Willdenowia Annals of the Botanic Garden and Botanical Museum Berlin-Dahlem
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Willdenowia Annals of the Botanic Garden and Botanical Museum Berlin-Dahlem JOACHIM W. KADEREIT1*, DIRK C. ALBACH2, FRIEDRICH EHRENDORFER3, MERCÈ GALBANY-CASALS4, NÚRIA GARCIA-JACAS5, BERIT GEHRKE1, GUDRUN KADEREIT6,1, NORBERT KILIAN7, JOHANNES T. KLEIN1, MARCUS A. KOCH8, MATTHIAS KROPF9, CHRISTOPH OBERPRIELER10, MICHAEL D. PIRIE1,11, CHRISTIANE M. RITZ12, MARTIN RÖSER13, KRZYSZTOF SPALIK14, ALFONSO SUSANNA5, MAXIMILIAN WEIGEND15, ERIK WELK16, KARSTEN WESCHE12,17, LI-BING ZHANG18 & MARKUS S. DILLENBERGER1 Which changes are needed to render all genera of the German lora monophyletic? Version of record irst published online on 24 March 2016 ahead of inclusion in April 2016 issue. Abstract: The use of DNA sequence data in plant systematics has brought us closer than ever to formulating well- founded hypotheses about phylogenetic relationships, and phylogenetic research keeps on revealing that plant genera as traditionally circumscribed often are not monophyletic. Here, we assess the monophyly of all genera of vascular plants found in Germany. Using a survey of the phylogenetic literature, we discuss which classiications would be consistent with the phylogenetic relationships found and could be followed, provided monophyly is accepted as the primary criterion for circumscribing taxa. We indicate whether and which names are available when changes in ge- neric assignment are made (but do not present a comprehensive review of the nomenclatural aspects of such names). Among the 840 genera examined, we identiied c. 140 where data quality is suiciently high to conclude that they are not monophyletic, and an additional c. 20 where monophyly is questionable but where data quality is not yet suicient to reach convincing conclusions. While it is still iercely debated how a phylogenetic tree should be trans- lated into a classiication, our results could serve as a guide to the likely consequences of systematic research for the taxonomy of the German lora and the loras of neighbouring countries. Key words: classiication, generic circumscription, German lora, monophyly, phylogeny Article history: Received 4 November 2015; peer-review completed 22 December 2015; received in revised form 10 January 2016; accepted for publication 14 January 2016. Citation: Kadereit J. W., Albach D. C., Ehrendorfer F., Galbany-Casals M., Garcia-Jacas N., Gehrke B., Kadereit G., Kilian N., Klein J. T., Koch M. A., Kropf M., Oberprieler C., Pirie M. D., Ritz C. M., Röser M., Spalik K., Susanna A., Weigend M., Welk E., Wesche K., Zhang L.-B. & Dillenberger M. S. 2016: Which changes are needed to render all genera of the German lora monophyletic? – Willdenowia 46: 39 – 91. doi: http://dx.doi.org/10.3372/wi.46.46105 1 Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universität Mainz, D-55099 Mainz, Germany; *e- mail: [email protected] (author for correspondence); [email protected]; [email protected]; [email protected]; [email protected] 2 Institute for biology and environmental sciences, Carl von Ossietzky-University Oldenburg, Carl von Ossietzky-Str. 9 – 11, D-26111 Oldenburg, Germany; e-mail: [email protected] 3 Department for Systematic and Evolutionary Botany, University of Vienna, A-1040 Wien, Rennweg 14, Austria; e-mail: friedrich. [email protected] 4 Departament de Biologia Animal, Biologia Vegetal i Ecologia, Facultat de Biociències, Universitat Autònoma de Barcelona, ES-08193 Bellaterra, Spain; e-mail: [email protected] 5 Botanic Institute of Barcelona (CSIC-ICUB), Pg. del Migdia s. n., ES-08038 Barcelona, Spain; e-mail: [email protected]. es; [email protected] 6 Institut für Allgemeine Botanik, Johannes Gutenberg-Universität Mainz, D-55099 Mainz, Germany; e-mail: clausing@uni-mainz. de 7 Botanischer Garten und Botanisches Museum Berlin, Freie Universität Berlin, Königin-Luise-Str. 6 – 8, D-14195 Berlin, Ger- many; e-mail: [email protected] 8 Centre for Organismal Studies, Biodiversity and Plant Systematics, Heidelberg University, D-69120 Heidelberg, Germany; e-mail: [email protected] 9 Institute for Integrative Nature Conservation Research, Universität für Bodenkultur Wien, Gregor Mendel-Str. 33, A-1180 Wien, Austria; e-mail: [email protected] (Addresses continued on next page) 40 Kadereit & al.: Rendering all genera of the German lora monophyletic Introduction ever, most plant systematists (hopefully) will agree that the use of DNA sequence data in plant systematics has “All a taxonomist’s decisions are subject to revision in brought us closer than ever to formulating solid hypoth- time” (Davis & Heywood 1973), and indeed even the eses about phylogenetic relationships, which could serve