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Fig. 1.

Lycopodiaceae (L.-B. Zhang)

Diphasiastrum was resolved as embedded withinLycopodium L. (Wikström & Kenrick 1997,2001). Thus recognition ofDiphasiastrum Holub as a separate genus would makeLycopodium paraphyletic, and Zhang & Iwatsuki (2013) suggested inclusion ofDiphasiastrum inLycopodium. However, this finding was based on plastidrbcL data and limited sampling only, and a final decision should await better sampling and use of additional DNA sequences.

Hymenophyllaceae (L.-B. Zhang)

A new classification of the family recognized only nine genera (Ebihara & al. 2006), andTrichomanes speciosum Willd. should now be known asVandenboschia speciosa (Willd.) G. Kunkel.Trichomanes L. in a new circumscription is a mainly neotropical genus with a few species in continental Africa, Madagascar and the Indian Ocean (Ebihara & al. 2006) and was resolved as sister toVandenboschia Copel. based on plastidrbcL data (Ebihara & al. 2007).

Aspleniaceae (L.-B. Zhang)

Plastid data resolved the family into two well-supported clades,Asplenium L. andHymenasplenium Hayata (van den Heede & al. 2003;Schneider & al. 2004), which have different chromosome base numbers as well as distinct root characters (Murakami 1995;Schneider 1996). All other small segregate genera are nested withinAsplenium (van den Heede & al. 2003;Schneider & al. 2004). Thus, synonymization ofCeterach Willd. andPhyllitis Hill withAsplenium is advocated (e.g.Smith & al. 2006;Lin & Vianne 2013). Consequently,P. scolopendrium (L.) Newman should beA. scolopendrium L. andC. officinarum Willd. should beA. ceterach L.

Thelypteridaceae (L.-B. Zhang)

Lastrea Bory was resolved as part ofOreopteris Holub based on plastid markers (He & Zhang 2012), andL. limbosperma (All.) Ching should now be known asO. limbosperma (All.) Holub.

Hydrocharitaceae (J. W. Kadereit)

A phylogenetic analysis of the family based on nuclear, plastid and mitochondrial DNA sequences (Chen & al. 2012) provides some evidence thatEgeria Planch, may not be monophyletic whenElodea Michx. is treated as a distinct genus. As only two of five species ofElodea were included in that study, and support in the relevant part of the tree is not entirely convincing, treatment of the two genera as separate is acceptable for the time being. If combined, as has been done in the past (for discussion seeLes & al. 2006),Elodea would be the name to be used.

Zosteraceae (J. W. Kadereit)

The finding thatHeterozostera tasmanica (M. Martens ex Asch.) Hartog is deeply nested inZostera L. (Les & al. 1997;Les & al. 2002;Kato & al. 2003;Tanaka & al. 2003) opens the option to maintainZostera includingHeterozostera (Setch.) Hartog as one genus, or to divide this group into two or three genera. In both latter options Z.marina L. would remain inZostera and Z.noltii Hornem. would have to be combined inNanozostera Toml. & Posl. asN. noltii (Hornem.) Toml. & Posl. Subdivision into three genera has been advocated and justified with morphological distinctness in inflorescence and vegetative characters by Tomlinson & Posluzny (2001), and maintainance ofZostera as one genus has been recommended by Les & al. (2002).

Potamogetonaceae (J. W. Kadereit)

It has been shown that species ofPotamogeton subg.Coleogeton (Rchb.) Raunk. constitute a monophyletic lineage which is well-supported sister to the remainder ofPotamogeton L. (Lindqvist & al. 2006). As this lineage is morphologically well characterized, as well summarized by Preston (2005; but see alsoWiegleb & Kaplan 1998), it could be separated at generic rank asStuckenia Börner, as argued by Lindqvist & al. (2006) and other authors (Les & Haynes 1996;Holub 1997;Haynes & al. 1998;Kaplan 2008), or could be maintained withinPotamogeton as argued by Wiegleb & Kaplan (1998). If treated as a distinct genus,P. pectinatus L. should be known asS. pectinata (L.) Börner andP. filiformis Pers. asS. filiformis (Pers.) Börner.

Dioscoreaceae (J. W. Kadereit)

Tamus L. is clearly nested inDioscorea L. (Caddick & al. 2002a,2002b;Wilkin & al. 2005). In consequence,T. communis L. should be treated asD. communis (L.) Caddick & Wilkin. An alternative option, to splitDioscorea into many smaller genera, as suggested by Huber (1998), was discussed but rejected by Caddick & al. (2002b).

Liliaceae (J. W. Kadereit)

According to studies by Peterson & al. (2008; see alsoPeterson & al. 2004) and Zarrei & al. (2009), all based on a broad species sample and both nuclear and plastid sequences, a non-monophyleticLloydia Rchb. is nested inGagea Salisb. IfLloydia should be included inGagea, as suggested by Peruzzi & al. (2008) and Zarrei & al. (2011), a name forL. serotina (L.) Rchb. inGagea would be available(G. serotina (L.) Ker Gawl.).

Orchidaceae (M. Kropf)

Initiated by molecular phylogenetic studies by Pridgeon & al. (1997) and Bateman & al. (1997), European orchids, and especially the genusOrchis L. s.l., have become a prime example for recent rearrangements in generic delimitations (Stace 2010). Although subsequent phylogenetic studies (cf.Cozzolino & al. 1998,2001;Aceto & al. 1999;Bateman 2001;Pridgeon & al. 2001;Bateman & al. 2003) generated support for (most of) these rearrangements (but almost exclusively based on ITS sequence variation only), most remained subject to fierce debates in the orchid community (cf.Wucherpfennig 1999,2002,2005;Bateman 2001,2009,2012a,2012b;Buttler 2001;Devos & al. 2006;Kretzschmar & al. 2007;Tyteca & Klein 2008,2009;Scopece & al. 2010;Paulus 2012;Tyteca & al. 2012). Possible and partially implemented rearrangements (cf.Jäger 2012) include either splitting of polyphyletic genera into smaller genera (e.g.Orchis s.l.), or inclusion of genera, either with several species (e.g.Nigritella Rich. inGymnadenia R. Br.) or monospecific (e.g.Aceras anthropophorum (L.) R. Br. inOrchis s.str.), in (otherwise) paraphyletic genera in order to obtain monophyletic entities.

Phylogenetic studies placed the (previously) monospecificAceras anthropophorum close toOrchis italica Poir. (Pridgeon & al. 1997;Bateman & al. 2003). This close relationship at the base of theOrchis s.str. group (see below) was not only supported by the original ITS sequence data, but also by seed ornamentation patterns (Gamarra & al. 2012), hybridization patterns (Klein 1989,2004;Scopece & al. 2007), and the nuclearOrcLFY,OrcPI, OrcP2 loci (Montieri & al. 2004;Cantone & al. 2009,2011), although support by the mitochondrialcoxl marker (Inda & al. 2010a) and chloroplastrpl16 intron data (Inda & al. 2012) was ambiguous due to low resolution. Therefore, Bateman (2012a: 111–114) noted that the “most obviously problematic taxa are the readily recognized anthropomorphic speciesOrchis (Aceras)anthropophora (L.) All. andO. italica … shown as the two earliest-diverging species, making the anthropomorphic species paraphyletic relative to a monophyletic nonanthropomorphic group”. Given the absence of a final solution to the question which taxon is indeed basally branching inOrchis s.str. (i.e. a sister group relationship betweenA. anthropophorum andOrchis s.str. is still possible; seePridgeon & al. 1997;Bateman & al. 2003; see alsoJacquemyn & al. 2011), and the still debated future treatment ofOrchis s.str. in general (see below), one could also retainA. anthropophorum as the only species ofAceras and the onlyOrchis-like species without a spur. On the other hand, the inclusion ofAceras inOrchis s.str. is one of the most widely accepted changes of controversial generic circumscriptions in European orchids (seeBateman 2009: Tab. 1).

ITS phylogenies implied inclusion ofCoeloglossumviride (L.) Hartm. in an otherwise paraphyleticDactylorhiza Necker ex Nevski (asDactylorhiza viridis (L.) R. M. Bateman & al.; seeBateman & al. 1997,2003;Pillon & al. 2007). Further molecular markers, especially chalcone synthase variation (Inda & al. 2010b; see alsoInda & al. 2010a,2012), supported this inclusion becauseC. viride was found nested inDactylorhiza. However, evidence against its inclusion exists, and a combined ITS and ETS phylogenetic tree resolvedC. viride as sister toDactylorhiza (Devos & al. 2006). The latter authors also compiled morphological characters differing between the two groups (Devos & al. 2006: Table 1; see alsoWucherpfennig 1999). Most strikingly,C. viride has a nectariferous spur (van der Pijl & Dodson 1966), whereasDactylorhiza has food-deceptive flowers. AsCoeloglossum Hartm. is the earlier name, a proposal to conserveDactylorhiza overCoeloglossum was needed (Cribb & Chase 2001).

As a consequence of studies uncovering the (morphological) heterogeneity and the phylogenetic intermingling of different infrageneric species groups of the closely related generaLiparis Rich. andMalaxis Sol. ex Sw., the monospecificHammarbya paludosa (L.) Kuntze, certainly a close relative of these two genera (although not sampled in the respective studies; e.g.Cameron 2005), “has often been included in a broadly defined genusMalaxis” (Pridgeon & al. 2005: 464–465). There are a number of unique features characterizingH. paludosa (e.g. incumbent anthers (Szlachetko & Margońska 2002), vegetative reproduction by bulbils at the leaf margin), which, however, have been doubted to be sufficient for differentiation at the generic level given the high variation inMalaxis s.l. (Wucherpfennig 2005). Independently, and referring to recent (but still unpublished) phylogenetic analyses by G. A. Salazar, Pridgeon & al. (2005: 464–465) stated thatH. paludosa “does not lie in the mainMalaxis clade (Salazar, pers. comm.) but rather is sister to a large clade that includes bothMalaxis s.str. andLiparis s.str.” Until comprehensive phylogenetic evidence on relationships amongMalaxis andLiparis becomes available (see below),H. paludosa presently can be maintained in a monospecific genus.

The monospecificChamorchis alpina (L.) Rich. and the dispecificTraunsteinera Rchb., represented by the widespreadT. globosa (L.) Rchb. in Germany, form an independent, well-supported clade (Cozzolino & al. 2001;Bateman & al. 2003). This surprising result refutes the originally hypothesized sister group relationship between the latter taxon andOrchis s.str. (Pridgeon & al. 1997) and induced Pridgeon & al. (2005: 228) to state: “However, the two morphologically distinct genera are sufficiently similar in ITS sequences to be potentially viewed as congeneric”. If treated as congeneric, Chamorchis represents the older genus name (cf.Alrich & Higgins 2011).

In (still unpublished) molecular phylogenetic analyses by Bateman and colleagues,Neottia nidus-avis (L.) Rich. is nested within a paraphyleticListera R. Br. as sister toL. ovata (L.) R. Br. (illustrated byPridgeon & al. 2005: 492). Already Chase & al. (2003) had consideredListera species as photosynthetic members ofNeottia Guett. (without presenting the respective phylogenetic analysis, except for a “summary”, i.e.Fig. 1 on p. 73) and suggested that the two genera should be combined (see alsoTesitelová & al. 2015, whereNeottia (n = 2) is nested withinListera (n = 3) based on ITS, 18S andtrnL^(UAA) intron data). A close relationship between the two genera has long been documented (e.g.Dressler 1990), and a combined generic treatment asNeottia, which is the older name, has already been published by Szlachetko (1995). In this treatment, however,Neottia ovata Bluff & Fingerh. is placed inN. subg.Listera (R. Br.) Szlach. The third species of a newly circumscribedNeottia native in Germany isN. cordata (L.) Rich. (L. cordata (L.) R. Br.).

Pridgeon & al. (1997) “took the controversial step of sinking the morphologically distinctNigritella back into synonymy withGymnadenia s.str., which would otherwise have been paraphyletic.” (Pridgeon & al. 2001: 229). These authors stressed that “despite superficial differences in flower form and resupination,Nigritella shares several morphological characters withGymnadenia: palmatedigitate tubers; two lateral, lobe-like stigmas; and two pollinia each with a caudicle…” (Pridgeon & al. 2001: 298). However, other authors, especially Wucherpfennig (1999,2002), advocated maintainingNigritella as a genus based on at least ten (“superficial”) morphological characters, but also based on allozyme data (Hedrén & al. 2000). It was noted that a study of character evolution acrossOrchidinae clearly showed thatNigritella is a morphologically derived lineage (Wucherpfennig 2002) arguing for keeping the genusNigritella even within a paraphyleticGymnadenia. However, in a more recent molecular analysis of ITS andrpl16 intron sequences, Pillon & al. (2006) documented a sister group relationship between theirNigritella (n = 2) andGymnadenia (n = 5) samples. This illustrates that molecular phylogenetic relationships obtained obviously depend on taxon sampling, type of data analyses performed and outgroup selection (seePillon & al. 2007). In consequence,Nigritella can still be recognized as a morphologically well-defined genus, until more comprehensive analyses are available.

Finally, species ofOrchis s.l. were placed in at least three major and only distantly related groups based on ITS data (Bateman & al. 1997,2003;Pridgeon & al. 1997). These three groups in principle correspond to hybridization patterns (Klein 1989,2004;Scopece & al. 2007). As regards the first group, the formerly monospecificNeotinea Rchb. f. was expanded by Pridgeon & al. (1997) and Bateman & al. (1997) to encompass the “… small-flowered, essentially trilobed-lipped species of theustulata-group that were formerly included inOrchis s.l. These could in theory have been treated as a genus separate from the more narrowly delimited original concept ofNeotinea, given the relatively long molecular branch and distinct vegetative markings ofN. maculata…” (Pridgeon & al. 2001: 228). Relevant for the German flora, the combinationsN. ustulata (L.) R. M. Bateman & al. (O. ustulata L.) andN. tridentata (Scop.) R. M. Bateman & al. (O. tridentata Scop.) were provided (Bateman & al. 1997). However, the small flowers ofN.maculata (Desf.) Stearn are different from theustulata-group by producing nectar (Pridgeon & al. 2001;Duffy & al. 2009), and by being 100 % autogamous (Duffy & al. 2009), while the species of the deceptiveustulata-group depend on pollinatormediated outcrossing. This would provide arguments for treatingN. maculata as an independent genus. If this approach is taken, the namesOdontorchis ustulata (L.) D. Tyteca & E. Klein andOdontorchis tridentata (L.) D. Tyteca & E. Klein are available (Tyteca & Klein 2008).

The second fairly well-supported clade encompasses all species of formerOrchis that have 2n = 36 (or 2n = 32 in the case ofO. papilionacea L.) chromosomes as well asAnacamptis pyramidalis (L.) Rich. (Pridgeon & al. 1997). Pridgeon & al. (2001: 255) stated that while “A. pyramidalis is distinctive… The other members of this newly circumscribed genusAnacamptis Rich. are difficult to distinguish morphologically fromOrchis s.str., but their flowering stems bear cauline sheathing leaves.” Members ofAnacamptis in this new circumscription in the German flora areA. coriophora (L.) R. M. Bateman & al. (O. coriophora L.),A. morio (L.) R. M. Bateman & al. (O. morio L.) andA. palustris (Jacq.) R. M. Bateman & al. (O. palustris Jacq.).

The remainingOrchis s.l. taxa should then, following Pridgeon & al. (1997) and Bateman & al. (1997), be treated asOrchis s.str. comprising an anthropomorphic species group (with flowers shaped like “little men”, i.e. sepals and petals forming a compact head and the labellum showing “arms” and “legs”; e.g.O. militaris L., the type ofOrchis) plusAceras R. Br. (see above) and a non-anthropomorphic group (e.g.O. mascula (L.) L.). However, suggestions have been put forward to splitOrchis s.l. further (Tyteca & Klein 2008,2009), partly ignoring problems of paraphyly (criticized, e.g. byScopece & al. 2010;Bateman 2012a). However, the two supported species groups withinOrchis s.str. (Bateman & al. 2003) could be treated asO. subg.Orchis (i.e.O. militaris, O. purpurea Huds. andO. simia Lam.) andO. subg.Masculae H. Kretzschmar & al. (i.e.O. mascula, O. pallens L. andO. spitzelii Saut. ex W. D. J. Koch;Kretzschmar & al. 2007). Tyteca & al. (2012) compiled morphological and pollinator assemblage data for these two groups and concluded that all their information as well as molecular (Bateman & al. 2003) and seed micromorphology data (Gamarra & al. 2012) are in favour of a separation at the generic level, i.e. asOrchis andAndrorchis D. Tyteca & E. Klein (Tyteca & al. 2012; see alsoTyteca & Klein 2008 for respective names, i.e.Androrchis mascula (L.) D. Tyteca & E. Klein,A. pallens (L.) D. Tyteca & E. Klein andA. spitzelii (Saut. ex W. D. J. Koch) D. Tyteca & E. Klein).

