolae. a. Marginal fultoportulae covered by a marginal lamina internally. The constant position of one-two rimoportulae on the mantle ... Tertiariopsis b. Marginal fultoportulae placed at the base of thin radial costae crossing valve mantle internally. The various location of one-several rimoportulae on the valve surface .................................................................. Stephanopsis II. Striae areolate-alveolate .................................................................................................. Tertiarius ACKNOWLEDGEMENTS This work was implemented as a part of Baikal Drilling Project supported by National Scientific Foundation of USA (NSF) grant EAR-96-14770, the Siberian Branch of Russian Academy of Sciences, the Russian Ministry of Geology, the Science and Technology Agency (STA) of Japan. Author thanks the entire NEDRA Scientific Drilling Team for the successful drilling effort. I am grateful to the scientists of the Institute of Geochemistry, Limnological Institute and the Institute of the Earth Crust (Irkutsk) who participated in primary description and sampling of the BDP-96-1 and BDP-98 drill cores, as well as in permanent slide preparation.

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REFERENCES ANONYMOUS 1975. Proposals for a standartization of diatom terminology and diagnoses. – Nova Hedwigia, Beih. 53: 325-354. BDP-Members 1997. Continuous paleoclimate record recovered for last 5 million years. – EOS American Geophysical Union, Transactions, 78(51): 597-604. BDP-MEMBERS 2000. Paleoclimatic record in the Late Cenozoic sediments of Lake Baikal (by 600 m deep-drilling data). – Russ. Geol. Geophys., 41(1): 3-32. (in Russian) BDP-MEMBERS 2001. The new BDP-98 600-m drill core from Lake Baikal: a key late Cenozoic sedimentary section in continental Asia. – Quatern. Intern., 80-81: 19-36. CANDE, S.C. and D.F. KENT 1995. Revised calibration of the geomagnetic polarity time scale for the Late Cretaceous and Cenozoic. – J. Geophys. Res., 100: 6093–6095. EDLUND, M.B. and E.F. STOERMER 2000. A 200,000-year, high-resolution record of diatom productivity and community makeup from Lake Baikal shows high correspondence to the marine oxygen-isotope record of climate change. – Limnol. Oceanogr., 45(4): 948-962. EHRLICH, A. 1966. Contributions à l’étude des gisements volcano-lacustres à Diatomées de la région de Rochessauve et de Saint Bauzile (Ardèche). – Bull. Soc. Géol. France, ser. 7 (7): 311-321. FOURTANIER, E. and F. GASSE 1988. Premiers jalons d’une biostratigraphie et évolution des diatomées lacustres d’Afrique depuis 11 Ma. - C. R. Acad. Sci. Paris, 306: 1401-1408. FOURTANIER, E., F. GASSE, O. BELLIER, M.G. BONHOMME and I. ROBLES 1993. Miocene non-marine diatoms from the western Cordillera basins of northern Peru. - Diatom Research, 8: 13-30. GLESER, Z.I., I. V. MAKAROVA, A.I. MOISSEEVA, and 6 OTHERS 1992. The diatoms of the USSR (fossil and recent). 2(2). Nauka, St.-Petersburg. (in Russian) GRACHEV, M.A., E.V. LIKHOSHWAY, S.S. VOROBYOVA, and 19 OTHERS 1997. Signals of the paleoclimates of the Upper Pleistocene in the sediments of Lake Baikal. – Russ. Geol. Geophys., 38: 957-980. (in Russian) HÅKANSSON, H. and G. KHURSEVICH 1997. Tertiarius gen. nov. – genus in the Bacillariophyceae, the transfer of some cyclotelloid species and a comparison to closely related genera. – Diatom Research, 12: 19-33. JULIUS, M.L., E.F. STOERMER, S.M. COLMAN and T.C. MOORE 1997. A preliminary investigations of siliceous microfossil succession in late Quaternary sediments from Lake Baikal, Siberia. – J. Paleolimnol., 18: 187-204. KHURSEVICH, G.K. 1989. Atlas of the species of Stephanodiscus and Cyslostephanos (Bacillarophyta) from the Upper Cenozoic sediments of the USSR. F. Ju. Velichkevich, editor. Nauka i tekhnika, Minsk. (in Russian) KHURSEVICH, G.K. 1999. Morphological peculiarities of some Stephanodiscus species from the bottom sediments of Lake Baikal. in: Mayama, Sh., M. Idei and I. Koizumi, editors. Proceedings of the 14th International Diatom Symposium: 603-612. Koeltz Scientific Books, Koenigstein. 86