most cursory comparison of taxa at any rank through as basis for classiication. Perhaps ironically, exactly the history of plant systematics reveals that their circum- how to translate a phylogenetic tree into a classiication scription has changed again and again. For example, as has resulted in ierce debates. Probably the majority of regards Rothmaler’s “Exkursionslora von Deutsch- authors will argue that the branching pattern of a phy- land”, used as basis for the present paper, well over 40 logenetic tree should be the primary criterion for clas- genera have been subject to changes in circumscription siication and that only monophyletic taxa (consisting when comparing the 19th and 20th editions of the Grund- of ALL descendants of one common ancestor) should band (Jäger 2012). Major reasons for changes in taxon be accepted. Some nevertheless maintain that non- circumscription, as described and discussed in great de- monophyletic taxa should also be accepted in order for tail by e.g. Davis & Heywood (1973) and Stuessy (2009), classiication to depict not only patterns of phylogenetic include the discovery of new species, availability of new relationship, i.e. tree topology, but also degree of (phe- data (characters), new approaches in data analysis, and, notypic) divergence. (Most of these latter authors will often related to the preceding point, changes in concepts describe the taxa they think of as paraphyletic; however, of classiication. At any point in time an author suggest- as is evident from Fig. 1, assessments of taxa as either ing change of taxon circumscription will have believed paraphyletic or as polyphyletic based on tree topology to provide something “better”, where “better” had to be alone are alternative ways to read a phylogenetic tree.) evaluated against the aim and purpose of the classiica- It is not our aim here to summarize or add to that discus- tion. sion. For that, the reader is referred to a recent review Post-Linnaean plant systematists (and indeed some by Schmidt-Lebuhn (2012), a proponent for recognizing systematists before Linnaeus) increasingly aimed at only monophyletic taxa, and a response to that review producing a “natural system” in which a priori selec- by Stuessy & Hörandl (2014), opponents to that view. It tion of characters used for classiication was replaced by is also not the aim of this paper to provide a general re- the simultaneous evaluation of many characters (Davis view of changes of genus concepts through time, which & Heywood 1973; Stuessy 2009). With the publication have been reviewed and discussed repeatedly (e.g. Hum- of Darwin’s (1859) “Origin of Species”, introducing the phreys & Linder 2009 and references therein). concept of evolution, “natural” obtained a new meaning, Instead, our aims are: (1) Taking the generic circum- and “natural” taxa were interpreted as groups of com- scriptions of the 19th edition of the Grundband of Roth- mon ancestry. Although “Post-Darwinian systems have maler (Jäger 2005) as starting point, to identify genera difered little in content, though they have difered in ar- which are not or not unequivocally monophyletic. Such rangement, from those of the later pre-Darwinian taxono- conclusions are based on a thorough survey of the phy- mists” (Davis & Heywood 1973), “After Darwin, virtu- logenetic literature. We make an efort to assess the qual- ally all comprehensive systems of classiication of plants ity of published phylogenies in terms of taxon sampling, were avowedly phylogenetic” (Stuessy 2009). DNA regions analysed and support for relationships We are far from having DNA sequences of all spe- identiied. This sometimes results in the conclusion that cies, and probably even farther from resolving all rela- a given genus may or may not be monophyletic, but that tionships among species and higher level lineages. How- the data available are too preliminary for drawing taxo- (Addresses continued from previous page) 10 Evolutionary and Systematic Botany, Institute of Plant Sciences, University of Regensburg, Universitätsstr. 31, D-93053 Regens- burg, Germany; e-mail: [email protected] 11 Department of Biochemistry, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa; e-mail: mpirie@sun. ac.za 12 Department of Botany, Senckenberg Museum of Natural History Görlitz, PF 300 154, D-02806 Görlitz, Germany; e-mail: [email protected]; [email protected] 13 Institut für Biologie/Geobotanik und Botanischer Garten, Martin-Luther-Universität Halle-Wittenberg, Neuwerk 21, D-06108 Halle, Germany; e-mail: [email protected] 14 Department of Molecular Phylogenetics and Evolution, Faculty of Biology, University of Warsaw, 101 Żwirki i Wigury str., 02-089 Warsaw, Poland; e-mail: [email protected] 15 Nees-Institut für Biodiversität