Several orchid genera have been shown not to be monophyletic:Liparis andMalaxis, both comprising about 250 species (Mabberley 2008), are to some extent intermingled (Cameron 2005);Platanthera Rich. should includePiperia Rydb. (Bateman & al. 2003; already implemented there); andHerminium L. is phylogenetically intermingled withPeristylus Blume orHabenaria Willd. (Douzery & al. 1999;Bateman & al. 2003). However, irrespective of exact phylogenetic relationships, which are not yet completely resolved, the nomenclature of the species occurring in the German flora will not be influenced if their respective monophyletic clades are preserved at the generic level, asLiparis loeselii (L.) Rich.,Platanthera bifolia (L.) Rich. andHerminium monorchis (L.) R. Br. are the types of the respective genus names (Alrich & Higgins 2011), andPlatanthera montana (F. W. Schmidt) Rchb. f. (P. chlorantha Cust. ex Rchb.), the second native species of this genus, is definitely closely related to the type,P. bifolia (Bateman & al. 2003). However,Malaxis monophyllos (L.) Sw. might be affected by future changes: a BLAST search of recently publishedmatK barcodes of this species (Kim & al. 2014;Xiang & al. 2014) revealed higher DNA sequence similarity to a group ofLiparis species around the type,L. loeselii, than to theMalaxis species group around the type,M. spicata Sw. (cf.Cameron 2005). On the other hand, this critical point in the systematics ofMalaxideae could alternatively be solved by choosing a wide genus concept. In this case,Malaxis would be an older name thanLiparis (andHammarbya; see above).

In summary, one major problem with respect to several recently suggested changes in generic circumscription in European orchids is that new molecular phylogenetic hypotheses often are based on only one molecular marker (i.e. ITS;Bateman & al. 2003). Other molecular markers often resulted in limited phylogenetic resolution given the probably young age of several European orchid lineages (cf.Inda & al. 2010a,2010b,2012). Although sometimes combined evidence of ITS plus cpDNA variation seems to improve results (e.g.Pillon & al. 2006), it does not in other cases, indicating the dominance of the ITS information (e.g.Inda & al. 2012). Moreover, it is striking that the overall taxon sampling, some 20 years after the first molecular phylogenetic publications, is still incomplete. Also, multiple samples of single taxa have rarely been included. In consequence, many molecular phylogenetic relationships have still not been solved satisfactorily, and some nomenclatural changes accordingly are premature, giving rise to frequent debate.

Amaryllidaceae (J. W. Kadereit)

Using a broad sample ofGalanthus L. andLeucojum L., Lledó & al. (2004) reported that the former genus is deeply nested in the latter. In order to maintain these two genera, the authors recommend to recognize the additional genusAcis Salisb. for large parts ofLeucojum. Generic allocation ofG. nivalis L.,L. aestivum L. andL. vernum L. would remain unaffected if this approach would be taken.

Cyperaceae (B. Gehrke)

Carex L. has been found to be paraphyletic and to include all other members of theCariceae, i.e.Cymophyllus Mack.,Kobresia Willd.,Schoenoxiphium Nees andUncinia Pers. (Roalson & al. 2001; Starr & al. 2004). The results of the molecular phylogenetic work are unambiguous. RetainingKobresia would lead to the necessity of describing a myriad of morphologically indistinguishable smaller genera and would also mean thatKobresia would have to be either greatly extended to include many species ofCarex subg.Psyllophora (Degl.) Peterm. (=Primocarex Kük.) or thatKobresia (and Uncinia) would have to be split into various smaller lineages. Combination of all names ofCymophyllus, Kobresia, Schoenoxiphium andUncinia inCarex are currently underway (GlobalCarex Group 2015). The namesCarexmyosuroides Vill. forKobresia myosuroides (Vill.) Fiori andCarex simpliciuscula Wahlenb. forK. simpliciuscula (Wahlenb.) Mack. should be used.

Eleogiton (L.) Link was recently discovered to be nested inIsolepis R. Br. (Muasya & al. 2001).Isolepis was thought to be characterized by having one or more pseudolateral spikelets and an erect culm, but the nodding culm of the single terminal spikelet, believed to characterizeEleogiton, is now known to have evolved from withinIsolepis (Muasya & al. 2001).Eleogiton fluitans (L.) Link was therefore recently changed toI. fluitans (L.) R. Br.

Recent studies suggest thatSchoenoplectus mucronatus (L.) Palla andS. supinus (L.) Palla are not part ofSchoenoplectus (Rchb.) Palla, but belong toSchoenoplectiella Lye, a cosmopolitan group, which is most closely related to the AfricanPseudoschoenus (C. B. Clarke) Oteng-Yeb. (Shiels & al. 2014).Schoenoplectiella differs morphologically fromSchoenoplectus by having an unbranched inflorescence (Jung & Choi 2010), whereasSchoenoplectus has a pseudo-lateral branched inflorescence. Both genera have culm-like primary bracts opposed to the inflorescence with leafy bracts inScirpus L. (Jung & Choi 2010). If recognition ofSchoenoplectiella as suggested by Lye (2003) should be accepted, bothS. mucronatus andS. supinus must be excluded fromSchoenoplectus asSchoenoplectiella mucronata (L.) J. Jung & H. K. Choi andSchoenoplectiella supina (L.) Lye. However, final decisions must await a better understanding of relationships betweenPseudoschoenus andSchoenoplectiella.

Poaceae (M. Röser)

A number of molecular phylogenetic studies employing nuclear and chloroplast DNA markers have shown thatFestuca L. s.l. is a large paraphyletic group that encompassesLolium L.,Micropyrum (Gaudin) Link,Vulpia C. C. Gmelin and a number of further genera (Torrecilla & Catalán 2002;Catalán & al. 2004,2007;Torrecilla & al. 2004;Inda & al. 2008).Lolium is nested within a more ancestral broad-leaved clade, whereasMicropyrum andVulpia belong to the presumably more recently derived fine-leavedFestuca lineages.Vulpia additionally appears to be polyphyletic and encompasses separate diploid and tetraploid/hexaploid lineages, which are not sufficiently understood to date. Because of several uncertainties concerning limited sampling of intermediate taxa and missing representation of severalFestuca groups, Catalán & al. (2007) argued for maintenance ofLolium, Micropyrum andVulpia. This would require no name changes for taxa of the German flora.Micropyrum andVulpia were included inFestuca by Soreng & al. (2015), but Lolium was kept separate and considered congeneric withSchedonorus P. Beauv. (syn.F. subg.Schedonorus (P. Beauv.) Peterm.), which was segregated fromFestuca.

Polyploidy and hybridization play an important role in the evolution ofSesleria Scop. and allies. Preliminary data from Amplified Fragment Length Polymorphisms (AFLPs) and plastid DNA (trnL-ndhF) sequences support the recognition ofOreochloa Link as a separate genus (with onlyO. disticha (Wulfen) Link represented in the German flora) and underline thatPsilathera ovata (Hoppe) Deyl diverges from the remainder ofSesleria (Lakušić 2013). Further study including a more comprehensive taxon sampling is needed to clarify whether the monospecificPsilathera Link (onlyP. ovata (Hoppe) Deyl in the German flora) can be maintained or should be merged withSesleria as was done in Jäger (2011) and by Lazarević & al. (2015).

Delimitation of genera allied withHelictotrichon Besser ex Schult. & Schult. f. is a long-term matter of debate. Molecular phylogenetic studies using different chloroplast DNA and nuclear ITS sequences of a sufficiently broad sample of relevant taxa suggest to acknowledge three genera occurring in the German flora, namelyAvenula (Dumort.) Dumort.,Helictochloa Romero Zarco andHelictotrichon s.str. (Döring & al. 2007;Quintanar & al. 2007;Schneider & al. 2009; Röser & al. unpubl. data).Avenula is represented byA. pubescens (Huds.) Dumort.,Helictochloa byH. pratensis (L.) Romero Zarco andH.versicolor (Vill.) Romero Zarco andHelictotrichon s.str. byH. parlatorei (Woods) Pilg. The description of the new genusHelictochloa, type designations and transfer of species toHelictochloa have been published by Romero Zarco (2011).

The distinctiveness ofAnthoxanthum L. andHierochloe R. Br. has repeatedly been questioned due to the occurrence of seemingly intermediate species in Africa and SE Asia. Following Schouten & Veldkamp (1985), the two genera have been merged by some authors (Wu & Phillips 2006;Allred & Barkworth 2007;Kellogg 2015;Soreng & al. 2015). The study by Pimentel & al. (2013), using AFLPs, chloroplast and nuclear DNA sequences, suggests that the intermediate taxa originated by ancient hybridization between the two genera. The question as to whetherAnthoxanthum andHierochloe should be kept separate or amalgamated in a single genus thus remains unanswered.

Ranunculaceae (E. Welk)

TraditionallyAconitum L.,Consolida (DC.) Gray andDelphinium L. (andAconitella Spach, seeSoják 1969) were grouped in tribeDelphinieae. Molecular phylogenetic research revealed threeDelphinium species (D. subg.Staphisagria J. Hill) to form the sister clade to all otherDelphinieae (Jabbour & Renner 2011a;2011b), andConsolida incl.Aconitella to be nested in Delphinium excl.D. subg.Staphisagria. The position ofD. subg.Staphisagria is supported by biochemical, karyological and morphological characters. Furthermore, Wang & al. (2013) found a sister position of the ChineseAconitum gymnandrum Maxim. toDelphinium (sensuJabbour & Renner 2012). In order to arrive at monophyletic Aconitum andDelphinium, name changes are required. Of these, inclusion ofConsolida (andAconitella) into Delphinium (Jabbour & Renner 2012) is relevant for the German flora. In consequence,C. ajacis (L.) Schur,C.hispanica (Costa) Greuter & Burdet andC. regalis Gray should be listed asD. ajacis L.,D. hispanicum Costa andD. consolida L., respectively.

Based on molecular phylogenetic analyses, Bittkau & Comes (2009) foundGaridella L. to be clearly monophyletic whileNigella L., its sister group, was not well supported as monophyletic. This may imply future inclusion ofGaridella inNigella, which, however, would not affect naming of the German species ofNigella.

Combined analyses of DNA sequence data, biochemical data and morphology by Compton & al. (1998) suggested to includeCimicifuga Wernisch andSouliea Franch. inActaea L. (alsoCompton & Culham 2002;Gao & al. 2008). However, it has also been argued to keep the genera separate based on the fleshy fruits ofActaea (e.g.Wang & al. 1997;Lee & Park 2004).Actaea andCimicifuga can also be distinguished using seed morphology and seed anatomical features (Ghimire & al. 2015). IfCimicifuga andSouliea should be included inActaea based on phylogenetic relationships, GermanActaea will not be affected becauseActaea L. is the oldest genus name.

Hoot & al. (1994) suggested thatHepatica Mill.,Knowltonia Salisb. andPulsatilla Mill. should be included inAnemone L. s.l. (cf.Ehrendorfer & Samuel 2001;Schuettpelz & al. 2002). However, Pfosser & al. (2011) argued that these genera could also be retained because of unsuitable outgroup selection (Clematis L.) in Hoot & al. (1994) and Schuettpelz & al. (2002). UsingRanunculusficaria L. as outgroup in their study, a position ofClematis withinAnemone s.l. became probable. The sister-group relationship of species ofA. subg.Anemonidium (Spach) Juz. (A. subsect.Anemonidium Spach,A. subsect.Himalayicae (Ulbr.) Tamura,A. subsect.Keiskea Tamura andA. subsect.Omalocarpus (DC.) Tamura) toHepatica found in all studies rendersAnemone paraphyletic in relation to the embeddedHepatica andPulsatilla. Similar to combined karyological and molecular phylogenetic analyses by Mlinarec & al. (2012), Hoot & al. (2012) found, again withClematis as outgroup, thatA. subg.Anemonidium containsAnemonastrum Holub andHepatica, whilePulsatilla is positioned withinA. subg.Anemone. Accordingly, they suggested to incorporateHepatica inAnemone asA. sect.Hepatica (Mill.) Spreng. orA. subg.Hepatica (Mill.) Peterm. ForPulsatilla they suggested inclusion inAnemone asA. sect.Pulsatilloides DC. orA. subg.Pulsatilloides (DC.) Juz. An alternative solution might be splittingAnemone into at least two genera corresponding to thex = 7/8 divergence seen inAnemoninae. At the moment, it seems best to wait for further analyses before combining the large number of taxa affected. However, from the results of all studies cited it seems inevitable forAnemonastrum Holub to be subsumed inAnemone again. The resulting combination isAnemone narcissiflora L.

Caltha L. has been divided into two sections: the monophyleticC. sect.Psychrophila (DC.) Bercht. & J. Presl in the S hemisphere and the paraphyleticC. sect.Caltha in the N hemisphere (Schuettpelz & Hoot 2004). Based on a broader sampling, Cheng & Xie (2014) showed thatThacla Spach (Caltha natans Pall.) diverged first in the genus, and that the other species fall into two monophyletic clades, i.e.Caltha s. str. andPsychrophila. Thus, it would be possible to raisePsychrophila to genus rank, but this would inevitably requireC. natans to be raised toThacla. Any decision here will not affect the name ofC. palustris L.

A number of molecular phylogenetic studies revealed thatRanunculus L. in a wide sense is polyphyletic (Lehnebach & al. 2007;Hoot & al. 2008;Wang & al. 2009). Although the entire tribeRanunculeae could be recognized as a very broadly circumscribedRanunculus, this would result in a morphologically highly heterogeneous group. The morphological and geographical independence ofFicaria Huds. andCeratocephala Moench is comparable to that ofMyosurus L. It thus seems to be justified to follow Emadzade & al. (2010) who proposed to recognizeCeratocephala, Ficaria andMyosurus (plus several other small genera) as separate genera, but to includeBatrachium (DC.) Gray andAphanostemma A. St.-Hil., sometimes recognized as separate genera in the past, in a then monophyleticRanunculus.

Berberidaceae (J. W. Kadereit)

MonophyleticBerberis L. with simple leaves clearly is nested in a paraphyletic grade ofMahonia Nutt. with compound leaves (Kim & al. 2004;Adhikari & al. 2015), a pattern of relationship already postulated by Ahrendt (1961). As the two genera are very similar to each other in many respects (for discussion seeAdhikari & al. 2012), and the different lineages ofMahonia would be difficult to justify at generic rank, they probably are best treated as one genus,Berberis, as was done by these authors.Mahonia aquifolium (Pursh) Nutt. had originally been described asB. aquifolium Pursh.

Papaveraceae (J. W. Kadereit)

Papaver L. is part of a group of four genera distributed almost entirely in the Old World (Schwarzbach & Kadereit 1995). The other three genera areMeconopsis Vig.,Roemeria Medik, andStylomecon G. Taylor. Subdivision into these four genera is based largely on capsule morphology. Various analyses of these four genera (Kadereit & al. 1997;Carolan & al. 2006;Kadereit & al. 2011;Xiao 2013;Liu & al. 2014) revealed that patterns of relationship cut across traditional generic delimitations (see alsoFig. 1). First, three subgroups ofPapaver, i.e. (1)Papaver s.str. (all sections exceptP. sect.Argemonidium Spach,P. sect.Californica Kadereit,P. sect.Horrida Elkan andP. sect.Meconella Spach), (2)P. californicum A. Gray (P. sect.Californica) and (3)P. aculeatum Thunb. (P. sect.Horrida) form a clade together withMeconopsiscambrica (L.) Vig. andStylomecon heterophylla (Benth.) G. Taylor. Second,P. sect.Argemonidium is most closely related toRoemeria. Third,P. sect.Meconella Spach is most closely related to one of three subgroups ofMeconopsis. While this pattern of relationships allows several classifications, the following option has partly been followed (Kadereit & Baldwin 2011;Kadereit & al. 2011). A newly circumscribedPapaver should containMeconopsis cambrica, Papaver s.str.,P. aculeatum, P.californicum andStylomecon heterophylla. Of the species found in Germany,P. confine Jord.,P. dubium L.,P. lecoqii Lamotte andP. rhoeas L. would remain inPapaver. Meconopsis cambrica was originally described asP. cambricum L., and the nameP. heterophyllum (Benth.) Greene is available forStylomecon heterophylla. Papaver sect.Argemonidium, represented byP. argemone andP. hybridum in the German flora, should be united withRoemeria, with which it shares sepal and pollen characters (Kadereit & al. 1997). The combinationR. argemone (L.) C. Morales & al. is available forP. argemone.

