Freshwater Benthic Diatoms of Central Europe: Over ...

Editors:

Marco CANTONATI, Martyn G. KELLY & Horst LANGE-BERTALOT

Freshwater Benthic Diatoms of Central Europe: Over 800 Common Species Used in Ecological Assessment English edition with updated taxonomy and added species [German edition: Hofmann G., Werum M. & Lange-Bertalot H. 2013. Diatomeen im Süßwasser-Benthos von Mitteleuropa. Bestimmungsflora Kieselalgen für die ökologische Praxis. Über 700 der häufigsten Arten und ihre Ökologie. Lange-Bertalot, H. (Ed.): 908 pp. 2. korrigierte Aufl.]

with 3578 Figures on 135 Plates

Authors: Horst Lange-Bertalot, Gabriele Hofmann, Marcus Werum & Marco Cantonati

Addresses of Authors and Editors: PD Dr Marco Cantonati Museo delle Scienze – MUSE, Limnology and Phycology Section Corso del Lavoro e della Scienza 3 I-38123 Trento - Italy

E-Mail: [email protected] Current address: Academy of Natural Sciences of Drexel University, Patrick Center for Environmental Research, Phycology Section, Philadelphia, PA, USA.

Dr Gabriele Hofmann Hirtenstraße 19 61479 Glashütten

E-Mail: [email protected] Dr Martyn G. Kelly Bowburn Consultancy Durham - UK E-Mail: [email protected]

Prof. Dr Dr h.c. Horst Lange-Bertalot Fb. Biowissenschaften the Goethe-Universität, Frankfurt am Main Institut for ecology, Evolution, Diversität and Forschungsinstitut Senckenberg Privat: Silberweg 3, 61350 Bad Homburg - Germany, Fax: +49 6172 301197

Dr Marcus Werum Karl-Bieber-Höhe 23 60437 Frankfurt am Main E-Mail: [email protected]

The Project for the German Edition was funded by the programme O 5.06/2006 with resources from the state (Länder) funding programme “Water, Soil, and Waste”.

Table of contents Foreword to the German Edition ....................................................................................................................7 Foreword to the English Edition .....................................................................................................................9 Acknowledgments ........................................................................................................................................11 Introduction ..................................................................................................................................................13 How to use this book ....................................................................................................................................15 Identification characters ...............................................................................................................................17 Glossary ........................................................................................................................................................27 Key to the genera ..........................................................................................................................................39 Key to the diatom genera covered by this book .........................................................................................40 Genera and species .......................................................................................................................................77 Achnanthes Bory 1822 ...............................................................................................................................77 Achnanthidium Kützing 1844 .....................................................................................................................79 Adlafia Lange-Bertalot in Moser et al. 1998 ...............................................................................................94 Amphipleura Kützing 1844 .........................................................................................................................97 Amphora Ehrenberg ex Kützing 1844 .........................................................................................................98 Aneumastus D.G. Mann et A.J. Stickle in Round et al. 1990 .....................................................................104 Anomoeoneis Pfitzer 1871 .......................................................................................................................107 Astartiella Witkowski, Lange-Bertalot et Metzeltin in Moser et al. 1998 .................................................108 Bacillaria Gmelin 1791 .............................................................................................................................109 Berkeleya Greville 1827 ............................................................................................................................110 Biremis D.G. Mann et E.J. Cox 1990 ..........................................................................................................111 Brachysira Kützing 1836 ...........................................................................................................................113 Caloneis P.T. Cleve 1894 ............................................................................................................................120 Campylodiscus Ehrenberg 1844 ...............................................................................................................128 Cavinula D.G. Mann et A.J. Stickle in Round et al. 1990 ...........................................................................129 Chamaepinnularia Lange-Bertalot et Krammer in Lange-Bertalot & Metzeltin 1996 ...............................133 Cocconeis Ehrenberg 1837 .......................................................................................................................137 Cosmioneis D.G. Mann et A.J. Stickle in Round et al. 1990 .......................................................................143 Craticula Grunow 1868 ............................................................................................................................144 Crenotia A.Z. Wojtal 2013 ........................................................................................................................149 Ctenophora (Grunow) Williams et Round 1986 ........................................................................................151 Cylindrotheca Rabenhorst 1859 ...............................................................................................................152 Cymatopleura W. Smith 1851 ...................................................................................................................153 Cymbella C. Agardh 1830 .........................................................................................................................155 Cymbellafalsa Lange-Bertalot et Metzeltin 2009 .....................................................................................171 Cymbellonitzschia Hustedt in A. Schmidt et al. 1924 ................................................................................172 Cymbopleura (Krammer) Krammer 1999 .................................................................................................173 Delicata Krammer 2003 ...........................................................................................................................180 Denticula Kützing 1844 ............................................................................................................................182 Diadesmis Kützing 1844 ...........................................................................................................................184 Diatoma Bory 1824 ..................................................................................................................................185 Didymosphenia M. Schmidt 1899 .............................................................................................................188 Diploneis Ehrenberg 1844 ........................................................................................................................190