KHURSEVICH, G.K. and G.P. CHERNYAEVA 1994. A new species of the genus Concentrodiscus (Bacillariophyta) from the Miocene deposits of the Transbaikal area. – Botanichesky Zhurnal, 79(1): 107-109 (in Russian). KHURSEVICH, G.K. and S.A. FEDENYA 2005. Morphology of new fossil species of Concentrodiscus and Alveolophora (Bacillariophyta) from bottom sediments of Lake Baikal. – Algologia (accepted). KHURSEVICH, G.K., S.A. FEDENYA, E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS and M.I. KUZMIN 2001a. Late Cenozoic diatom record from Lake Baikal sediments. in: A. Economou-Amilli, editor. Proceedings of the 16th International Diatom Symposium: 451-460. University of Athens, Athens. KHURSEVICH, G.K., S.A. FEDENYA, E.B. KARABANOV, D.F. WILLIAMS and M.I. KUZMIN (2000): Stephanopsis Khursevich & Fedenya – new genus of class Centrophyceae (Bacillariophyta) from the Pliocene deposits of Lake Baikal (Russia). – Algologia, 10(1): 106-109. KHURSEVICH, G.K., S.A. FEDENYA, M.I. KUZMIN, E.B. KARABANOV, D.F. WILLIAMS and A.A. PROKOPENKO 2002a. Morphology of new species of Concentrodiscus and Mesodictyon (Bacillariophyta) from the Upper Miocene deposits of Lake Baikal. Algologia, 12(3): 361-370. KHURSEVICH, G.K., S.A. FEDENYA, M.I. KUZMIN, E.B. KARABANOV, D.F. WILLIAMS and A.A. PROKOPENKO 2003a. Morphology of new taxa of the class Centrophyceae (Bacillariophyta) from the Pliocene and Pleistocene deposits of Lake Baikal, Siberia. – Algologia, 13(3): 305-321. KHURSEVICH, G.K., S.A. FEDENYA, M.I. KUZMIN, E.B. KARABANOV, D.F. WILLIAMS and A.A. PROKOPENKO 2003b. New species of Stephanodiscus (Bacillariophyta) from the Pleistocene sediments of Lake Baikal. – Algologia, 13(4): 389-401. KHURSEVICH, G., E. KARABANOV, M. KUZMIN, D. WILLIAMS, A. PROKOPENKO and S. FEDENYA 2003c. Diatom succession in Upper Miocene sediments of Lake Baikal from BDP-98 drill core. in: K. Kashiwaya, editor. Long Continental Records from Lake Baikal: 271-282. Springer Verlag, Tokyo. KHURSEVICH, G.K., E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, M.I. KUZMIN and S.A. FEDENYA 2001b. Biostratigraphic significance of new fossil species of the genera Stephanodiscus and Cyclotella from Upper Cenozoic deposits of Lake Baikal, Siberia. – Micropaleontology, 47(1): 47-71. KHURSEVICH, G.K., E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, M.I. KUZMIN, S.A. FEDENYA and A. N. GVOZDKOV 2001c. Insolation regime in Siberia as a major factor controlling diatom production in Lake Baikal during the past 800,000 years. – Quatern. Intern., 80-81: 47-58. KHURSEVICH, G.K. and J. KOCIOLEK 2002. New Tertiarius (Bacillariophyta: Stephanodiscaceae) species from Western North America. in: J. John, editor. Proceedings of the 15th International Diatom Symposium: 331-349. Biopress Limited, Bristol. KHURSEVICH, G.K., J.P. KOCIOLEK and S.A. FEDENYA 2002b. A new genus of fossil freshwater diatoms (Bacillariophyta: Stephanodiscaceae) from the sediments of Lake Baikal. - Proceedings of the California Academy of Sciences, 53(1): 1–10. 87