AlthoughPapaver alpinum L. as part ofP. sect.Meconella should clearly be excluded fromPapaver, the exact relationships ofP. sect.Meconella to HimalayanMeconopsis are not sufficiently clear yet to suggest a formal name. However, it seems to be sister clade to a newly circumscribedMeconopsis (excl.Cathcartia Hook. f. andM. cambrica) and probably is best treated as a distinct genus.

Crassulaceae (J. T. Klein)

Sedum L. has repeatedly been shown to be highly polyphyletic (van Ham & al. 1994;van Ham & 't Hart 1998;Mort & al. 2001;Mayuzumi & Ohba 2004;Gontcharova & al. 2006;Carrillo-Reyes & al. 2009). In the most recent phylogenetic analysis ofCrassulaceae based on combined nuclear ITS and chloroplast DNA (Klein & Kadereit in prep.), the 20 species ofSedum found in Germany fall into several lineages.

The remaining species, including the type,Sedum acre L., fall into a large clade of the family that contains a large number of other genera (see below). If the species discussed above were to remain in a monophyleticSedum, essentially two thirds of the family would have to be included in that genus. Accordingly, segregation of three of the above four genera, i.e.Rhodiola, Phedimus andPetrosedum, is likely to be stable irrespective of future name changes in other parts of the family. As regardsHylotelephium, future name changes are conceivable because relationships between this genus and Meterostachys,Orostachys andSinocrassula are not yet fully resolved.

The large clade of the family containing the type consists of two subclades, known as the Leucosedum-clade and the Acre-clade (van Ham & 't Hart 1998), respectively.

The Leucosedum-clade, which also includesDudleya Britton & Rose,Mucizonia A. Berger,Pistorinia DC.,Prometheum (A. Berger) H. Ohba,Rosularia Stapf, Telmissa Fenzl andSedella Britton & Rose, contains seven species of GermanSedum, i.e.S. album L.,S. atratum L.,S. cepaea L.,S. dasyphyllum L.,S. hispanicum L.,S. rubens L. andS. villosum L. These seven species are scattered across a number of subclades, which partly contain one or more of the genera listed above.

The remaining five species of GermanSedum, i.e.S. acre, S. alpestre Vill.,S. annuum L.,S. sexangulare L. andS. sarmentosum Bunge fall into the Acre-clade, which also includesCremnophila Rose,Echeveria DC.,Graptopetalum Rose,Lenophyllum Rose,Pachyphytum Link, Klotsch & Otto,Thompsonella Britton & Rose andVilladia Rose. In this Acre-clade,S. acre is supported sister to all remaining taxa, andS. alpestre, S. annuum,S. sarmentosum andS. sexangulare again are scattered across a number of subclades.

In view of the relationships described above, several potential options exist for a monophyleticSedum. (1)Sedum could be treated as monospecific with only its type,S. acre. Of course, species that have not been sampled yet may fall into this clade. (2) The entire Acre-clade could be treated asSedum. This, however, would imply inclusion ofCremnophila, Echeveria, Graptopetalum,Lenophyllum, Pachyphytum, Thompsonella andVilladia.Sedum in such circumscription would contain c. 500 species. (3) The Acre-clade and the Leucosedum-clade (with c. 160 species) could be combined intoSedum, which would require additional inclusion ofDudleya, Mucizonia,Pistorinia, Prometheum, Rosularia, Telmissa andSedella.

Whereas recognition of a monospecificSedum (option 1) would require description of a large number of genera for former species of that genus, options 2 and 3 would require combination in one genus of morphologically very different genera that are geographically widely distributed. Of these three options, option 1 appears best to us, although the new genera that will have to be described partly may not be easy to differentiate morphologically or geographically. However, asSedum has not yet been completely sampled, and many relationships within the Acre- and Leucosedum-clades are not supported, I recommend to retain allSedum species of the Acre- and Leucosedum-clades inSedum until relationships are understood better. However, suchSedum clearly is not monophyletic.

Saxifragaceae (J. W. Kadereit)

The non-monophyly ofSaxifraga L., first shown by Soltis & al. (1993), has been confirmed in several studies (for discussion seeFernández Prieto & al. 2013;Tkach & al. 2015). In particular, a group of 70–90 species from North America and Eurasia is only very distantly related to the remainder ofSaxifraga and has to be treated as the genusMicranthes Haw. The one species affected in the German flora isSaxifraga stellaris L., which should be treated asM. stellaris (L.) Galasso & al. Following Soltis (2007), the two genera are clearly distinct in pollen and ovule characters.

Linaceae (J. W. Kadereit)

A broadly sampled phylogeny ofLinum L. and relatives by McDill & al. (2009) showed that the South AmericanCliococca Bab., the North AmericanHesperolinon (A. Gray) Small andSclerolinon C. M. Rogers and the EurasianRadiola Hill (with onlyR. linoides Roth) are nested withinLinum. McDill & al. (2009) proposed to return these four genera toLinum, in which they have been classified before.Radiola linoides should then beL. radiola L.

Euphorbiaceae (J. W. Kadereit)

Following Webster (2014; see discussion of literature there),Euphorbia L. is best treated as one large genus with >2000 spp. as the four major clades recognized (Chamaesyce, Esula, Euphorbia and Rhizanthium), partly treated as subgenera (see, e.g.,Bruyns & al. 2006;Zimmermann & al. 2010;Horn & al. 2012;Yang & al. 2012), cannot be defined morphologically. Accordingly, segregation of subclades, e.g. the Chamaesyce Clade (Webster 2014;E. subg.Chamaesyce Raf., e.g.Yang & al. 2012), at genus rank would result in a paraphyleticEuphorbia. In consequence, all species ofEuphorbia growing in Germany should be retained in that genus.

Fabaceae (C. M. Ritz)

The circumscription of the generaCytisus Desf. (40 spp.) andGenista L. (90 spp.) has been subject to long-standing discussions. The first published molecular phylogenies based on plastid (rbcL) and ITS data revealed two well-supported lineages,Cytisus andGenista, each containing numerous segregate taxa of uncertain position (Käss & Wink 1995,1997). Reviewing the published phylogenies, Cristofolini and Troia (2006) proposed a new sectional classification ofCytisus. Since raising all monophyletic entities withinCytisus s.l. to generic rank would lead to an impracticably high number of small and often monospecific genera, the authors advocated inclusion ofChamaecytisus Link. (30 spp.),Lembrotropis Griseb. (monospecific) andSarothamnus Wimm. (five spp.) inCytisus. Molecular studies did indeed not separateChamaecytisus fromCytisus s.str. (Käss & Wink 1995;Cubas & al. 2002;Pardo & al. 2004), and species with an intermediate morphology blur the boundaries between the two genera (Cristofolini & Conte 2002).Lembotropis nigricans (L.) Griseb. (Cytisus nigricans L.) is morphologically very distinct by its elongate racemes, calyx shape and naviculate hairs, but is phylogenetically nested withinCytisus (Käss & Wink 1995,1997).Sarothamnus scoparius (L.) W. D. J. Koch (C. scoparius (L.) Link), which is widespread in Europe, is part ofC. sect.Spartiopsis Dumort. with four more species distributed in the Iberian Peninsula (Cristofolini & Troia 2006).

Molecular phylogenies based on plastid and ITS markers support the monophyly of three subgenera ofGenista, but the segregate generaGenistella Ortega (Genista sagittalis L. /Genistella sagittalis (L.) Gams) andUlex L. (20 spp.;U. europaeus L. in Germany) are nested in theGenista clade (Pardo & al. 2004). However, a comprehensive revision of the complex is still missing.

The neophyticAmorpha fruticosa L. represents a poorly understood polyploid complex within the North American genusAmorpha L. (16 spp.). The monophyly of the genus is questionable: it is supported by plastid sequences, while phylogenies based on nuclear genes suggest its paraphyly because the clade also contains the North American shrubsErrazurizia rotundata (Wooton) Barneby andParryella filifolia Torr. & A. Gray (McMahon & Hufford 2004,2005;Straub & Doyle 2014). However,Amorpha is a Linnaean genus, and accordingly the name of the introducedA. fruticosa will remain unchanged if the above-named species are included inAmorpha.

Planted as ornamentals in Europe,Wisteria Nutt. contains four to seven deciduous lianas distributed in E Asia and North America. Phylogenetic reconstructions based on plastid and nuclear genes suggest the inclusion of the evergreen lianasAfgekia Craib andCallerya Endl. inWisteria (Li & al. 2014). SinceWisteria is the oldest name, the names of the cultivated species in Germany will be not affected.

Coronilla L. (nine spp.) andSecurigera DC. (13 spp.) each represent monophyletic entities in a highly supported clade that is sister toHippocrepis L. according to an ITS-based phylogeny (Sokoloff & al. 2007). However, detailed analyses based on other genetic markers so far are missing. Based on present knowledge, two options, either adopting a largeCoronilla s.l. includingSecurigera (Sokoloff 2003) or recognizing two genera (Lassen 1989), are equally possible. In the first case the nameC. varia L. and in the second caseS. varia (L.) Lassen should be used.

The most comprehensive study ofAnthyllis L., based on plastid and nuclear markers, support its monophyly whenHymenocarpus Rchb. is included inAnthyllis and the Mediterranean generaDorycnopsis Boiss. (two spp.) and the monospecificTripodion Medik, are segregated (Degtjareva & al. 2012). Contradicting results were reported in an ITS-based phylogenetic study with a relatively small taxon sampling (Nanni & al. 2004). This study placed two annual species ofAnthyllis, which were clearly part ofAnthyllis in the study by Degtjareva & al. (2012), together withTripodion nearLotus L. However, this result is questionable because resolution and taxon sampling were much lower than in the study by Degtjareva & al. (2012). In any case, the name of the GermanA. vulneraria L. would not be affected.Lotus (190 spp.) in its traditional circumscription is polyphyletic and divided into an Old World clade and several New World lineages (Allan & Porter 2000;Allan & al. 2003). The latter have now been recognized as four genera (Arambarri & al. 2005). Studies focusing on the highly supported Old World clade revealed that the segregate generaTetragonolobus Scop. (five spp.) andDorycnium Mill. (ten spp.) should be returned toLotus (Degtjareva & al. 2006;Degtjareva & al. 2008), a result already suggested by morphological studies (Polhill 1981). However, the phylogenies published so far rely on nuclear ribosomal DNA only. Since incongruencies between markers are a common phenomenon inFabaceae, additional genetic data are required. WhenDorycnium andTetragonolobus are included inLotus, the namesL. germanicus (Gremli) Peruzzi (D. germanicum (Gremli) Rikli),L. herbaceus (Vill.) Jauzein (D. herbaceum Vill.),L. hirsutus L. (D. hirsutum (L.) Ser.) andL. maritimus L. (T. maritimus (L.) Roth) should be used.

All phylogenies based on plastid and nuclear markers published so far suggest a close relationship betweenCalophaca Fisch.,Caragana L. and the monospecific AsianHalimodendron DC. (Sanderson & Wojciechowski 1996;Wojciechowski & al. 2000;Zhang & al. 2009;Duan & al. 2015). The morphologically distinctCalophaca andHalimodendron are probably nested withinCaragana, but statistical support for this was low and more research is needed (Zhang & al. 2009). In any case, the name of the introducedCaragana arborescens Lam. will not be affected by any changes in generic circumscriptions becauseCaragana is the oldest genus name.

The monophyly ofHedysarum L., containing c. 180 spp. distributed in the N hemisphere, still remains questionable. Two N African species have been excluded fromHedysarum and recognized asGreuteria Amirahmadi & Kaz. Osaloo (Amirahmadi & al. 2014). According to plastid phylogenies,Hedysarum (including the monospecific genusSartoria Boiss. from Turkey) is monophyletic (Amirahmadi & al. 2014;Duan & al. 2015). The close relationship ofHedysarum andSartoria has also been corroborated by biochemical analyses (Arslan & Ertuğrul 2010). In contrast, trees based on ITS sequences separatedH. sect.Hedysarum (containing the type,H. alpinum L.) fromH. sect.Multicaulia (Boiss.) B. Fedtsch. andH. sect.Stracheya (Benth.) B. H. Choi. The latter two were sister to a clade comprisingOnobrychis Mill. (Amirahmadi & al. 2014;Duan & al. 2015). Further studies including sequences of nuclear low-copy genes are needed to unravel the reasons for these incongruencies. If non-monophyly ofHedysarum should obtain further support, either all species ofOnobrychis and some other smaller genera should be transferred to a very largeHedysarum, orHedysarum should be split into several genera. In the latter case the nameH. hedysaroides (L.) Schinz & Thell. would remain unchanged because this species is closely related to the type of the genus name.

Similar results were obtained forOnobrychis Mill. Plastid phylogenies supportedOnobrychis as a monophyletic entity but ITS phylogenies failed to do so (Amirahmadi & al. 2014;Duan & al. 2015).

All published phylogenies revealed a close relationship betweenTrigonella L. (60 spp.) andMelilotus Mill. (20 spp.), which is supported by morphology (e.g. incised margin of stipules, notched apex of standard, smooth surface of seed coat). Most reconstructions based on either plastid, ITS or nuclear low-copy genes revealedTrigonella as paraphyletic in relation toMelilotus (Bena 2001;Steele & Wojciechowski 2003;Steele & al. 2010;Dangi & al. 2015). In contrast, a combined analysis of ITS and plastid data showed well-supported monophyly of both genera (Dangi & al. 2015). However, taxon sampling in both genera has not been sufficiently exhaustive to solve this problem. The so-called medicagoid species ofTrigonella (23 spp.) distributed in the Mediterranean area share a complex explosive tripping mechanism of pollination withMedicago (Small & al. 1987). In support of this, nuclear ribosomal sequences corroborate the inclusion of these species inMedicago (Bena 2001).

A recent comprehensive study of tribeVicieae based on plastid and ITS sequences revealed that neitherVicia L. (140 spp.) norLathyrus L. (160 spp.) are monophyletic in their current delimitation (Schaefer & al. 2012). Comparable results were also obtained by earlier studies based onmatK sequences of a small number of species (Steele & Wojciechowski 2003;Wojciechowski & al. 2004).Lathyrus is paraphyletic in relation to two monophyletic groups: the CaucasianVavilovia Fed. (two spp.) andPisum L. (three spp.;Smykal & al. 2011;Schaefer & al. 2012).Vicia appears to be paraphyletic because annual species ofV. sect.Ervum (L.) Taub. (e.g.V. tetrasperma (L.) Schreb.) andV. sect.Ervilia (Link.) W. Koch (includingV. sect.Ervoides (Godr.) Kupicha andTrigonellopsis Rech. f. andV. hirsuta (L.) Gray) were sister toLathyrus s.l. and the remaining species ofVicia includingLens Mill, (four spp.;Schaefer & al. 2012). Schaefer & al. (2012) recommended the inclusion ofPisum andVavilovia inLathyrus. Vicia could be then recognized as a monophyletic entity by includingLens and re-transferringV. articulata Hornem.,V. ervilia (L.) Willd.,V. hirsuta (L.) Gray,V. parviflora Cav.,V. sylvatica L. andV. tetrasperma (L.) Schreber to eitherErvilia Link orErvum L.

Polygalaceae (J. W. Kadereit)

Several studies (Eriksen 1993;Persson 2001;Forest & al. 2007;Abbott 2011), of which the study by Abbott (2011), although not including full results, used a large sample and both nuclear and plastid sequences, have shown that the large genusPolygala L. is highly polyphyletic. In consequence, several segregate genera of groups formerly included inPolygala have been proposed (for summary seePastore 2012). Of the species ofPolygala known in Germany,P. chamaebuxus L. should be removed fromPolygala. According to Abbott (2011), this species is part of a lineage which should be calledPolygaloides Haller and be treated asP. chamaebuxus (L.) O. Schwarz. Although not all other German species of the genus were sampled in any of the published phylogenies, their close relationship to each other has been documented (Lack 1995) and it seems safe to assume that they all will remain inPolygala.

Rosaceae (B. Gehrke)

Many genera of thePotentilleae, such asComarum L.,Dasiphora Raf.,Duchesnea Sm. and evenFragaria L., have at some point been included inPotentilla L. (Mabberley 2002). However, recent molecular phylogenetic work clearly showed thatFragariinae andPotentillinae are distinct lineages. Based on molecular work authors tend to recognizePotentillinae as comprising only two genera. These are (1)Potentilla s.str. excludingP. fruticosa L.,P. palustris (L.) Scop, andP. rupestris L. (see below) and including, amongst others,Duchesnea indica (Andrews) Teschem (asP. indica (Andrews) Th. Wolf), which is deeply nested inPotentilla s.str., and (2)Argentina Hill (Feng & al. 2015), a mostly Asian group, includingP. anserina L. (as A.anserina (L.) Rydb.). The separation ofArgentina s.l. andPotentilla s.str., which are sister lineages, is based on differences in the insertion of the styles, withPotentilla s.str. having subterminal styles, whereasArgentina has lateral ones (Dobes & Paule 2010;Sojak 2010;Feng & al. 2015). However, considering the relationship between these two genera, it would also be possible not to recognizeArgentina as a separate genus and use the namePotentilla for all species of thePotentillinae (Eriksson & al. 2015).