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Inhaltsverzeichnis Ellerbeckia R.M. Crawford 1988 ...............................................................................................................200 Encyonema Kützing 1833 .........................................................................................................................201 Encyonopsis Krammer 1997 .....................................................................................................................211 Entomoneis Ehrenberg 1845 ....................................................................................................................216 Epithemia Brébisson ex Kützing 1844 .......................................................................................................217 Eucocconeis P.T. Cleve ex F. Meister 1912 .................................................................................................222 Eunotia Ehrenberg 1837 ...........................................................................................................................225 Fallacia A.J. Stickle et D.G. Mann in Round et al. 1990 .............................................................................260 Fistulifera Lange-Bertalot 1997 ................................................................................................................265 Fragilaria Lyngbye 1819 ...........................................................................................................................267 Fragilariforma Williams et Round 1988 ...................................................................................................278 Frustulia Rabenhorst 1853 .......................................................................................................................281 Geissleria Lange-Bertalot et Metzeltin 1996 ............................................................................................285 Gliwiczia M. Kulikovskiy, Lange-Bertalot et A. Witkowski 2013 ................................................................288 Gomphocymbellopsis Krammer 2003 .......................................................................................................289 Gomphoneis Cleve 1894 ...........................................................................................................................290 Gomphonema Ehrenberg 1832 ................................................................................................................291 Gomphosphenia Lange-Bertalot 1995 ......................................................................................................325 Gyrosigma Hassall 1845 ...........................................................................................................................327 Halamphora (Cleve) Levkov 2009 .............................................................................................................331 Hannaea R.M. Patrick 1966 ......................................................................................................................336 Hantzschia Grunow 1877 .........................................................................................................................338 Hippodonta Lange-Bertalot, Metzeltin et Witkowski 1996 .......................................................................340 Humidophila Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot et Kopalová 2014 ........................344 Karayevia F.E. Round et L. Bukhtiyarova ex F. E. Round ............................................................................347 Khursevichia M.S. Kulikovskiy, Lange-Bertalot et Metzeltin 2012 .............................................................349 Kobayasiella Lange-Bertalot 1999 ............................................................................................................350 Kolbesia F.E. Round et L. Bukhtiyarova ex F.E. Round 1998 ......................................................................351 Lemnicola Round et Basson 1997 .............................................................................................................354 Luticola D.G. Mann in Round et al. 1990 ..................................................................................................355 Mastogloia Thwaites in W. Smith 1856 ....................................................................................................360 Mayamaea Lange-Bertalot 1997 ..............................................................................................................364 Melosira C. Agardh 1824 ..........................................................................................................................368 Meridion C. Agardh 1824 .........................................................................................................................370 Microcostatus Johansen et Sray 1998 ......................................................................................................372 Navicula Bory 1822 ..................................................................................................................................373 Navicymbula Krammer 2003 ....................................................................................................................417 Neidiomorpha Lange-Bertalot et Cantonati 2010 .....................................................................................418 Neidium Pfitzer 1871 ................................................................................................................................420 Nitzschia Hassall 1845 ..............................................................................................................................425 Nupela Vyverman et Compère 1991 ........................................................................................................466 Odontidium Kützing 1844 .........................................................................................................................468