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KHURSEVICH, G.K., J.P. KOCIOLEK, T. IWASHITA, S.A. FEDENYA, M.I. KUZMIN, T. KAWAI, D.F. WILLIAMS, E.B. KARABANOV, A.A. PROKOPENKO and K. MINOURA 2004. Mesodictyopsis Khursevich, Iwashita, Kociolek & Fedenya – new genus of class Centrophyceae (Bacillariophyta) from Upper Miocene sediments of Lake Baikal, Siberia. – Proceedings of the California Academy of Sciences, 55(15): 336–355. KHURSEVICH, G.K., A.I. MOISSEEVA and G.A. SUKHOVA 1989. New genus of diatom algae of the family Stephanodiscaceae. - Botanichesky Zhurnal, 74(11): 1600-1601. (in Russian) KHURSEVICH, G.K., A.A. PROKOPENKO, S.A. FEDENYA, L.I. TKACHENKO and D.F. WILLIAMS 2005. Diatom biostratigraphy of Lake Baikal during the past 1.25 Ma: new results from BDP-96-2 and BDP-99 drill cores. – Quatern. Intern., 136: 95-104. KHURSEVICH, G.K. and S.L. VANLANDINGHAM 1995. Morphology and stratigraphy of some Mesodictyon species (Bacillariophyta) from upper Miocene freshwater deposits of Idaho and Nevada, U.S.A. – Nova Hedwigia, 60(3-4): 467-478. KREBS, W.N. 1994. The biochronology of freshwater planktonic diatom communities in western North America. in: J.P. Kociolek, editor. Proceedings of the 11th International Diatom Symposium: 485-499. California Academy of Sciences, San Francisco. KREBS, W.N., J.P. BRADBURY and E. THERIOT 1987. Neogene and Quaternary lacustrine diatom biochronology, western USA. – Palaios, 2: 505—513. LIKHOSHWAY, YE.V. 1996. Stephanodiscus khurseviczae sp. nov. from Pleistocene sediments of Lake Baikal. – Diatom Research, 11(2): 273-281. NIKITEEVA, T.A. and YE.V. LIKHOSHWAY 1994. Cyclotella gracilis sp. nov. from Pleistocene material of Lake Baikal. – Diatom Research, 9(2): 349 – 353. NIKOLAEV, V.A. and D.M. HARWOOD 2001. Diversity and classification of centric diatoms. in: A. Economou-Amilli, editor. Proceedings of the 16th International Diatom Symposium: 127-152. University of Athens, Athens. OGNJANOVA–RUMENOVA, N.G. 1996. Cyclotella iris Brun & Héribaud – a group from the Upper Miocene sediments of the Sofia basin, Bulgaria. – Geologica Carpathica, 47(5): 301–310. OGNJANOVA–RUMENOVA, N.G. 2001. Neogene diatom assemblages from lacustrine sediments of F.Y.R.O.M. (Macedonia) and their distribution in correlative formations of south-western Bulgaria. in: A. Economou-Amilli, editor. Proceedings of the 16th International Diatom Symposium: 423-432. University of Athens, Athens. RASSKAZOV, S.V., N.A. LOGACHEV, A.V. IVANOV, V.A. MISHARINA, G.P. CHERNYAEVA, I.S. BRANDT, S.B. BRANDT, V.M. SKOBLO and N.A. LYAMINA 2001. Palynological and diatom analyses of sediments from the late Cenozoic Amalat valley. – Russ. Geol. Geophys., 42(5): 773-785. (in Russian) ROSS, R., E.J. COX, N.I. KARAYEVA, D.G. MANN, T.B.B. PADDOCK, R. SIMONSEN and P.A. SIMS 1979. An emended terminology for the siliceous component of the diatom cell. – Nova Hedwigia, 64: 513-533. ROUND, F.E., R.M. CRAWFORD and D.G. MANN 1990. The diatoms: biology and morphology of the genera. – Cambridge University Press, Cambridge. 88