The other monophyletic subtribe in thePotentilleae, theFragariinae, has its highest species diversity in Asia and includes numerous smaller lineages as well asAlchemilla L.,Fragaria andSibbaldia L. Well nested inFragariinae and more closely related toFragaria than toAlchemilla or evenPotentilla areP. fruticosa andP. rupestris. These should be treated asDasiphora fruticosa (L.) Rydb., a monospecific genus, andDrymocallis rupestris (L.) Soják.Drymocallis Soják is a small genus confined to the N hemisphere. Alternatively,Fragaria could be extended to includeDasiphora andDrymocallis, amongst some other Asian groups, but the genus then would no longer be united by its characteristic fleshy receptacle. LeavingD. fruticosa andD. rupestris inPotentilla would necessitate includingAlchemilla, Fragaria andSibbaldia inPotentilla as well, which is obviously not desirable. Most authors therefore seem to prefer to recognizeDasiphora andDrymocallis as genera separate fromFragaria.

Alchemilla forms a clade withAphanes L. and the mainly South AmericanLachemilla Rydb., easily recognizable by the lack of petals and the presence of only four calyx and epicalyx lobes (Notov & Kusnetzova 2004;Gehrke & al. 2008). Molecular phylogenetic work revealed the existence of a fourth, previously unknown clade withAlchemilla species from Africa (Gehrke & al. 2008).Aphanes is clearly nested amongAlchemilla, Lachemilla and AfricanAlchemilla (Gehrke & al. 2008). As there are no obvious morphological features to separate the African clade ofAlchemilla from the European clade, and the entire clade is readily recognizable by floral morphology despite differences in life cycle, size and leaf morphology, I would like to recommend to includeAphanes inAlchemilla leading to reusing the namesAlchemilla arvensis (L.) Scop. forAphanes arvensis L. andAlchemilla microcarpa Boiss & Reut. forAphanes inexspectata W. Lippert. Irrespective of this,Alchemilla, Aphanes, andLachemilla in their traditional circumscriptions differ in habit and some details of floral morphology. WhereasAlchemilla andLachemilla species are perennial and usually have four introrse stamens inserted at the outer side of the discus (Alchemilla) or 2(-4) extrorse stamens inserted at the inner side of the discus (Lachemilla), Aphanes species are annual or short-lived and have only a single extrorse stamen at the inner side of the discus.

Potentilla palustris is most closely related toAlchemilla as circumscribed above according to chloroplast data and toSibbaldia using nuclear data. Unless included in either of these two genera, which is not desirable from a morphological point of view, it should be reinstated asComarum palustre L. It seems that especially the Asian species ofSibbaldia require more work (Eriksson & al. 2015), but it is most likely thatS. procumbens L. can retain its name.

Molecular phylogenetic work in combination with morphological character optimization has shown thatRosaceae contain only three major lineages (Potter & al. 2007):Dryadoideae, Rosoideae andSpiraeoideae. The last includes the formerly recognizedAmygdaloideae, Maloideae, Prunoideae as well asPyrinae. Evolution of derived fruit types (pome, drupe, achene) has been shown to be more complex than traditionally hypothesized (Morgan & al. 1994;Potter & al. 2002;Potter & al. 2007).

In the newly definedSpiraeoideae, the most prominent result of molecular phylogenetic work is the recognition that the species ofSorbus L. fall into two major clades. As part of the first major clade,Sorbus s.str., which is closely related toPyrus L., should include only pinnate-leaved species (Campbell & al. 2007;Potter & al. 2007;Lo & Donoghue 2012). In this clade,S. domestica L. should be placed in the monospecific genusCormus Spach according to Lo & Donoghue (2012) because this species is sister to a clade formed bySorbus s.str. andMicromeles Decne. according to chloroplast data, with a weakly supported incongruent placement ofMi-cromeles as sister toAria (Pers.) Host (see below) according to nuclear ITS sequences. If this approach were taken, the only species remaining inSorbus found in Germany would be the type,S. aucuparia L. Chloroplast and combined chloroplast and nuclear data suggest thatSorbus species with simple leaves are not closely related toSorbus s.str., but are a subclade of the second major clade also includingCydonia Mill.,Malus Mill. and others. In this subclade of simple-leavedSorbus species, Lo & Donoghue (2012) suggested to recognize the monospecific generaAria (withS. aria (L.) Crantz apparently asA. nivea Host),Chamaemespilus Medik. (withS. chamaemespilus (L.) Crantz asC. alpina (Mill.) K. R. Robertson & J. B. Phipps) andTorminalis Medik. (withS. torminalis (L.) Crantz asT. clusii (M. Roem.) K. R. Robertson & J. B. Phipps). However,Chamaemespilus andTorminalis form a well-supported clade together withAria and could also be included inAria (Li & al. 2012a;Lo & Donoghue 2012;Sennikov 2014). Lo & Donoghue (2012) pointed out that the former inclusion ofAria and satellite genera inSorbus reflects the finding that numerous apomictic microspecies in Europe and W Asia are of apparent hybrid origin involving species ofAria (incl.Torminalis) andSorbus s.str. (Aas & al. 1994;Nelson-Jones & al. 2002). Maintainance ofSorbus as one genus would require sinkingCotoneaster Medik.,Crataegus L.,Malus Mill. and many other genera inPyrus L. (Sennikov 2014), which is evidently even less desirable.

Rhamnaceae (J. W. Kadereit)

Phylogenetic studies inRhamnaceae, focusing onFrangula Mill. andRhamnus L., suggested thatFrangula andRhamnus are distinct genera, and thatRhamnus is best divided intoRhamnus s.str., the Old World genusOreoherzogia W. Vent and the New World genusVentia Hauenschild (Hauenschild & al. 2016). Of the German species ofRhamnus, R. pumila Turra falls intoOreoherzogia, in which it should be known asO. pumila (Turra) W. Vent. Following Hauenschild & al. (2016),Rhamnus s.str. andOreoherzogia can be distinguished by the number of lateral leaf vein pairs (3–5 inRhamnus, 6–20 inOreoherzogia) and by the position of a seed furrow (lateral-medial inRhamnus, dorso-medial inOreoherzogia).

Urticaceae (J. W. Kadereit)

Evidence has been presented that the generic circumscription ofParietaria L. in relation toGesnouinia Gaudich. andSoleirolia Gaudich. may require modification (Wu & al. 2013). However, no sufficiently well-sampled phylogeny is available yet to tackle this problem.

Myricaceae (J. W. Kadereit)

Myrica L. has been found to be diphyletic by Huguet & al. (2005). Following these authors (for discussion of nomenclature see their paper),M. gale L. lectotypifies the genus nameMyrica, andM. pensylvanica Mirb. should be treated asMorella pensylvanica (Mirb.) Kartesz.

Onagraceae (C. M. Ritz)

Heterogeneity ofEpilobium L. in stamen characters had already been noticed by Linnaeus. Several sections are recognized in the genus, of which onlyE. sect.Chamaenerion Ség. andE. sect.Epilobium grow in Germany. While the former has alternate leaves, weakly zygomorphic flowers with only a very short hypanthium, almost entire petals, recurved stamens of almost equal length, a recurved style and pollen in monads (type:E. angustifolium L.),E. sect.Epilobium has opposite leaves, actinomorphic flowers with a distinct hypanthium, emarginate petals, erect stamens of different length, an erect style and pollen in tetrads (lectotype:E. hirsutum L.). All phylogenetic analyses of the family, partly using a broad taxon sampling and both nuclear and plastid sequences (Baum & al. 1994;Levin & al. 2003,2004) invariably demonstrated thatE. sect.Chamaenerion is sister to the remainder of the genus. Considering this pattern of relationship, it is both possible to treatE. sect.Chamaenerion at generic rank on account of its morphological distinctness, as was done in most North American Floras, or to include it inEpilobium. If treated as a distinct genus, this would affect classification ofE. angustifolium, E. dodonaei Will. andE.fleischeri Hochst. The nameChamaenerion has long been discussed controversially.Chamaenerion Ség. instead ofChamaenerion Hill orChamerion Raf. has to be used according to Sennikov (2011).

As shown in the well-sampled phylogeny ofOnagraceae by Levin & al. (2004),Oenothera L. is only monophyletic whenCalylophus Spach,Gaura L. andStenosiphon Spach are included, as was done by Wagner & al. (2007).

Lythraceae (J. W. Kadereit)

A phylogenetic analysis ofLythraceae including several species ofLythrum L. andPeplis portula L. (Morris 2007) clearly showed thatPeplis L. is deeply nested inLythrum and should, as already done by Webb (1967), be treated asL. portula (L.) D. A. Webb.

Malvaceae (J. W. Kadereit)

A well-sampled phylogenetic analysis ofAlcea L.,Althaea L.,Lavatera L. andMalva L. using nuclear and plastid sequences by Escobar García & al. (2009) revealed that, probably with the exception ofAlcea, these genera are not monophyletic. This had been shown before forLavatera andMalva by Ray (1995). The two species ofAlthaea found in Germany fall into two only distantly related clades, withAlthaea hirsuta L. as representative of one clade more closely related toMalva /Lavatera andAlthaea officinalis L. as representative of the second clade more closely related toAlcea. Species ofMalva fall into three separate clades, of which the one containingM. alcea L. andM. moschata L. is more closely related to one of two clades ofLavatera that containsL. thuringiaca L. than to a second clade ofMalva withM. verticillata L.,M. sylvestris L. andM. neglecta Wallr. As is evident, these patterns of relationship require taxonomic changes. Escobar García & al. (2009) did not present a new classification of this “Malva alliance”, but both Banfi & al. (2005,2011) and Stace (2010) suggested to recognize an enlargedMalva containingLavatera andAlthaea hirsuta and relatives.

Resedaceae (J. W. Kadereit)

A phylogenetic analysis of a broad sample ofResedaceae using nuclear and plastid sequences by Martín-Bravo & al. (2007) demonstrated thatReseda L. is paraphyletic in relation to the generaOchradenus, Oligomeris andRandonia. This group of genera consists of two major lineages, and the four species ofReseda found in Germany fall into both.Reseda alba L. andR. luteola L. fall into two different subclades of the lineage that also containsOligomeris, whereasR. lutea L. andR. odorata L. fall into two different subclades of the lineage that also containsOchradenus andRandonia. Although the authors argued thatOchradenus andRandonia should be recognized at generic rank, they do not propose subdivision ofReseda into smaller genera. If this should eventually be proposed, the nameReseda would have to be applied to a clade containingR. lutea, the type of the genus name.

Brassicaceae (M. A. Koch)

Brassicaceae, currently recognized to contain 325 genera in 51 tribes (Al-Shehbaz 2012;Koch & al. 2012;Kiefer & al. 2014), show high levels of homoplasy in almost every morphological character used in the circumscription of tribes and genera in the past. Consequently, reliable systematic concepts often have to be obtained from molecular data, and many changes of tribal and generic circumscriptions have become necessary.

Based on molecular data,Erophila DC. is nested inDraba L. (Jordon-Thaden & al. 2010) and should be included in that genus, andE. verna (L.) Chevall. should be known asD. verna L. If recognized at species rank,E. praecox (Stev.) DC. andE. spathulata Lang should beD. praecox (Stev.) andD. spathulata (Lang) Sadler, respectively.

Several species of a formerly widely definedArabis L. have to be transferred to other genera:A. glabra (L.) Bernh. has to be treated asTurritis glabra L.,A. pauciflora Garcke asFourraea alpina (L.) Greuter & Burdet andA. turrita L. asPseudoturritis turrita (L.) Al-Shehbaz (Koch & al. 1999,2000,2001;Karl & Koch 2014). None of these three genera groups in tribeArabideae any longer (Koch & al. 2007;Couvreur & al. 2010). Even after these changes,Arabis is still a paraphyletic taxon. SinceA. alpina L. is the type of the genus name, all remainingArabis species might be transferred to newly introduced genera in the future.

Cardaminopsis Hayek is the sister group ofArabidopsis thaliana (L.) Heynh. (Koch & Matschinger 2007;Hohmann & al. 2014), and it has been widely accepted to includeCardaminopsis inArabidopsis Heinh. The German species ofCardaminopsis will beA. arenosa (L.) Lawalrée,A. halleri (L.) O'Kane & Al-Shehbaz andA. lyrata subsp.petraea (L.) O'Kane & Al-Shehbaz (Al-Shehbaz 2012;Kiefer & al. 2014).

A new classification ofThlaspi L. was proposed four decades ago (Meyer 1973,1979), recognizing the generaMicrothlaspi F. K. Mey.,Noccaea Moench andThlaspi for species ofThlaspi s.l. in the German flora. This concept has been confirmed by a series of molecular studies (e.g.Mummenhoff & al. 1997a,1997b;Koch & Mummenhoff 2001).Microthlaspi andNoccaea do not group in tribeThlaspideae, but are members of tribeColuteocarpeae (Koch & German 2013). For the German flora,T. caerulescens J. Presl & C. Presl,T. cepaeifolia (Wulfen) Koch andT. montanum L. were transferred toNoccaea and should be recognized asN. caerulescens (J. Presl & C. Presl) F. K. Mey.,N. cepaeifolia (Wulfen) Rchb. andN. montana (L.) L. K. Mey., respectively.Thlaspi perfoliatum, with its two morphologically slightly differentiated cytotypes (T. erraticum Jord. andT. improperum Jord.;Koch & Bernhardt 2004), has to be included inMicrothlaspi asM. perfoliatum (L.) F. K. Mey. It has also been proposed to combine most genera of tribeColuteocarpeae in a broadly definedNoccaea (Al-Shehbaz 2012). However, since comprehensive molecular analyses of the entire tribe with its more than 125 species (Koch & German 2013) are lacking, this concept should not be followed at the moment.

Considering the German flora,Alyssum saxatile L. has been shown to be member of a clade including various species ofAurinia Desv., which is sister toBornmuellera Hausskn. andClypeola L. (Cecchi & al. 2010;Resetnik & al. 2013). Consequently,A. saxatile is best treated asAurinia saxatilis (L.) Desv. All otherAlyssum species in Germany belong to a then monophyleticAlyssum.

Integration ofDentaria L. inCardamine L. (Carlsen & al. 2009) and ofCoronopus Mill. inLepidium L. (Al-Shehbaz & al. 2002;Mummenhoff & al. 2008) is strongly supported and both are nested in the respective genera in molecular analyses. The fourDentaria species of the German flora should be known asCardamine bulbifera (L.) Crantz,C. enneaphyllos (L.) Crantz,C. heptaphyllos (Vill.) O. E. Schulz andC. pentaphyllos (L.) Crantz.Coronopus didymus (L.) Sm. andC. squamatus (Forrsk.) Asch. are now best treated asLepidium didymum L. andL. coronopus (L.) Al-Shehbaz, respectively.

Pritzelago Kuntze andHymenolobus Nutt. of tribeErysimeae are best included inHornungia Bernh. These three genera form a well-supported clade (Mummenhoff & al. 2001;Kropf & al. 2003), and it has been demonstrated that there is no single character that reliably distinguishes the three genera (Al-Shehbaz & Appel 1997). Consequently, the following names should be used:Hornungia alpina (L.) O. Appel,H. petraea (L.) Rchb. andH. procumbens (L.) Hayek.

Maximum-likelihood trees derived from ITS1 and ITS2 sequences available from BrassiBase (Koch & al. 2012;Kiefer & al. 2014;  http://brassibase.cos.uni-heidelberg.de/ clearly show thatCheiranthus cheiri L. is nested inErysimum L., where it should be calledE. cheiri (L.) Crantz.Hirschfeldia Moench of tribeBrassiceae consists of one species only:H. incana (L.) Lagr.-Fossat is most closely related toErucastrum C. Presl. (including its type,E. virgatum C. Presl.;Arias & al. 2014). However, sinceErucastrum as currently treated is a polyphyletic genus, and various otherErucastrum species might be transferred to different genera in future (Arias & Pires 2012), it seems best to keepHirschfeldia separate until various phylogenetic hypotheses have been tested in more detail.