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Inhaltsverzeichnis Orthoseira Thwaites 1848 ........................................................................................................................470 Paraplaconeis M.S. Kulikovskiy, Lange-Bertlot et Metzeltin 2012 ............................................................471 Parlibellus E.J. Cox 1988 ...........................................................................................................................473 Peronia Brébisson et Arnott ex Kitton 1868 .............................................................................................475 Pinnularia Ehrenberg 1843 .......................................................................................................................476 Placoneis Mereschkowsky 1903 ...............................................................................................................495 Planothidium Round et Bukhtiyarova 1996 ..............................................................................................504 Platessa Lange-Bertalot 2004 ...................................................................................................................512 Prestauroneis K. Bruder et Medlin 2008 ..................................................................................................517 Psammothidium Bukhtiyarova et Round 1996 .........................................................................................518 Pseudofallacia Liu, Kociolek et Wang 2012 ..............................................................................................528 Pseudostaurosira Williams et Round 1988 ...............................................................................................530 Reimeria Kociolek et Stoermer 1987 ........................................................................................................533 Rhoicosphenia Grunow 1860 ...................................................................................................................535 Rhopalodia O. Müller 1895 ......................................................................................................................536 Rossithidium Bukhtiyarova et Round 1996 ...............................................................................................538 Sellaphora Mereschkowsky 1902 .............................................................................................................541 Simonsenia Lange-Bertalot 1979 ..............................................................................................................557 Skabitschewskia Kulikovskiy et Lange-Bertalot 2015 ................................................................................558 Stauroforma Flower, Jones et Round 1996 ...............................................................................................560 Stauroneis Ehrenberg 1843 ......................................................................................................................561 Stauronella Mereschkowsky 1901 ............................................................................................................570 Staurosira Ehrenberg 1842 .......................................................................................................................571 Staurosirella Williams et Round 1988 ......................................................................................................574 Stenopterobia Brébisson ex Van Heurck 1896 ..........................................................................................578 Surirella Turpin 1828 ................................................................................................................................579 Tabellaria Ehrenberg ex Kützing 1844 ......................................................................................................589 Tabularia (Kützing) D.M. Williams et Round 1986 ....................................................................................591 Tetracyclus Ralfs 1843 ..............................................................................................................................593 Tryblionella W. Smith 1853 .......................................................................................................................594 Ulnaria (Kützing) P. Compère 2001 ...........................................................................................................601 Selected brackish-water taxa found along the northern Germany coastline ............................................604 References ..................................................................................................................................................609 Plates ..........................................................................................................................................................627 Index to the species ....................................................................................................................................901

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Foreword to the German Edition