RYLOVA, T.B., T.V. YAKUBOVSKAYA and G.K. KHURSEVICH 1999. Palaeobotanical evidence for correlating the stratigraphy of the Neogene deposits of Belarus. – Acta Palaeobotanica, Suppl. 2.: 359-363. SAPOTA, T., A. ALDAHAN, G. POSSNERT, J. PECK, J. KING, A. PROKOPENKO and M. KUZMIN 2004. A late Cenozoic Earth’s crust and atmosphere dynamics record from an active continental rift system. – J. Paleolimnol., 32: 341-349. SERIEYSSOL, K. 1981. Cyclotella species of Late Miocene age from St. Bauzile, France. in: R. Ross, editor. Proceedings of the 6th Symposium on Recent and Fossil Diatoms: 27-42. O. Koeltz, Koenigstein. SERIEYSSOL, K. 1984. Cyclotella iris Brun & Héribaud. in: D.G. Mann, editor. Proceedings of the 7th International Diatom Symposium: 197-212. O. Koeltz, Koenigstein. SERIEYSSOL, K. and F. GASSE 1991. Diatomées néogènes du Massif Central Français: quelques faits biostratigraphiques. - C. R. Acad. Sci. Paris, 312: 957-964. SERIEYSSOL, K., I. ISRADE GARDUNO and F. GASSE 1998. Thalassiosira dispar comb. nov. and T. cuitzeonensis spec. nov. (Bacillariophyceae) found in Miocene sediments from France and Mexico. – Nova Hedwigia, 66 (1-2): 177-186. TEMNISKOVA-TOPALOVA, D. and N. OGNJANOVA-RUMENOVA 1997. Description, comparison and biostratigraphy of the nonmarine Neogene diatom floras from Southern Bulgaria. – Geologica Balcanica, 27(1-2): 57-81. THERIOT, E. 1990. New species of Mesodictyon (Bacillariophyta: Thalassiosiraceae) in Late Miocene lacustrine deposits of the Snake River basin, Idaho. - Proceedings of the Academy of Natural Sciences of Philadelphia, 142: 1-19. THERIOT, E. and J.P. BRADBURY 1987. Mesodictyon, a new fossil genus of the centric diatom family Thalassiosiraceae from the Miocene Chalk Hills Formation, western Snake River Plain, Idaho. – Micropaleontology, 33: 356-367. WILLIAMS, D.F., J. PECK., E.B. KARABANOV, A.A. PROKOPENKO, V.KRAVCHINSKY, J. KING and M.I. KUZMIN 1997. Lake Baikal record of continental climate response to orbital insolation during the past 5 million years. – Science, 278: 1114 –1117.

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A new Cymbella from the Neogene in Bulgaria and its stratigraphic significance Nadja Ognjanova-Rumenova1 and Ditmar Metzeltin2 Department of Palaeontology and Stratigraphy, Institute of Geology, Bulgarian Academy of Sciences, Acad. G.Bonchev str. 24, 1113 Sofia, Bulgaria E-mail: [email protected] 2 Am Stegskreuz 3b, D-65719 Hofheim, Germany 1

ABSTRACT A new species, Cymbella serdica is described and illustrated from the Upper Miocene Novi Iskar Formation of the Sofia Neogene Basin. Key words: Bacillariophyta, Cymbella serdica sp. nov., Sofia Neogene Basin, Bulgaria INTRODUCTION The Novi Iskar Formation consists of lacustrine sediments which were deposited in the Sofia Basin, South Bulgaria, in the Late Miocene – Early Pliocene, and contains very diverse diatom flora (Ognjanova-Rumenova 1991). The diatom flora includes 365 extinct and extant species, varieties and forms, which are significant from ecological and evolutional viewpoints. In previous studies of this material the biochronological scheme was based only on the development of the different planktonic genera of the Centrophyceae (Ognjanova-Rumenova and Popova 1992). In this scheme the most diverse group of the Pennatophyceae was neglected. Some of these pennatae species have a short stratigraphic range and even if benthic – they could be used as index species, for example the extinct species of the genera Fragilaria Lyngbye, Tetracyclus Ralfs, and Eunotia Ehrenberg. The objective of this study was to describe and illustrate a new species, of the genus Cymbella Ehr. from the Upper Miocene Novi Iskar Formation, Sofia Basin, and to determine its biostratigraphic significance.