Santalaceae (J. W. Kadereit)

Thesium L. was found to be monophyletic only when the generaAustroamericum Hendrych andThesidium Sonder are included (Moore & al. 2010). Discussing the options of either sinking these two genera intoThesium or maintaining them, requiring splitting ofThesium in its traditional circumscription into several smaller genera, Moore & al. (2010) preferred the former option for morphological reasons.

Polygonaceae (K. Wesche)

InRumex L. two monophyletic subgenera can be distinguished:R. subg.Acetosa (Mill.) Rech. f. andR. subg.Rumex. This is the approach currently chosen in most C European floras, although it is possible (but not mandatory) to raise these subgenera to generic rank (Hejný & Slavik 1990). According to molecular analyses,R. subg.Acetosa includes the sometimes separately treatedR. subg.Acetosella (Meisn.) Rech. f. (Schuster & al. 2015). This is supported by shared morphological characters, e.g. the presence of hastate leaves.

The taxonomy ofPolygonum L. has posed particular challenges. The traditional broad concept had survived two centuries in spite of repeated criticism including calls to split the genus into up to nine sections, which commenced as early as 1856 (Meisner 1856). Based on morphological evidence, Haraldson (1978) reinforced these earlier proposals for splittingPolygonum, which have since been confirmed by studies of both plastid and nuclear DNA markers (Lamb Frye & Kron 2003;Galasso & al. 2009;Schuster & al. 2015).Polygonum s.l. clearly is polyphyletic and should be split into several genera, partly even belonging to different tribes. Some details, however, are still controversial, given that new molecular studies continue to differ from preceding ones, and no final conclusions have been reached. Accordingly, all inferences remain somewhat tentative.

Species of tribePolygoneae have outer tepals with one primary vein and include a range of life forms.Polygonum s.str. is characterized by a distinct pollen morphology and by outer tepals that do not develop large appendages in fruit (Schuster & al. 2011a). In Germany it comprises few, mainly ruderal species(P. aviculare L. agg. - includingP. arenastrum Borean,P. oxyspermum Ledeb. andP. raii Bab., the latter sometimes treated as a subspecies ofP. oxyspermum). In our context, these species are distinct by having essentially solitary or at the most approximate flowers in axillary glomerules and a silvery ochrea. Genetic studies support the monophyly ofPolygonum L. s.str. (Schuster & al. 2015).

A clade related toPolygonum L. s.str. contains the generaReynoutria Houtt. andFallopia Adans. Their taxonomy is notorious for frequent changes and their treatment is inconsistent among C European Floras (Fischer & al. 2008;Jäger 2011;Tison & de Foucoult 2014).Fallopia in its traditional circumscription contains mostly lianas, whileReynoutria includes extremely tall herbs that are invasive in many regions. Both taxa share the presence of extrafloral nectaries and have wings on the floral bracts. Viable intergeneric hybrids are known, and polyploidy and extreme morphological variability add to the taxonomic difficulties. In consequence,Reynoutria has often been included in a broaderFallopia s.l., where it was treated as a section. Uncertainty about the treatment of the two genera pertains, although molecular approaches have used both chloroplast and nuclear markers for a very good taxonomic coverage. These studies support the monophyly of each of the two genera (Schuster & al. 2011b,2015). The S hemisphereMuehlenbeckia Meisn., however, has been identified as closely related (Haraldson 1978), and recent molecular studies implied that it is indeed sister toFallopia (Schuster & al. 2011b,2015). This is in line with the fact that bothFallopia andMuehlenbeckia share a base chromosome number of 10 (11 inReynoutria) and contain flavones (absent inReynoutria). IncludingReynoutria but notMuehlenbeckia in a broadly circumscribedFallopia would thus result in a polyphyletic group. In view of this, keepingFallopia,Muehlenbeckia andReynoutria as separate genera currently is the best — but not necessarily final — solution.

The second large tribe relevant for relationships ofPolygonum s.l. in Germany are thePersicarieae, which are monophyletic and morphologically distinct by the presence of three veins arising from the base of the tepals, of nectaries and of non-dilated stamen filaments (Lamb Frye & Kron 2003;Kim & Donoghue 2008;Sanchez & Kron 2008). The tribe includesAconogonon (Meisn.) Rchb.,Bistorta Mill, andPersicaria (L.) Mill.Persicaria is characterized by spicate or capitate panicles, a usually entire but often ciliate or pectinate ochrea, and has 4–8 stamens and 4 or 5 tepals. All recent treatments agree that it is monophyletic and should be excluded from tribePolygoneae (Kim & Donoghue 2008;Fan & al. 2013). Thus, the following combinations should be used for the German species:Persicaria amphibia (L.) Delarbre,P.hydropiper (L.) Delarbre,P. lapathifolia (L.) Delarbre,P.maculosa Gray,P. minor (Huds.) Opiz,P. mitis (Schrank) Assenov andP. pensylvanica (L.) M. Gómez. Except forP. maculosa (formerlyPolygonum persicaria L.), epithets could be directly adopted from former names inPolygonum. Although the highly variableP. amphibia is a morphologically distinct taxon withinPersicaria (Kim & Donoghue 2008), there is no need to raise it to genus level (Galasso & al. 2009).

Bistorta Mill, is morphologically distinct (with a rosette of basal leaves and usually only one terminal, spicate panicle), and both chloroplast and nuclear data imply that it is monophyletic withinPersicarieae (Kim & Donoghue 2008;Fan & al. 2013). Molecular approaches, however, are not fully consistent with respect to its exact relationships toAconogonon andKoenigia L. Nonetheless, most current Floras and also molecular studies (Galasso & al. 2009;Sanchez & al. 2011;Schuster & al. 2011a) accept its generic rank. The German species thus have to be namedB. officinalis Delarbre andB. vivipara (L.) Delarbre (formerlyPolygonum bistorta L. andP. viviparum L., respectively).

The taxonomy ofAconogonon is particularly complicated. Species in this group have been placed inPersicaria,Polygonum orRubrivena M. Krái (the last forA. polystachyum (Meisn.) Small as the only species ofAconogonon s.l. occurring in Germany). Recent molecular studies implied thatAconogonon species are distinct fromBistorta andPersicaria, but also revealed their close relationship with the mostly boreal and polarKoenigia (Galasso & al. 2009;Sanchez & al. 2009).Aconogonon andKoenigia have broadly similar pollen, and the two genera cannot easily be separated by morphological characters. Studies based on cpDNA have suggested thatKoenigia in its traditional circumscription may be nested betweenAconogonon andRubrivena (Sanchez & al. 2011). The so-far most comprehensive study covering many taxa and employing both cpDNA and nuclear markers (Schuster & al. 2015) confirms this close relationship and finds one large clade that comprises all analysed species ofAconogonon andKoenigia (andRubrivena). While mostKoenigia species form a distinct group, some (but not all!) accessions ofK. delicatula (Meisn.) H. Hara are sister group to a clade comprising otherAconogonon andKoenigia species (incl.A. polystachyum). This implies thatKoenigia in its traditional sense is not monophyletic. Relationships ofAconogonon are even more puzzling, with a number of polyploid Aconogonon species being more closely related toKoenigia than to other members of the genus. Moreover, different accessions of someAconogonon species appear on very different branches in theAconogonon /Koenigia clade. Details of the evolution of this group clearly are not fully understood, and thus Schuster & al. (2015) advocate the fusion of all taxa in one large genus. They propose to unite them under the nameKoenigia, which was chosen for priority reasons. These authors also draw the necessary taxonomic consequences and provide the new combinationK. polystachya (Meisn.) T. M. Schust. & Reveal.

Though using a somewhat smaller species set, Fan & al. (2013) also presented a comprehensive molecular study, which confirmed the odd position ofK. delicatula (plus oneAconogonon species). In their analysis, A.polystachyum is nested within otherAconogonon species, which jointly form the sister clade to the coreKoenigia species. Fan & al. (2013) also discussed the possibility to adopt a broad concept ofKoenigia. However, they acknowledged that merging the largerAconogonon in the smallerKoenigia is somewhat impractical and also remarked on the apparently different chromosome base numbers in the two groups. They advocated keeping the two genera independent and placing the oddK. delicatula in a new monospecific genus, for which no valid name is available yet. This would also be supported by some of its morphological features that are transient toPersicaria. Splitting the whole complex into several, partly new genera indeed is an alternative solution to the problem implied by the tree of Schuster & al. (2015), but would presumably result in the formation of many small genera such asRubrivena. Given that details of the evolution ofAcononogon /Koenigia remain unclear, I opt for an intermediate position. The special position ofAconogonon andKoenigia inPersicarieae is undebated, but instead of drawing far-reaching taxonomic consequences, I rather acknowledge the level of uncertainty by keepingAconogonon as a separate genus for the time being. In line with Fan & al. (2013), I regard evidence for a separate genusRubrivena as questionable and maintain the established name A.polystachyum for the taxon occurring in C Europe.

Caryophyllaceae (M. S. Dillenberger)

Regarding generic delimitations in theCaryophyllaceae, Greenberg & Donoghue (2011) stated: “none of the eight largest genera (Arenaria, Cerastium, Dianthus, Gypsophila,Minuartia, Paronychia, Silene, Stellaria) appear to be strictly monophyletic”. For some genera taxonomic adjustments have already been made (e.g.Dillenberger & Kadereit 2014), but not for all. All taxonomic changes that were recently made for taxa in the German flora, or that need to be made in the future, are related to these eight genera.

There are several problems concerning the monophyly ofCerastium L. andStellaria L.Cerastium is an almost cosmopolitan genus with about 100 species.Stellaria is cosmopolitan, too, and contains c. 120 species (Mabberley 2008). Both genera have emarginate to deeply lobed petals, but this character is shared with other genera, e.g.Myosoton Moench (Bittrich 1993).Myosoton is a monospecific genus, withM. aquaticum (L.) Moench as its only species. This species has recently (Jäger 2011;Seybold 2011) been treated as part ofStellaria, asS. aquatica L. This is congruent with the findings of Greenberg & Donoghue (2011), whereS.aquatica is nested with good support in a clade of severalStellaria species, includingS. media (L.) Vill. but not the type ofStellaria, S. holostea L., and is closely related toS. bungeana Fenzl. Unfortunately,Stellaria does not become monophyletic by includingMyosoton. With good support,S. holostea is sister to a clade containing the largest part ofStellaria, but alsoCerastium, Holosteum L. andMoenchia Ehrh. Furthermore,Cerastium is not monophyletic since a well-supported clade of two species,C. cerastoides (L.) Britton andC. dubium (Bastard) Guépin, is sister toHolosteum. This position is poorly supported, butMoenchia is sister to the rest ofCerastium with good support, making it impossible to retain the twoCerastium species inCerastium without including at leastMoenchia. To amend these various violations of monophyly there are at least two possible solutions:

InGypsophila L. and relatives of interest (i.e.Dianthus L.,Petrorhagia (Ser.) Link andVaccaria Wolf.) two issues need to be discussed. The first is the treatment ofVaccaria. Vaccaria is a monospecific genus containing onlyV. hispanica (Mill.) Rauschert. This species is native to Eurasia, especially the Mediterranean region, but has become naturalized in large parts of the world (S Africa, Australia and North and South America). The phylogeny of Greenberg & Donoghue (2011) unambiguously placedVaccaria withinGypsophila. It differs fromGypsophila mainly by its calyx wings. The position in the phylogeny allows two alternative solutions.

I prefer the second solution for different reasons.Vaccaria is quite similar toGypsophila and its inclusion in that genus will not require large changes in the circumscription ofGypsophila. The other point is that it is difficult to justify splittingGypsophila into different genera only to retainVaccaria as an independent genus. As Greenberg & Donoghue (2011) included only few of the 150Gypsophila species in their phylogeny, I cannot foresee to what extent an independentVaccaria would affect subdivision ofGypsophila.

The second issue concerns the paraphyly ofPetrorhagia in relation toDianthus, and the position ofGypsophila muralis L. and several otherGypsophila species from outside Germany as sister toDianthus andPetrorhagia instead of being part of the rest ofGypsophila. Petrorhagia is a genus with 33 species distributed from the Canary Islands across the Mediterranean region to Kashmir (Mabberley 2008). Although the phylogeny of Greenberg & Donoghue (2011) contains only three species ofPetrorhagia, it unambiguously shows that the genus is paraphyletic. Two solutions seem possible:

Minuartia L. (sensuMcNeill 1962) comprises about 175 species that are distributed in the N hemisphere. It was delimited from most other genera ofCaryophyllaceae by a combination of three styles and three capsule valves. Molecular phylogenies revealed that the genus consists of ten independent lineages (Fior & al. 2006;Harbaugh & al. 2010;Greenberg & Donoghue 2011;Dillenberger & Kadereit 2014), each of which is closest relative of another genus or group of genera. According to Dillenberger & Kadereit (2014) the genus is best divided into 11 genera instead of including other genera inMinuartia. The ten lineages were divided into 11 genera because in one case there was no morphological or karyological character or combination of characters to define this clade as one genus. Therefore two subclades with more uniform morphologies were described as genera. Including other genera inMinuartia would have affected most genera of subfam.Alsinoideae or subfam.Alsinoideae and subfam.Caryophylloideae. In consequence, several species ofMinuartia in the C European flora need to be treated as part of other genera.Minuartia species transferred to other genera are:Cherleriasedoides L.(M. sedoides (L.) Hiern),Facchinia cherlerioides (Sieber) Dillenb. & Kadereit(M. cherlerioides (Sieber) Bech.), present in the German flora only withF. cherlerioides subsp.aretioides (Port, ex J. Gay) Dillenb. & Kadereit,F. rupestris (Scop.) Dillenb. & Kadereit(M. rupestris (Scop.) Schinz & Thell.),Sabulina austriaca (Jacq.) Rchb.(M. austriaca (Jacq.) Hayek),S. stricta (Sw.) Rchb.(M. stricta (Sw.) Hiem),S. tenuifolia (L.) Rchb.(M. hybrida (Vill.) Schischk.),S. verna (L.) Rchb.(M. verna (L.) Hiem) andS. viscosa (Schreb.) Rchb.(M.viscosa (Schreb.) Schinz & Thell.). The only two species in Germany that remain inMinuartia areM. rubra (Scop.) McNeill andM. setacea (Thuill.) Hayek.

Silene L. contains c. 700 species that are restricted to the N hemisphere (Mabberley 2008). Although the genus is large, there exist only small problems with its monophyly. One point concernsLychnis L., which contains c. 20 species distributed in N-temperate and arctic regions (Bittrich 1993). Its treatment as separate fromSilene L. has repeatedly been regarded as doubtful (seeOxelman & Lidén 1995).Lychnis has usually five styles and five capsule teeth, whereasSilene has three or five styles and six or ten capsule teeth. Even the most recent phylogeny of theCaryophyllaceae could not unambiguously determine the position ofLychnis (Greenberg & Donoghue 2011). In that studySilene seems to be paraphyletic in relation toLychnis. However, this position is not well supported, and a change of position is possible. For the moment, the species ofLychnis in the German flora, i.e.L. coronaria (L.) Desr. andL. flos-cuculi L., should be maintained, but future inclusion inSilene, asS. coronaria (L.) Clairv. andS. flos-cuculi (L.) Clairv., seems likely.

The second problem is related toCucubalus baccifer L. AlthoughSilene is not sufficiently well supported, the position ofC. baccifer seems to be clearly withinSilene (Greenberg & Donoghue 2011). Therefore it seems advisable to treat this species asS. baccifera (L.) Roth.

Several problems hinge on the acceptance ofHeliosperma Rchb. and other smaller genera. When acceptingHeliosperma, several smaller genera need to be recognized in order to keepSilene monophyletic. One of these genera isAtocion Adans. Based on a molecular phylogeny, Lidén & al. (2001) excluded five species, includingS. armeria L. andS. rupestris L., fromSilene and included them inAtocion. These results were verified with a large sample ofSilene and related genera by Greenberg & Donoghue (2011) and should have taxonomic consequences.Atocion is sister toViscaria Bernh, and the names for the two species areA.armeria (L.) Raf. andA.rupestre (L.) Oxelman. An inclusion of Atocion inSilene would also affectEudianthe Rchb.,Heliosperma andViscaria and is therefore not desirable.Atocion is glabrous, has elliptic or oblanceolate leaves, a regular dichasium, and flowers with entire or emarginate petals and three stigmas (Lidén & al. 2001).Silene species with the same character combination of hairiness, inflorescence type and stigma number have lower leaves that are spathulate and petals that are usually lobed. Furthermore, theseSilene species have anastomosing calyx veins, butAtocion has non-anastomosing veins (Lidén & al. 2001).

Irrespective of the inclusion ofLychnis inSilene or its separate treatment,L. viscaria L. is not part of either of these two genera. The species clearly belongs to a wellsupported clade that is sister toAtocion (Greenberg & Donoghue 2011). The correct genus name for the species of this clade isViscaria Bernh., andL. viscaria should be known asV. vulgaris Bernh.Viscaria vulgaris is the type ofViscaria.