According to the EU Water Framework Directive (EU-WFD) different biological components have to be considered in order to assess running and standing freshwater habitats. In this context, benthic diatoms, along with macrophytes, other phytobenthos organisms, and the phytoplankton, represent a very important group of photoautotrophic organisms. Diatoms are proven indicators of the quality of freshwater environments and are successfully used for the assessment of organic pollution and trophic status, as well as of anthropogenic acidification and salinization. The EU-WFD requires, in the first instance, an assessment of the overall ecological quality, that offers a comprehensive representation of the situation of freshwater environments by means of the integration of the relevant influencing factors. Each of the Member States of the European Union developed their own national procedures for its implementation. In the German evaluation procedures for aquatic vegetation, PHYLIB (SCHAUMBURG et al. 2006, 2007, 2008), use of the 'diatoms' modules requires a comprehensive knowledge of benthic diatoms due to their high species richness. The application of the procedure is further complicated by the continuous increase of knowledge on the species and their ecology in the past years not least as a consequence of the enhanced use of benthic diatoms as bioindicators in flowing and standing freshwater habitats. This has led to a strong differentiation of the identification literature. The Süßwasserflora of central Europe volumes 2/1-4 (KRAMMER & LANGE-BERTALOT 1986-1991) provided an international standard of the situation until the mid 1990s; however, since that time the identification literature has increased due to many new descriptions and revisions, and has become complex and often difficult to access. With the present text, developed on behalf of the German Working Group of the Federal States on Water Issues (LAWA), an identification book covering over 700 diatom species relevant for the assessment of freshwater habitats is provided for the first time. Identification characters, ecology, and similar taxa are documented along with species profiles and micrographs. In this way, almost half of all benthic taxa recorded in central Europe were covered. Existing uncertainties were removed, to some extent, by means of new descriptions. Where necessary, problems related to genus affiliation were explained, but without uncritically following every new proposal in the literature. As an additional aid for identification, an overview of the available records of the species in German freshwater habitat types is provided.

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Foreword to the English Edition

After the time known informally by diatomists as the “Era of the Green Books” (i.e. the Süßwasserflora) which lasted about until the mid 1990s and during which all the information one needed to have to address the central European diatom microflora was included in four books, there was an explosion of new publications as fundamental concepts underpinning the taxonomy of diatoms were revised, a large number of new species were described, and many familiar taxa bounced between genera. These are exciting times for those interested in the phylogeny, biogeography and diversity of diatoms, but also hugely frustrating times for those concerned with the routine identification of diatoms for practical purposes. Application of the EUWFD needs a consistent approach to the naming of the organisms upon which assessments depend. HOFMANN et al. (2011) represented a welcome exception to the trend of an ever-more diffuse literature on diatoms, since it gave analysts, once again, a single identification text for the species most likely to be found during routine assessments in central Europe. In order to facilitate use outside the German-speaking area, we decided to translate this work into English. At the same time, the lead Editor used this opportunity to update the work to include the latest information on the taxonomy and ecology. Decisions on revisions were made following a very empirical criterion: all taxonomy and ecology papers of which we are aware published until autumn 2016 were considered, and all genera and species (if quantitatively relevant) were included if they appeared to be recognized in the main online resources (in particular, ‘Diatoms of the Unites States’, ‘AlgaeBase’ etc.), and by the community of diatomists. This is a very practical criterion, which might sometimes lead to inclusion of taxa whose merits are still being debated, as long as it appears that they still might have some useful application in everyday practice and some recognition in applied ecology. Following this process, 26 genera were included in the updated English edition of the book whilst in just three cases genera that were included in the first (German) edition have been removed (although they are still mentioned and briefly explained in the book). 10 species have been added to the book, 27 more have been newly mentioned under ‘Similar taxa’, taxonomic concepts were clarified / updated for 36, and 39 were transferred from one genus to another.

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The clarification and updating made it necessary to produce new identification keys for two genera (Diploneis and Sellaphora). Information on plastids (only occasionally available in the German edition) has been added whenever possible to the descriptions of the genera. Terminology of both taxonomic features and also ecological concepts was updated in the Glossary. We have also improved the information on ecology and distribution for 22 species. These improvements to the text are accompanied by several additional references. Formal taxonomic changes were carried out only when strictly necessary in order to simplify difficult situations or improve the rationale of genus affiliation: one new species description (of a morphologically distinct Sellaphora of waters of high saprobic level), seven formal transfers (comb. nov.), one formal name changes (nom. nov.). The full text of the species descriptions of the German edition has been replaced by more schematic profiles, in which the different points are provided in a “bullet point” style. This replaces the rich German prose, capable of conveying even nuances of the morphology, with something far more accessible to international audiences. This English edition, like the earlier German ones, is intended primarily as a tool to facilitate consistent and accurate identification of diatoms in central Europe. We are, however, confident that it might be of general interest for all those looking for an up-to-date overview of most freshwater benthic diatom genera and of the characteristics of their most commonly encountered species.