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MATERIAL AND METHODS The sediments studied were collected from two boreholes C-1 and C-14, situated in the northern part of the Sofia Basin. The two boreholes cut across the Novi Iskar Formation (Kamenov and Kojumdgieva 1983). Borehole C-1, village of Goljanovci, 18 km northern of Sofia, 42o51’00'’ north and 23o13’48'’ east, is represented by irregular alternation of diatomaceous clays (0-48.8 m), lignite and lignite clays (48.8-57.1 m), chalk (57.1-67.4 m), lignite clays (67.4-76.7 m) and gray limestones (76.7-87.3 m). Borehole C-14, village of Katina, 15 km northern of Sofia, 42o50’01'’ north and 23o14’13'’ east, cuts across 10 m sands (not sampled), gray-bluish silty clays alternated by lignite, sands and sandy silty clays (10.0-110.0 m), gray clays with diatomacous clays (110.0-199.0 m), lignite with lignite clays (199.0-260.0 m), chalk (260.0-262.4). Diatom bearing sediments from boreholes C-1 (1.548.8 m) and C-14 (10.0-190.0 m) were investigated. (Fig. 1) The samples were prepared for diatom analyses following standard laboratory procedures (Ognjanova-Rumenova 1991), which combined parts of methods of Schrader (1973) and Hasle and Fryxell (1970). The relative abundance of the taxon was given according to Schrader’s scale (Schrader 1973). OBSERVATIONS AND DISCUSSION Cymbella serdica Metzeltin & Ognjanova-Rumenova nov. spec. (Pl. 1: Figs 1-3) Valvae valde dorsiventrales, semi-lanceolatae-ellipticae valvis maioribus fortiter lunatis, cum margine dorsali magis convexa et margine ventrali magis concava cum levi inflatione in medio, apicibus non protractis modice late rotundatis, 110-165 µm longae, 38-48 µm latae. Area axialis modice angusta, linearis. Area centralis fere parva. Raphe modice lateralis, ad aream centralem sensim filiformis. Stigmata nulla vel non bene aspectabilia. Striae transapicales modice radiatae, punctatae-lineolatae, 8-9 in 10 µm in parte dorsali, parte ventrali 6,5-7 in 10 µm, puncta 12 in 10 µm.

Typus: Praep. Core 1/20, 30.0 m depth in Coll. Ognjanova-Rumenova, Institute of Geology, Bulgarian Academy of Sciences, Sofia Locus typicus: 18 km in the north of Sofia, “Neogene Sofia basin”, near the village of Goljanovci; Borhole is 42o51'00'’ north and 23o13'48'’ east. Etymology: Serdica is the first recorded name of the modern town of Sofia. The Thracian Serdi tribe settled here in the 7th century BC and gave this name. Diagnosis difference in comparison with Cymbella aspera (Ehr.) Peragallo and C. neogena (Grunow) Krammer. (Krammer 2002) Valves much more strongly dorsiventral, crescent-shaped. Dorsal margin more convex than the ventral margin concave, the latter commonly slightly inflated in the middle. Ends bluntly rounded, not protracted. Length 110-165 µm, breadth 38-48 µm (not 26-35). Raphe 92

A new Cymbella from the Neogene in Bulgaria and its stratigraphic significance

Fig. 1. Biostratigraphic distribution of Cymbella serdica in the lacustrine sediments of Novi Iskar Formation, Sofia Neogene Basin, South Bulgaria.

93


1

2

3 10 µm

Plate 1. Figs 1-3 Cymbella serdica sp. nov. Scale bar 10 µm.