Another problem is related toSilene pusilla Waldst. & Kit., which is nested in the well-supportedHeliosperma (Rchb.) Rchb. The inclusion ofS. pusilla inHeliosperma asH. pusillum (Waldst. & Kit.) Rchb. is justified and necessary.

Chenopodiaceae (G. Kadereit)

Chenopodium L. in its traditional wide circumscription, comprising c. 150 spp. worldwide, has been shown to be highly polyphyletic withChenopodium lineages spread all over the phylogeny of subfam.Chenopodioideae (Kadereit & al. 2010;Fuentes-Bazán & al. 2012a,2012b). According to Fuentes-Bazán & al. (2012a,2012b), species ofChenopodium belong to six different genera:Blitum L.,Chenopodiastrum S. Fuentes & al.,Chenopodium L. s.str.,Dysphania R. Br.,Lipandra Moq. andOxybasis Kar. & Kir. Although the sampling for the molecular analyses was far from complete, the polyphyly ofChenopodium seems well supported and future studies will reveal where unsampled species belong. Twenty of the 23 species of formerChenopodium occurring in the German flora were included in the molecular studies by Fuentes-Bazán & al. (2012a,2012b), and these are distributed among all six genera.Blitum is represented by three species:B.bonus-henricus (L.) Rchb. (C.bonus-henricus L.),B.capitatum L. (C.capitatum (L.) Aschers.) andB. virgatum L.(C. foliosum Aschers.).Chenopodiastrum is represented byChenopodiastrum hybridum (L.) S. Fuentes & al.(Chenopodium hybridum L.) andChenopodiastrummurale (L.) S. Fuentes & al.(Chenopodium murale L.). Species with glandular hairs and an aromatic odour clearly need to be classified inDysphania, which is only distantly related to coreChenopodium. In the German flora these areD. ambrosioides (L.) Mosyakin & Clemants(C.ambrosioides L.),D. botrys (L.) Mosyakin & Clemants (C. botrys L.),D. pumilio (R. Br.) Mosyakin & Clemants (C. pumilio R. Br.) andD. schraderiana (Schult) Mosyakin & Clemants (C. schraderianum Schult).Lipandra is represented byL. polysperma (L.) S. Fuentes & al.(C. polyspermum L.) andOxybasis byO. chenopodioides (L.) S. Fuentes & al. (C. botryodes Sm.),O. glauca (L.) S. Fuentes & al. (C.glaucum L.),O. rubra (L.) S. Fuentes & al. (C. rubrum L.) andO. urbica (L.) S. Fuentes & al.(C. urbicum L.). Of the remaining species present in the German flora,Chenopodium album L.,C. berlandieri Moq.,C. ficifolium Sm.,C. opulifolium Schrader ex Koch & Ziz,C. patericola Rydb. andC. vulvaria L. belong toChenopodium s.str.Chenopodium hircinum Schrader,C. strictum Roth andC. suecicum Murr have not yet been included in molecular analyses.Chenopodium aristatum L. (Dysphania aristata (L.) Mosyakin & Clemants) is a neophyte in the German flora and should be treated asTeloxysaristata (L.) Moq. This monospecific genus is closely related toCycloloma Moq.,Dysphania andSuckleya A. Gray (Kadereit & al. 2010;Fuentes-Bazán & al. 2012a).

Halimione Aellen is well-supported sister group of the large genusAtriplex L., from which it can be distinguished by unique seed and fruit characters (Kadereit & al. 2010). Inclusion ofHalimione intoAtriplex as proposed in Sukhorukov (2006) is possible, but not recommended by the present author (G. Kadereit).

Bassia All. andKochia Roth were both found to be polyphyletic in molecular studies (Kadereit & Freitag 2011;Kadereit & al. 2014). Most species ofKochia including the two species present in Germany,K. laniflora (S. G. Gmelin) Borbás andK. scoparia (L.) Schrader, have been included inBassia, and the remaining species were classified in two new genera,Eokochia Freitag & G. Kadereit andGrubovia Freitag & G. Kadereit. Other species ofBassia (B. dasyphylla Kuntze,B. hirsuta (L.) Kuntze andB. sedoides (Schrad.) Asch.) needed to be transferred to new genera (Grubovia dasyphylla (Fisch. & C. A. Mey.) Freitag & G. Kadereit,Spirobassia hirsuta (L.) Freitag & G. Kadereit andSedobassia sedoides (Schrad.) Freitag & G. Kadereit) in order to define monophyletic genera inCamphorosmeae (Kadereit & Freitag 2011). Of these new genera onlySpirobassia (S. hirsuta) occurs in Germany.

Salsola L. is a large and highly polyphyletic genus (Akhani & al. 2007). Unfortunately there is disagreement among experts concerning the typification ofSalsola. Mosyakin & al. (2014) proposed a conserved type,S. kali L., while Akhani & al. (2014) argued in favour of the current type,S. soda L. IfS. soda is accepted as type ofSalsola, S. kali has to be included inKali Mill., asKalisoda Moench (Akhani & al. 2007).

Nyctaginaceae (J. W. Kadereit)

As shown by Levin (2000),Oxybaphus Willd. is clearly part ofMirabilis L. where it should have the rank of section. Accordingly,O. nyctagineus (Michx.) Sweet should be treated asM. nyctaginea (Michx.) MacMill.

Hydrangeaceae (J. W. Kadereit)

AlthoughPhiladelphus L. appears to be paraphyletic in relation to the monospecificCarpenteria Torr. (Guo & al. 2013), classification ofP. coronarius L. as aPhiladelphus would not be affected asP. coronarius is the type of the genus name.Philadelphus inodorus L. falls into the same clade asP. coronarius.

Primulaceae (J. W. Kadereit)

Mast & al. (2001) demonstrated thatCortusa L. is deeply nested inPrimula L. Accordingly, it should be treated asP. matthioli (L.) V. A. Richt.

As summarized by Manns & Anderberg (2009), several studies using either nuclear, plastid or both nuclear and plastid sequences have shown that a non-monophyleticAnagallis L. (incl.Centunculus L.),Glaux L. andTrientalis L. (as well as the non-C-European generaAsterolinon Hoffsgg. & Link andPelletiera A. St. Hil.) are all nested in a highly paraphyleticLysimachia L. Based on a careful consideration of morphological variation in this group of genera, and facing the choice between including all inLysimachia or splittingLysimachia in such a way that at least some of the above genera can be maintained, Manns & Anderberg (2009) argue: “It is, however, difficult to establish morphological characters to distinguish between different subgroups withinLysimachia and the morphological distinctiveness of these subgroups is not very high. Furthermore, the characters used to recognizeLysimachia are also present inAnagallis and to large extent also inAsterolinon, Pelletiera andTrientalis. Consequently, proposal of new genera for someLysimachia (e.g.L. nemorum L. andL. serpyllifolia Schreb.), or transfer ofL. nemorum and allied taxa toAnagallis would inevitably result in poorly diagnosed genera. Choosing among alternatives, we find it better to merge the smaller segregate genera withLysimachia, rather than splittingLysimachia further.” Through earlier work and the work by Banfi & al. (2005) and Manns & Anderberg (2009) combinations are available for C EuropeanAnagallis (plusCentunculus), Glaux andTrientalis as species ofLysimachia. These would beL. arvensis (L.) U. Manns & Anderb. (Anagallis arvensis L.),L. europaea (L.) U. Manns & Anderb. (Trientalis europaea L.),L.foemina (Mill.) U. Manns & Anderb. (A.foemina Mill.),L.maritima (L.) Galasso & al. (Glaux maritima L.) andL.tenella L. (A.tenella (L.) L.).

Ericaceae (M. D. Pirie)

Three genera have been recently re-delimited to make them monophyletic. The first isKalmia L., which becomes monophyletic only after inclusion ofLoiseleuria Desv. (Gillespie & Kron 2013). Accordingly,Loiseleuriaprocumbens (L.) Desv. should be known asKalmiaprocumbens (L.) Gift & al. ex Galasso & al.

The second isRhododendron L., with c. 850 species, which should includeLedum L. based on morphological evidence by Kron & Judd (1990) and molecular evidence by, e.g., Goetsch & al. (2005). In Germany, the nativeL. palustre L. should be known asR. tomentosum Harmaja and the introducedL. groenlandicum Oeder asR. groenlandicum (Oeder) Kron & Judd.

The third isMonotropa L., which is replaced byHypopitys Hill, in Jäger (2011). Evidence from nuclear encoded markers suggests that the type ofMonotropa, M. uniflora L., and that ofHypopitys, H. monotropa Crantz (M. hypopitys L.), are more closely related to other monotropoid genera than to each other (Bidartondo & Bruns 2001). Species delimitation withinHypopitys is controversial, but resolution of the precise number and delimitation of species (includingH. hypophegea G. Don in Germany) across its broad geographic range seems unlikely to further affect generic boundaries.

Problems in generic delimitation remain inVaccinium L. A number of different genera are apparently nested between its c. 450 species, with no evidence to suggest that the type,V uliginosum L., is closely related to any of the other species of the German flora, and clear indication thatV myrtillus L. is more closely related to species elsewhere (Powell & Kron 2002). As the specialists are apparently not in favour of expanding the circumscription ofVaccinium it is likely that name changes will yet be required, but the current phylogenetic hypothesis is insufficiently resolved and sampled to offer a solution.

Rubiaceae (F. Ehrendorfer)

Since more than 20 years ago, DNA-analytical phylogenetic studies on the critical tribeRubieae (Rubiaceae) have become available (e.g.Ehrendorfer & al. 1994;Manen & al. 1994;Natali & al. 1995,1996;Soza & Olmstead 2010a,2010b; and particularlyEhrendorfer & Barfuss 2014:Fig. 1 & 2, with clades and their reference numbers). These studies have made it increasingly clear that the traditional generaAsperula L. andGalium L., both well represented in the flora of Germany (Jäger 2005), are polyphyletic in their present circumscriptions. Monophyly was documented only forCruciata Mill.,Rubia L. andSherardia L. In order to achieve monophyly forAsperula andGalium, one would have to lump all these genera (and several others exceptRubia) into a giantGalium s.latiss. with about 900 species worldwide and a very complex infrageneric classification.

If a more narrow generic concept for C EuropeanRubieae is preferred,Asperula would have to be restricted to its type, the annual A.arvensis L., and its perennial sister taxon A.taurina L. (clade V-B). The large A. sect.Cynanchicae (DC.) Boiss. (with A.cynanchica L. and A.neilreichii Beck), centred in the Mediterranean area, is more closely related toSherardia (both in clade V-A) than toAsperula s.str. and might also deserve separate generic status. This also applies to A.tinctoria L., a member of the traditional A. sect.Glabella Griseb. (clade V-C) with a disjunct Eurasian distribution. Also into clade V-C fallsGalium sect.Aparinoides (Jord.) Gren., a sube lade of limnic habitats with a worldwide distribution, typified byG. palustre L., a well-known element of the European flora. The morphological distinctness and deviating chromosome base numberx = 12 (otherwise mostlyx = 11 inRubieae) also suggest generic separation of this subclade.

It was no surprise to find twoGalium species(G. boreale L. andG. rotundifolium L.) fromG. sect.Platygalium (DC.) W. D. J. Koch in the same clade (V-D) as the generally recognized generaCruciata andValantia L.: they all are characterized by whorls of two leaves and only two additional leaf-like stipules. This and the relevant DNA data could justify the transfer ofG. boreale andG. rotundifolium to a separate genus, corresponding toG. sect.Platygalium s.latiss. (also including the former European genusTrichogalium Fourr., the American genusRelbunium (Endl.) Benth. & Hook, and probably also the monotypieMicrophysa Schrenk from C Asia) with a worldwide distribution and up to 230 other, clearly related formerGalium andRelbunium species centred in E Asia and the Americas.

The majority of the remaining C EuropeanGalium species (24 in Germany;Jäger 2005) always have leaves and leaf-like stipules in whorls of more than four (and up to 12). They are clearly verified as members of a worldwide “monophylum” that corresponds to clade VI and the genusGalium s.str. with about 350 predominantly Old World species. The relationships of its species in Germany correspond quite well with the following more or less DNA-supported taxonomic sections:G. sect.Aparine W. D. J. Koch(G. aparine L. andG. spurium L.),G. sect.Aspera (DC.) W. D. J. Koch, syn.:G. sect.Microgalium Griseb.(G. parisiense L.),G. sect.Galium (G. album Mill.,G. aristatum L.,G. glaucum L.,G. intermedium Schultes [G.schulte sii Vest], G.lucidum All., G.mollugo L., G.xpomeranicum Retz., G.sylvaticum L., G.truniacum (Ronn.) Ronn. and G.verum L.), G. sect.Hylaea (Griseb.) Ehrend. (G.odoratum (L.) Scop.), G. sect.Kolgyda Dumort. (G.tricomutum Dandy and G.verrucosum Huds.), G. sect.Leptogalium (G.anisophyllon Vill., G.megalospermum All, G.noricum Ehrend., G.pumilum Murray, G.saxatile L., G.sterneri Ehrend, and G.valdepilosum H. Braun) and G. sect.Trachygalium K. Schum. (G.uliginosum L.).

A more detailed presentation of our current knowledge concerning relationships within tribeRubieae in C Europe can be found in Kästner & Ehrendorfer (in press). Before one can begin to execute the possible and DNA-supported taxonomic and nomenclatural changes within theRubieae discussed above, further critical research appears obligatory.

Gentianaceae (J. W. Kadereit)

Several phylogenetic studies ofGentianaceae-Swertiinae (Chassot & al. 2001;von Hagen & Kadereit 2001,2002) have shown that generic circumscriptions in this group require substantial revision. Thus, it is evident thatGentianella ciliata (L.) Borkh. and G.tenella (Rottb.) Börner are only distantly related toGentianella s.str., and should be treated asGentianopsis ciliata (L.) Ma andComastoma tenellum (Rottb.) Toyok., respectively. Even after exclusion of these (and related) species,Gentianella is polyphyletic, as isSwertia L. If this eventually should result in the recognition of several smaller genera, the generic identity of the remaining German species ofGentianella would remain unaffected as they fall into the same clade as the type of the genus name,G. campestris (L.) Börner. AsS. perennis L. is the type ofSwertia, recognition of segregate genera will not affect the generic identity ofS. perennis. For descriptions and discussion of genera see Struwe & al. (2002). Inclusion ofComastoma (Wettst.) Toyok. andLomatogonium A. Braun inGentianella, as suggested by Banfi & al. (2005), who in consequence provided a combination forL. carinthiacum (Wulfen) Rchb. inGentianella, is not justified by the data available unless a much larger number of lineages, including several lineages ofSwertia, are included inGentianella.

Oleaceae (J. W. Kadereit)

As first suspected by Wallander & Albert (2000) on the basis of plastid sequences, a monophyleticLigustrum L. was found deeply nested in a paraphyleticSyringa L. using nuclear sequences (Li & al. 2002). In consequence, inclusion ofLigustrum inSyringa may have to be considered once stronger evidence for such relationship is available. Interestingly, one species ofLigustrum, L. sempervirens (Franch.) Lingelsh., sometimes classified as a separate genus, is intermediate in fruit morphology betweenSyringa (capsules) andLigustrum (berries or drupes) by having berries that become leathery and eventually dehisce.

Plantaginaceae (D. C. Albach)

A hundred years ago,Veronica L. included allScrophulariaceae with a tetramerous flower and short corolla tube, two stamens and a flattened capsule. In that circumscription the genus included approximately 300 species. Subsequent authors treated more and more groups of distinct species as separate genera, such asHebe Juss. mainly from Australasia,Pseudolysimachion Opiz from Eurasia(V. longifolia L. andV. spicata L. in the German flora) andVeronicastrum Farw. from E Asia and E North America. The first DNA-based phylogenetic analyses (e.g.Albach & Chase 2001; Wag staff & al. 2002;Albach & al. 2004a) supported the separation of some genera(Paederota L. andVeronicastrum), but demonstrated that most genera split off in the 19^th and 20^th centuries are nested in a lineage that should be recognized as a monophyleticVeronica. These results caused a sometimes heated discussion on whether autapomorphies need to be considered as important as symplesiomorphies (e.g.Brummitt 2006). However, subsequent analyses added support to the molecular results and demonstrated that autapomorphies of these segregate genera are not as clear as sometimes believed, and that morphological transitions betweenVeronica and groups considered distinctive commonly exist. For example, such transitional species betweenPseudolysimachion andVeronica occur in E Asia and Japan (Albach 2008). Thus, based on molecular and morphological arguments, these analyses suggest inclusion of these genera inVeronica rather than further splitting (Albach & al. 2004b;Garnock-Jones 2007). In C Europe, reintegration of AustralasianHebe and relatives and North AmericanSynthyris Benth. will be of interest mainly to horticulturists, but reintegration ofPseudolysimachion, the species ofVeronica with dense, spicate inflorescences, reverses a split adopted by many European Floras since the 1960s (Holub & Pouzar 1967). All European species ofPseudolysimachion were originally described as species ofVeronica. Therefore, only taxonomic changes at the intraspecific level were necessary (Albach 2008).