Marco Cantonati & Martyn Kelly

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Acknowledgments

German Edition: Our sincere thanks go to all the institutions and colleagues, who made this book possible supporting us in many ways: the German Working Group of the Federal States on Water Issues (LAWA), that funded and supervised the initiative (Eva Bellack, Jochen Schaumburg, Klaus Wendling), the competent authorities and the employees of the Federal Länder, Andreas Hoppe (Baden-Württemberg), Jörg Schönfelder (Brandenburg), Marina Carstens (MecklenburgVorpommern), Eva Abée (Lower Saxony), Ilona Arndt-Dietrich and Karl-Heinz Christmann (Nordrhein-Westfalen), Susanne Wanner (Rheinland-Pfalz), Angela Doege, Jens Kroker, Markus Paul and Andreas Meybohm (Saxony), Martina Jährling (Saxony-Anhalt), Mandy Bahnwart (Schleswig-Holstein) and Kerstin Wyrwa (Thüringen), who provided additional diatom data. Special thanks go to our “diatom Colleagues”, who supported us with their expertise in the work up of problematic genera in a friendly and uncomplicated way: the late Frithjof Sterrenburg (Gyrosigma), Erwin Reichardt (Gomphonema), Ditmar Metzeltin and Ilka Schönfelder (Pinnularia). We are grateful to Kurt Krammer, Zlatko Levkov, Ditmar Metzeltin, Erwin Reichardt, and the late Frithjof Sterrenburg for permission to use micrographs. Heartfelt thanks go also to Silvia Hofmann, who examined the manuscript in such an in depth and meticulous way.

English Edition: We are grateful to Per & Sven Koeltz (Koeltz Scientific Books) for encouraging, funding, and supporting the production of an updated English edition of this book. MC is grateful to Don Charles (Leader of the Phycology Section, Patrick Center for Environmental Research, Academy of Natural Sciences of Drexel University, Philadelphia, PA, U.S.A.) for continuous encouragement to accomplish this task. For micrographs (or comments on specific parts) thanks go to: Małgorzata Bąk (Univ. Szczecin, Poland), Luc Ector (LIST, Luxembourg) and 11

Valérie Peeters (DIREN Bourgogne), Ionel Ciugulea (ANSP Drexel, PA, U.S.A.), Andre Fuhrmann (Univ. Frankfurt, Germany), Regine Jahn & Jonas Zimmermann (BGBM Univ. Berlin, Germany), Ingrid Jüttner (National Museum of Wales, UK), Bryan Kennedy (Environment Protection Agency, Ireland), Lydia King (Freiburg, Germany), Nicola Angeli & Chiara Defrancesco (Limnology & Phycology Sect., MUSE Trento), Zlatko Levkov (Univ. Skopje, Macedonia), Ditmar Metzeltin (Hofheim, Germany), Erwin Reichardt (Treuchtlingen, Germany), Bart Van de Vijver (Botanic Garden Meise, Belgium), Carlos Wetzel (LIST, Luxembourg).

We dedicate this book posthumously to Ute Rumrich, who inspired generations of students for the world of diatoms.

„... Sonne, Bäume, Blumen, Wasser and Liebe. Freilich fehlt letztere im Herzen des Beschauers, so mag das Ganze wohl einen schlechten Anblick gewähren, und die Sonne hat dann bloß noch so and soviel Meilen im Durchmesser, und die Bäume sind gut zum Einheizen, und die Blumen werden nach Staubfäden klassifiziert, and das Wasser ist naß.“ Heinrich Heine (Die Harzreise, 1826) [... Sun, trees, flowers, water, and love. However, if the latter is missing from the heart of the observer, the whole might indeed provide a poor view, and the sun, as a result, only has a diameter of a certain number of miles, the trees are only good for firewood, the flowers are characterised only in terms of stamens and filaments, and the water is wet.] Heinrich Heine (Die Harzreise, 1826)

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