94

A new Cymbella from the Neogene in Bulgaria and its stratigraphic significance

moderately lateral, becoming filiform proximally, central ends somewhat far-standing and very distinctly deflected (not weakly) to the ventral side (without large, somewhat round central pores). Axial area moderately narrow; central area comparatively small rather than large. Stigmata absent or at least not discernible in LM. Striae moderately radial becoming subparallel near the ends, 8-9 in 10 µm at the ventral side, 6.5-7 at the dorsal side (nowhere more than 10/10 µm. Areolae 12 in 10 µm. Biostratigraphic distribution: Cymbella serdica has been found as ‘’rare’’ in samples: C-1, village of Goljanovci, 4,5m, 9.0m, 12.0m, 30.0m and 31.5m; C-14, village of Katina, 140.0145.0m, 183.0-190.0m. (Fig. 1) These levels come from diatom zone Concentrodiscus sp. (Ognjanova-Rumenova and Popova 1992), the Middle Pontian stage of Late Miocene age, corresponds to mammal zone MN-13 (Kojumdgieva et al. 1984, Spassov 2000), about 6 MA B.P. (Steininger and Wessely 2000). According to the hypothesis for the palaeoecological development of the Sofia Neogene basin – this was the first stage of the deposition of sediments of Novi Iskar Formation – an oligotrophic phase (Ognjanova-Rumenova and Popova 1996). ACKNOWLEDGEMENTS Nadja Ognjanova-Rumenova dedicates this paper to Prof. Dobrina Temniskova, who inspired me to study fossil diatoms and to thank her for our fruitful cooperation for many years. The authors acknowledge the help our colleague and friend, Prof. Dr. Dr. h. c. Horst Lange-Bertalot for the latin description of this new species, and also our colleague and friend Dr. Kurt Krammer for the numerous discussions of Cymbella species. REFERENCES HASLE, G. and G. FRYXELL 1970. Diatoms: Cleaning and mounting for light and electron microscopy. – Transaction of the American Microscopical Society, 89, 4: 469-474. KAMENOV, B. and E. KOJUMDGIEVA 1983. Stratigraphy of the Neogene in Sofia Basin. Palaeontology, stratigraphy and lithology. – Bulgarian Academy of Sciences, 18: 6985 (in Bulgarian). KOJUMDGIEVA, E., I. NIKOLOV and P. MEIN 1984. Les associations de grands mamifères du Miocene superieur en Bulgarie et leur correlation avec l’echelle regionale de la Paratethys. – Comptes rendus de l’Academie bulgare des Sciences, 37, 3: 341-343. KRAMMER, K. 2002. Diatoms of Europe. Diatoms of the European Inland Waters and Comparable Habitats. (ed. Lange-Bertalot, H.) Vol. 3. Cymbella. 584 p., 194 pl. A.R.G. Gantner Verlag K.G. FL 9491 Ruggell. Distributed by Koeltz Scientific Books, Koenigstein. OGNJANOVA-RUMENOVA, N. 1991. Neogene diatoms from sediments of Sofia Valley and their stratigraphic significance. Ph.D. Thesis, Geological Institute, Bulgarian Academy of Sciences, 330 pp. (in Bulgarian).

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OGNJANOVA-RUMENOVA, N. and E. POPOVA 1992. Diatom biostratigraphy and comparative core correlation within Sofia basin, Bulgaria. in: J. Eder-Kovar, editor. Palaeovegetational development in Europe and regions relevant to its palaeofloristic evolution, Proceedings of the 1st Pan-European Palaeobotanical Conference, Vienna, 1991: 197-203. OGNJANOVA-RUMENOVA, N. and E. POPOVA 1996. Palaeolimnological reconstruction of the Sofia Neogene Basin, Southern Bulgaria – a review of palaeoenvironmental diatom studies. – Phytologia Balcanica, 2, 2:43-53. SCHRADER, H-J. 1973. Proposal for a standardized method of cleaning diatom bearing deep-sea and land-exposes marine sediments. – Beihefte zur Nova Hedwigia, 45: 403-409. SPASSOV, N. 2000. The Turolian Hipparion-fauna and the character of the environment in the Late Miocene of West Bulgaria. – Review of the Bulgarian Geological Society, 61, 1-3: 47-61. STEININGER, F. and G. WESSELY 2000. From the Tethian ocean to the Paratethys sea: Oligocene to Neogene stratigraphy, palaeogeography and palaeobiogeography of the circum Mediterranean region and the Oligocene to Neogene Basin evolution in Austria. Aspects of geology in Austria. – Mitt. Österr. Geol. Ges., 92 (1999): 95-116.

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Nitzschia toskalensis sp. nov. a new diatom (Bacillariophyceae) from the sediments of Toskaljavri, northwestern Finland Paul B. Hamilton1 & Jan Weckström2 Phycology Section, Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada, e-mail: [email protected] 2 Environmental Change Research Unit (ECRU), Department of Biological and Environmental Sciences, P.O. Box 65 (Viikinkaari 1), FIN-00014, University of Helsinki, Finland, e-mail: [email protected] 1

ABSTRACT Nitzschia toskalensis Hamilton & Weckström sp. nov. is identified and described from the Holocene sediments of Lake Toskal, northwestern Finland. The new species was observed sporadically in the sediments from present to 6500 cal. yr BP. Nitzschia toskalensis is a rare taxon (

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