Lamiaceae (M. S. Dillenberger)

Ballota L. contains c. 30 species that occur in Europe, the Mediterranean area, W Asia and, with one species, S Africa (Mabberley 2008). Several species ofBallota were included in a phylogenetic analysis of subfam.Lamioideae (Bendiksby & al. 2011b). This phylogeny unambiguously showed thatBallota is not monophyletic. The type ofBallota, B. nigra L. (the only species of the genus in Germany), is well-supported sister toMarrubium L., represented in the German flora byM. peregrinum L. andM. vulgare L. The otherBallota species are sister to thisB. nigra—Marrubium clade. Only two otherBallota species,B. frutescens (L.) Woods andB. integrifolia Benth., form a separate clade that is sister to the former clade and a clade containing species ofMoluccella L.,Otostegia Benth. andSulaimania Hedge & Rech. f. There are two solutions to obtain a monophyleticBallota. The first is to merge all species of this clade (i.e.Ballota, Marrubium, Moluccella, Otostegia andSulaimania) in one genus. The second solution is to excludeB. frutescens andB. integrifolia fromBallota and to combine the rest ofBallota includingB. nigra andMarrubium in one genus. In order to avoid creation of one very large and heterogeneous genus, it seems reasonable to take the second approach. Since both genera were described by Linnaeus (1753b), it remains unclear at this point which genus name should be used.Marrubium contains 40 species (Mabberley 2008), so that a comparable number of new combinations would be needed when using either name.

The treatment and circumscription ofClinopodium L. is very different in different Floras of Germany (e.g.Jäger & Werner 2005;Jäger 2011;Seybold 2011).Clinopodium in its broad circumscription, includingAcinos Mill.,Bancroftia Billb.,Calamintha Mill., New WorldMicromeria Benth. andSatureja L. contains c. 100 species and is almost cosmopolitan (Mabberley 2008). Seybold (2011) includedAcinos andCalamintha, but notSatureja, inClinopodium, and Jäger & Werner (2005) treatedAcinos, Calamintha, Clinopodium andSatureja as separate genera. A molecular phylogeny of subtribeMenthinae illustrates the whole dimension of the problem (Bräuchler & al. 2010). In this phylogeny,Clinopodium is highly polyphyletic and numerous genera are nested among differentClinopodium clades. The species ofAcinos form a well-supported clade together withZiziphora L., a genus of c. 20 species distributed from the Mediterranean area to C Asia, Afghanistan and Himalaya (Mabberley 2008). In this clade,Acinos andZiziphora are not supported as monophyletic.Calamintha species are in a well-supported clade with the type and other species ofClinopodium. Another genus that causes problems with respect to the monophyly ofClinopodium isMonarda L., a small genus of c. 16 mostly North American species (Mabberley 2008) occurring in Germany with one introduced species,M. didyma L. (Jäger & Werner 2005). The large number of genera, species and clades makes several solutions possible. For the German species only two solutions need to be discussed. The first is to include all species ofAcinos, Calamintha, Clinopodium andMonarda in one genus, together with the whole or parts ofAcanthomintha (A. Gray) A. Gray,Blephilia Raf.,Bystropogon L′Hér.,Conradina A. Gray,Cuminia Colla,Cunila D. Royen ex L.,Cyclotrichium Mandenova & Schengelia,Dicerandra Benth.,Glechon Spreng.,Hedeoma Pers.,Hesperozygis Epling,Hoehnea Epling,Killickia Bräuchler & al.,Mentha L., New WorldMicromeria Benth.,Minthostachys (Benth.) Spach,Monardella Benth.,Obtegomeria Doroszenko & P. D. Cantino,Piloblephis Raf.,Poliomintha A. Gray,Pycnanthemum Michx.,Rhododon Epling,Stachydeoma (Benth.) Small andZiziphora. Alternatively,Clinopodium can be split into clades which could be treated as morphologically recognizable genera. In view of substantial morphological variation of the lineages concerned it is not meaningful to merge so many genera only to preventClinopodium from being split. Although it is not clear how exactlyClinopodium will be split in the future, the impact of this approach on German species can easily be seen. SinceCalamintha is very closely related to the type ofClinopodium, and this relationship is well supported, there is no other solution than to transferCalamintha toClinopodium. TheCalamintha species in Germany,C. menthifolia Host andC. nepeta (L.) Savi, will have to become known asClinopodium menthifolium (Host) Stace andClinopodium nepeta (L.) Kuntze. No species name inClinopodium is available for the hybrid taxonCalamintha xfoliosa Opiz; at subspecies levelClinopodium nepeta nothosubsp.subisidoratum (Borbás) Govaerts has been used. It is not possible to treat the species ofAcinos as part ofClinopodium without including inClinopodium all genera listed above. Although the relationships betweenAcinos andZiziphora are not fully resolved, it seems necessary to combine these two genera in one genus.Ziziphora has priority overAcinos, and the species ofAcinos accordingly need new names inZiziphora. These are not yet available. The only GermanClinopodium species,C. vulgare L., is the type of the genus name and will therefore most likely not be affected by any changes of generic circumscriptions. The only genus of this group that seems to be unproblematic isSatureja. This genus, together withGontscharovia Boriss., is part of a polytomy with theClinopodium-clade (the numerous genera listed above) and a clade of Old WorldMicromeria (Bräuchler & al. 2010). Even ifSatureja is paraphyletic in relation toGontscharovia, Satureja has priority overGontscharovia and no taxonomic changes will be necessary in the German flora.

A long-discussed problem is the correct placement and naming of species belonging toGaleobdolon Adans. /Lamiastrum Heist, ex Fabr. (Dandy 1967;Holub 1970;Rauschert 1974;Mennema 1989;Krawczyk & al. 2013). Choice of genus name is a nomenclatural problem, which will not be dicussed here. In recent Floras of or covering Germany, either both names were used:Galeobdolon (Jäger & Werner 2005;Jäger 2011) andLamiastrum (Heywood & Richardson 1972;Seybold 2009), or the species ofGaleobdolon /Lamiastrum were included inLamium L. (Seybold 2011). Molecular phylogenies of subfam.Lamioideae (Bendiksby & al. 2011b) and ofLamium (including species ofGaleobdolon /Lamiastrum;Bendiksby & al. 2011a) clearly showed that a well-supported clade of species ofGaleobdolon /Lamiastrum is sister to a well-supportedLamium. Accordingly, both inclusion ofGaleodolon /Lamiastrum inLamium and treatment as two distinct genera would result in monophyletic genera. When included inLamium, G. argentatum Smejkal,G. flavidum (F. Herm.) Holub,G. luteum Huds. andG. montanum (Pers.) Pers. ex Rchb. should beL. argentatum (Smejkal) Henker ex G. H. Loos,L. flavidum F. Herm.,L. galeobdolon (L.) L. andL. montanum (Pers.) Hoffm. ex Kabath, respectively. The treatment of these four taxa at species level has been questioned. When treated as subspecies ofLamium galeobdolon (e.g. byBendiksby & al. 2011a), the names to be used would beL. galeobdolon subsp.argentatum (Smejkal) J. Duvign.,L. galeobdolon subsp.flavidum (F. Herm.) Á. Löve & D. Löve,L. galeobdolon subsp.galeobdolon andL. galeobdolon subsp.montanum (Pers.) Hayek, respectively.

Majorana Mill, andOriganum L. are two genera containing commonly used spices.Origanum is distributed in Eurasia and contains c. 38 species (Mabberley 2008).Majorana hortensis Moench was first described asO. majorana L. A phylogenetic analysis by Katsiotis & al. (2009) showed thatM. hortensis is nested among other species ofOriganum, so that recognition ofM. hortensis would makeOriganum paraphyletic. Therefore the inclusion ofM. hortensis inOriganum, asO. majorana, is appropriate.

Salvia L. in its traditional circumscription is a large genus of 800–900 tropical to temperate species (Mabberley 2008). Recent molecular studies in the genus clearly showed thatSalvia is highly polyphyletic (Walker & Sytsma 2007;Will & Claßen-Bockhoff 2014) and will have to be split into several genera (Will & Claßen-Bockhoff 2014; M. Will pers. comm.). The only alternative would be to include several smaller genera inSalvia, e.g.Rosmarinus L., which would inflate this large genus even more. The German flora is largely unaffected by these changes. According to the different phylogenies available, only one species of the German flora,S. glutinosa L., will have to be transferred to a new genus.

Stachys L. is a large genus of c. 450 species distributed in temperate and warm regions of the world, including tropical mountains but excluding Australasia (Mabberley 2008). Molecular phylogenies have shown thatStachys is highly polyphyletic, with many different genera nested among differentStachys clades (Bendiksby & al. 2011b;Salmaki & al. 2013). In Germany eight species and one hybrid taxon ofStachys can be found. These fall into four largerStachys clades (Salmaki & al. 2013). OnlyS. arvensis L. andS. palustris L. fall into the clade containing the type ofStachys, S. sylvatica L. This clade also containsHaplostachys Hillebr.,Phyllostegia Benth.,Stenogyne Benth. andSuzukia Kudô.Sideritis L. is one of those genera nested among differentStachys clades (Bendiksby & al. 2011b;Salmaki & al. 2013). This genus of c. 140 N-temperate species of the Old World and Macaronesia (Mabberley 2008) occurs in Germany with only one species,S. montana L. Relationships among the genera listed above and several others are complicated and not fully resolved. At this point two solutions seem possible. One is to include all species of a clade calledEurystachys Salmaki & Bendiksby (includingStachys andSideritis;Salmaki & al. 2013) in one genus. The other is to splitStachys into a large number of smaller genera. Both solutions are problematic. The first would result in a large genus that is morphologically heterogeneous and, according to Salmaki & al. (2013), c. 194 new combinations would have to be made. The second solution would allow maintaining morphologically distinct genera. However, it would require dividingStachys into several genera that would be difficult to delimit (Salmaki & al. 2013). In this second approach a similarly high number of combinations would be necessary. The first solution would allow keeping all German species ofStachys inStachys, which, however, would also have to includeSideritis. In the second solution,Stachys alpina L.,S. annua L.,S. byzantina K. Koch,S. germanica L. andS. recta L. most likely would need to be excluded fromStachys. The relationships ofSideritis are unresolved. It therefore remains unclear whetherSideritis montana would need a new name when opting for the second solution. A further difficulty of the second solution is a high level of incongruence between the nuclear and plastid data sets analysed (Salmaki & al. 2013). Future changes in this group are clearly necessary. They will affect large numbers of species on a global scale, but only few species of the German flora.

Orobanchaceae (D. C. Albach)

There has been some debate about the monophyly ofOrobanche L., and some publications re-used the namePhelipanche Pomel, introduced for some morphologically deviant species more commonly treated asO. sect.Trionychon Wallr. (in GermanyO. arenaria Borkh.,O. purpurea Jacq. andO. ramosa L.; see lead 1 in the key toOrobanche inJäger 2011). The group differs from the type section not only in flower morphology but also in seed ultrastructure (Plaza & al. 2004) and pollen morphology (Abu Sbaih & al. 1994). Whereas first cpDNA-based phylogenetic analyses suggestedO. ramosa to be nested in the rest ofOrobanche (Young & al. 1999;Manen & al. 2004), subsequent analyses using ITS (Schneeweiss & al. 2004) revealed a biphyleticOrobanche withO. sect.Trionychon and New World species of the genus forming a clade andO. sect.Orobanche sister toDiphelypaea Nicolson. More detailed analyses of cpDNA sequences demonstrated that the nested position ofO. sect.Trionychon is due to horizontal gene transfer (Park & al. 2007). However, analyses of another nuclear marker (PhyA;Bennett & Mathews 2006) as well as cpDNA analyses removing introgressed sequences (Park & al. 2008) agree on phylogenetic relationships withOrobanche being monophyletic andO. sect.Trionychon and New World species being sister toO. sect.Orobanche. Thus, no taxonomic changes will be necessary.

Linderniaceae (D. C. Albach)

Only two species ofLinderniaceae occur in Germany and are commonly still recognized underLindernia L., one being the type of the genus name,L. procumbens (Krock.) Borbás.Lindernia dubia (L.) Pennell, though, has been demonstrated to be more closely related toMicranthemum Michx. than toLindernia (Fischer & al. 2013). However, no generic realignment has been proposed so far.Lindernia dubia had been recognized as separate fromLindernia before under the namesGratiola dubia L. orIlysanthes riparia Raf., but was included inLindernia by Pennell (1935).Ilysanthes Raf. had been separated fromLindernia based on the reduction of the androecium to two stamens, which Pennell (1935) did not consider stable enough to merit generic rank. Fischer & al. (2013) seemed to favour inclusion ofMicranthemum inLindernia. However, sinceMicranthemum also has only two stamens and occurs sympatrically withL. dubia in North America, combiningL. dubia inMicranthemum remains a possibility.

Convolvulaceae (J. W. Kadereit)

A monophyleticCalystegia R. Br. is clearly nested withinConvolvulus L. (Stefanović & al. 2002;Carine & al. 2004;Williams & al. 2014) and should be classified inConvolvulus following Stefanović & al. (2002). Combinations are available for most German species ofCalystegia, andC. pulchra Brummitt & Heywood should beConvolvulus dubius J. L. Gilbert,C. sepium (L.) R. Br. should beConvolvulus sepium L.,C. silvatica (Kit.) Griseb. should beConvolvulus silvaticus Kit. andC. soldanella (L.) Roem. & Schult, should beConvolvulus soldanella L.

Solanaceae (J. W. Kadereit)

Lycopersicon Mill, is clearly nested inSolanum L. (Spooner & al. 1993) and should be treated in that genus. Accordingly, the tomato should be calledS. lycopersicum L.

Whitson & Manos (2005) demonstrated that the two species ofPhysalis L. listed for Germany,P. alkekengi L. andP peruviana L., fall into two distantly related clades ofPhysalinae. The authors argued: “To correct the paraphyly ofPhysalis, nomenclatural changes are required. Options include restricting the namePhysalis toP. alkekengi, the type, and renaming the 75+ species of New WorldPhysalis, or broadening the circumscription ofPhysalis by uniting the majority of thePhysalinae into a single genus. However, the least taxonomically disruptive approach for dealing with this problem is to re-typifyPhysalis using a Linnaean species that is a member of the morphologically typicalRydbergis clade, such asP. pubescens. The atypical species could then be recognized as four small genera (forP. carpenteri, P. alkekengi, P. microphysa, and subgenusPhysalodendron), which would produce a morphologically homogeneousPhysalis. A proposal to re-typifyPhysalis is currently in progress.” This proposal has been made by Whitson (2011), and conservation ofPhysalis L. with conserved type has been recommended (Applequist 2012). If accepted,P. alkekengi should be known asAlkekengi officinarum Moench.

Boraginaceae (M. Weigend)

Generic limits in the large familyBoraginaceae (1500— 1600 spp.) are highly problematic and numerous re-alignments of generic limits are required, but few of these problems concern the German flora. The genusOmphalodes Moench is represented only by two species in Germany:O. scorpioides (Haenke) Schrank andO. verna Moench.Omphalodes scorpioides has been shown to be more closely related toMertensia Roth than to the typical representatives ofOmphalodes (Weigend & al. 2013), and is now accommodated in the monospecific genusMemoremea A. Otero & al. asMemoremea scorpioides (Haenke) A. Otero & al. (Otero & al. 2014). This is clearly supported by molecular data, but also by gross differences in habit and its aberrant fruit morphology (circular wing of the nutlet forming a hollow ring, not a flat appendage).Omphalodes in the narrowest sense is restricted to those perennial, rhizomatous herbs which are closely related toO. verna, the type of the genus name. This group ranges from N Spain to N Iran. Other groups from Asia and the Americas previously assigned toOmphalodes either have already been segregated from the genus (Otero & al. 2014) or will likely be removed to other genera.

The genusBuglossoides Moench is also represented by only two species in Germany.Buglossoides arvensis (L.) I. M. Johnst., an annual weed with tiny white flowers and four triangular-ovate, verrucose nutlets andB. purpurocaerulea (L.) I. M. Johnst., a perennial herb with large, blue, hypocrateriform flowers and single, smooth, spherical nutlets. They represent the C European representatives of two highly natural and monophyletic species groups, which are retrieved as sister groups in molecular studies (Weigend & al. 2009;Cecchi & al. 2014). These species groups have recently been segregated into two different, easily distinguished genera (Cecchi & al. 2014):Buglossoides s.str., essentially comprising the two speciesB. arvensis andB. incrassata and largely restricted to the circum-Mediterranean region and Europe (and introduced as weeds elsewhere), andAegonychon Gray with a total of three species, one narrow S Italian endemic and the widespreadA. purpurocaeruleum (L.) Holub, in W Eurasia as sister to the morphologically barely distinguishable Japanese endemicA. zollingeri (A. DC.) Holub (Cecchi & al. 2014). The clear morphological differences between these two groups justify their separation into two well-defined genera, but phylogenetic data would equally permit a broader delimitation ofBuglossoides, includingAegonychon.

The generaEritrichium Schrad. ex Gaudin,Hackelia Opiz andLappula Moench have a confused taxonomic history, butHackelia was finally segregated fromLappula by Johnston (1923). The only C European species ofHackelia andLappula and the types of those names,H. deflexa (Wahlenb.) Opiz andL. squarrosa (Retz.) Dumort., have recently often been treated as belonging to a single genus, i.e.Lappula. Recent molecular studies retrieved these two species in widely separate clades in tribeEritrichieae, together with the bulk of the species currently assigned to the respective genera. There is therefore both morphological (Johnston 1923) and molecular (Weigend & al. 2013) evidence supporting the recognition of the two genera. The exact limits betweenEritrichium andHackelia andLappula still require additional work, with several extra-European segregate genera apparently nested in them, and some species incorrectly placed. This, however, does not concern the German or European flora.

The delimitation ofAnchusa L., characterized by radially symmetrical flowers, fromLycopsis L. with curved, slightly zygomorphic flowers, has been contentious in the past. Morphological differences are small but striking, and the segregateLycopsis is currently not generally recognized. Hilger & al. (2004) advocated the subdivision ofAnchusa into several smaller genera, including the separation ofLycopsis. However, their molecular data failed to retrieve the two species ofLycopsis as monophyletic, and there was no statistical support forAnchusa excludingLycopsis. Generic limits inAnchusa s.1. clearly require more work, and it seems more sensible at this stage to recognize a single, more widely defined genusAnchusa until much better data are available.

Two other genera represented in Germany will likely be subject to re-definition in the near future, without affecting the taxonomy of German species: Both species ofCynoglossum L. were retrieved in the core-clade ofCynoglossum s.1. (Weigend & al. 2013) and certainly will remain part of a redefinedCynoglossum. However,Cynoglossum likely will have to include a whole range of W Eurasian segregate genera (Hilger & al. 2015). Similarly,Heliotropium europaeum L. is the type ofHeliotropium L. and therefore will not be affected by name change, irrespective of how the limits ofHeliotropium, with the large genusTournefortia L. deeply nested in it (Luebert & al. 2011), will ultimately be redefined.

Apiaceae (K. Spalik)

Hacquetia DC. is nested withinSanicula L. (Valiejo-Roman & al. 2002;Calvifio & Downie 2007) and should therefore be sunk into synonymy; for its only species,H. epipactis (Scop.) DC., the nameS. epipactis (Scop.) E. H. L. Krause is available.

Apium L. s.l. is polyphyletic and among its European species only the type, A.graveolens L., is retained in the genus; the other true celeries are distributed throughout the S hemisphere (Spalik & al. 2010). For its other European members, the genusHelosciadium W. D. J. Koch has been reinstated (Hardway & al. 2004;Spalik & al. 2009;Ronse & al. 2010) includingH. inundatum (L.) W. D. J. Koch (A. inundatum (L.) Rchb. f.),H. nodiflorum (L.) W. D. J. Koch (A. nodiflorum (L.) Lag.) andH. repens (Jacq.) W. D. J. Koch (A. repens (Jacq.) Lag.). The species ofHelosciadium are hydrophytes or helophytes and are closely related to the morphologically and ecologically similarBerula W. D. J. Koch andSium L., members of tribeOenantheae (Spalik & al. 2014).

Carum L., the type ofCareae, includes c. 30 species that in molecular analyses are located in several disparate clades interspersed with species ofChamaesciadium C. A. Mey.,Fuernrohria K. Koch andGrammosciadium DC., with only few species closely related to the type ofCarum, C. carvi L. (Zakharova & al. 2012).Carum verticillatum (L.) W. D. J. Koch is a very distant relative of its nominative congeners and, therefore, was placed in the reinstated monospecific genusTrocdaris Raf.; its proper name isT. verticillata (L.) Raf. (Zakharova & al. 2012). This species forms an isolated lineage in a clade of hydrophytic umbellifers constituting tribeOenantheae (Spalik & al. 2014).

The generaAngelica L.,Cnidium Cusson,Libanotis Haller ex Zinn,Peucedanum L.,Selinum L.,Seseli L. andTrinia Hoffm. are part of the taxonomically difficult tribeSelineae (Spalik & al. 2004;Downie & al. 2010). Many of its genera are polyphyletic while at the same time many monophyletic lineages have unnecessarily been split into small segregates. Numerous species have not yet been included in molecular phylogenetic studies, and the generic boundaries remain unclear. Phylogenetic relationships within this tribe were mostly examined using only nuclear ITS sequences that have some limitations. Moreover, the tribe originated relatively recently, c. 12 Mya, and underwent rapid radiation (Banasiak & al. 2013: Appendix S2). In effect, internal branches of the phylogenetic trees obtained from molecular data are short and often poorly supported, precluding unambiguous taxonomic inferences.

Seseli sensu amplo encompasses 100–120 species and is obviously polyphyletic: its species occur in tribesApieae, Pimpinelleae andSelineae (Downie & al. 2010), and inSelineae they are placed in several clades (Spalik & al. 2004).Seseli hippomarathrum Jacq. together with three other congeners forms a clade that is not most closely related toS. tortuosum L., the type of the genus name; for this group, a restitution ofHippomarathrum G. Gaertn. & al. has been considered (Spalik & al. 2004). Depending on taxonomic sampling and the method of phylogenetic inference, this clade was placed sister to theSeseli clade (Spalik & al. 2004) or sister toPeucedanum s.l. (see Appendix S2 inBanasiak & al. 2013). Detailed molecular and morphological studies are necessary to elucidate the taxonomic status of this group. Upon restitution ofHippomarathrum the nameH. pelviforme G. Gaertn. & al. would be available forS. hippomarathrum. Seseli annuum L. has not yet been included in molecular analyses; therefore, its phylogenetic affinities remain unknown.

Libanotis pyrenaica (L.) Bourg eau is closely related toL. montana Crantz, the type ofLibanotis, and inFlora iberica (Aedo & Vargas 2003) the former was synonymized with the latter. In molecular analyses, the clade containing these two species is sister to a clade containing the type ofSeseli (Spalik & al. 2004;Banasiak & al. 2013). If a broad definition ofSeseli is adopted, e.g. based on theSeseli clade in Spalik & al. (2004), thenLibanotis should be sunk intoSeseli and the species is to be namedS. libanotis (L.) W. D. J. Koch.

Ligusticum mutellinoides (Crantz) Vill. (Pachypleurummutellinoides (Crantz) Holub) is also closely related to theLibanotis—Seseli clade in tribeSelineae, whereas the types ofLigusticum L. andPachypleurum Ledeb. are placed in theAcronema clade, which deserves rank as a separate tribe (Downie & al. 2010;Banasiak & al. 2013). Depending on the delineation ofLibanotis andSeseli, Ligusticum mutellinoides may be included in either of these two genera. Alternatively,Neogaya Meisn. may be reinstated. Its type isN. simplex (L.) Meisn., a taxonomic synonym ofL. mutellinoides. In molecular phylogenetic trees,Ligusticum mutellina (L.) Crantz is placed in theConioselinum chinense clade far from the type ofLigusticum and, therefore, should be excluded from the genus and placed in the reinstatedMutellina Wolf, asM. purpurea (Poir.) Reduron & al. (Valiejo-Roman & al. 2006).

Cnidium dubium (Schkuhr) Schmeil & Fitschen is not most closely related to the type ofCnidium, C. monnieri (L.) Spreng., and should therefore be recognized asKadenia dubia (Schkuhr) Lavrova & V. N. Tikhom. (Valiejo-Roman & al. 2006).

Trinia is exceptional inApiaceae due to its dioecious breeding system, and this feature seems to be synapomorphic for the genus. So far, onlyT. hispida Hoffm. has been included in molecular phylogenetic analyses and it was placed in theSeseli clade very close to the type ofSeseli (Spalik & al. 2004). If this placement is confirmed upon extended sampling of species and molecular markers, then eitherTrinia is to be included into the synonymy ofSeseli or the latter is to be restricted to a clade of only a few closest relatives of its type.

Peucedanum sensu amplo includes c. 100–120 species worldwide and is a “dustbin” genus encompassing taxa that do not fit elsewhere. The European species have often been transferred to small segregate genera includingCervaria Wolf,Dichoropetalum Fenzl (=Holandrea Reduron & al.),Imperatoria L.,Oreoselinum Mill.,Thysselinum Adans. andXanthoselinum Schur. Of these, however, onlyCervaria andDichoropetalum are unambiguously supported by molecular data because their types are distant relatives ofPeucedanum officinale L., the type of the genus name. The remaining segregates form thePeucedanum s.l. clade that can be retained as one genus (Spalik & al. 2004). Therefore, the use of the namesDichoropetalum carvifolia (Vill.) Pimenov & Kljuykov (P. carvifolia Vill.) andCervaria rivini Gaertn. (P. cervaria (L.) Lapeyr.) is advocated. If a very narrow definition ofPeucedanum is adopted, the namesImperatoria ostruthium L. (P. ostruthium (L.) W. D. J. Koch),Oreoselinum nigrum Delarbre (P. oreoselinum (L.) Moench),Thysselinum palustre (L.) Hoffm. (P. palustre (L.) Moench) andXanthoselinum alsaticum (L.) Schur (P. alsaticum L.) are available for the respective species ofPeucedanum.

Molecular data have demonstrated thatLaserpitium L. is polyphyletic (Weitzel & al. 2014;Lyskov & al. 2015), and this polyphyly is strongly supported by nrDNA and cpDNA markers (Banasiak & al. in press). Six closely related species including the type,L. gallicum L., as well asL. latifolium L. constituteLaserpitium s.str.Laserpitium siler L. forms an isolated lineage that is not closely related to the type and, therefore, the restitution of the monospecificSiler Crantz has been postulated; the respective name forL. siler isS. montanum Crantz.Laserpitium prutenicum L. is more closely related toDaucus L. than toLaserpitium s.str. and, together with its closest relative,L. hispidum M. Bieb., it deserves to be placed in a new genus,Silphiodaucus (Koso-Pol.) Spalik & al. (Banasiak & al. in press). The respective name forL. prutenicum would beS. prutenicus (L.) Spalik & al.

Dipsacaceae (J. W. Kadereit)

Virga Hill, withV. pilosa (L.) Hill andV. strigosa (Roem. & Schult.) Holub clearly groups inDipsacus L. (Avino & al. 2009;Carlson & al. 2009) and these two species should be known asDipsacus pilosus L. andD. strigosus Roem. & Schult., respectively.

Valerianaceae (J. W. Kadereit)

NeitherValerianella Mill, norValeriana L. are monophyletic according to Hidalgo & al. (2004) and Bell & Donoghue (2005). However, inclusion ofFedia Gaertn. emend. Moench inValerianella, and both inclusion ofPlectritis (Lindl.) DC. inValeriana and exclusion of some species ofValeriana could make the two genera monophyletic.

Campanulaceae (N. Kilian)

The two German species ofLobelia L.,L. dortmanna L. andL. erinus L., fall into two different clades of a highly paraphyleticLobelia (Antonelli 2008). If this should result in splitting ofLobelia, an approach considered premature by Lammers (2011),L. erinus would belong to a different genus.

Wahlenbergia Roth has been shown to be polyphyletic (Haberle & al. 2009;Roquet & al. 2009;Prebble & al. 2012;Cupido & al. 2013).Wahlenbergia hederacea (L.) Rchb., the only species present in Germany, is not only misplaced inWahlenbergia (typified byW. elongata (Willd.) Schrad., a synonym of the S AfricanW. capensis (L.) A. DC.;Lammers 2007) but also in the otherwise monophyletic wahlenbergioid group of genera (Cupido & al. 2013). It appears instead to be a close relative ofFeeria Buser andJasione L. (Prebble & al. 2012;Cupido & al. 2013;Crowl & al. 2014; but not so inMansion & al. 2012), but its systematic position still needs clarification.

A number of molecular phylogenetic studies ofCampanulaceae (Eddie & al. 2003;Park & al. 2006;Roquet & al. 2008,2009;Borsch & al. 2009;Haberle & al. 2009;Mansion & al. 2012;Crowl & al. 2014) have shown thatCampanula L. in its present circumscription is not monophyletic, and that the species of this genus fall into at least four major clades, each containing other genera of the family. Referring, with a view on the German flora, to the analysis based on the most comprehensive sampling by Mansion & al. (2012), which also provides the best resolution so far, the three largest major clades are relevant. These are: (1) theCampanula s.str. clade (Park & al. 2006;Roquet & al. 2008,2009;Borsch & al. 2009;Mansion & al. 2012), including the type of the genus name,C. latifolia L., and comprising clades 13–17 in Mansion & al. (2012), contains the majority of theCampanula species in Germany (C. alliariifolia Willd.,C. alpina Jacq.,C. barbata L.,C. bononiensis L.,C. cervicaria L.,C. glomerata L.,C. latifolia, C. medium L.,C. rapunculoides L.,C. sibirica L.,C. thyrsoides L. andC. trachelium L., all nested in clade 17). TheCampanula s.str. clade also includes the species of the S EuropeanTrachelium L., but the different analyses demonstrate that this genus does not constitute a natural group but is found dismembered in clades 13 and 16 in Mansion & al. (2012). (2) The Rapunculus clade (clades 5–12 inMansion & al. 2012) includes all but one of the remaining species in Germany (C. baumgartenii Becker,C. cochleariifolia Lam.,C. rhomboidalis L.,C. rotundifolia L. [incl.C. gentilis Kovanda] andC. scheuchzeri Vill. in clade 12;C. patula L. andC. rapunculus L. in clade 9) and also contains (in clade 6) the generaAdenophora Fisch, andHanabusaya Nakai. (3) The third major clade, which has low support, comprises the well-supported clades 2–4 in Mansion & al. (2012), in which severalCampanula lineages (among them the last German memberC. persicifolia L. in clade 3) are mixed withAsyneuma Griseb. & Schenk,Legousia Durande andPhyteuma L. as well as with the American generaGithopsis Nutt.,Heterocodon Nutt, andTriodanis Raf. Faced with different classificatory options, i.e. (1) treating all clades containing species ofCampanula as one genus, (2) limitingCampanula to theCampanula s.str. clade, and (3) splittingCampanula into numerous small genera, an option briefly discussed by Park & al. (2006), Roquet & al. (2008) concluded: “We favor the first option in order to arrive at a generic delimitation that reflects the evolutionary history ofCampanula. This approach is more consistent with previous taxonomic work,Campanula has always been very rich in number of species, and it does not seem to us reasonable to divide it ad nauseam. … However, a comprehensive study of the currently recognized genera that fall withinCampanula should be conducted before changing their taxonomic status.” If this approach would be taken, all species ofAdenophora, Legousia andPhyteuma would have to be treated asCampanula.

Menyanthaceae (J. W. Kadereit)

AlthoughNymphoides Ség. was found to be non-monophyletic, with one species more closely related to one clade of a non-monophyleticVillarsia Vent, than to the remaining species ofNymphoides (Tippery & al. 2008),N. peltata (S. G. Gmel.) Kuntze will not change name even when combined withVillarsia becauseNymphoides is the older name.

Cardueae (A. Susanna & N. Garcia-Jacas)

Extensive molecular analyses in subtribeCentaureinae have demonstrated thatCentaurea L., as defined in classic terms, was a polyphyletic assemblage (Susanna & al. 1995;Garcia-Jacas & al. 2001). As regards naming of the two main lineages, problems originated with an inadequate type of the genus name proposed by Britton & Brown (1913), a decision later ratified by Dittrich (1993):C. centaurium L. This species belongs to a group of some 20–25 taxa that are not closest relative of the largest part of the genus. Two alternate solutions were possible for achieving a natural delineation of the two genera that should be recognized: first, to keep the old type and limit the use ofCentaurea to this group of species, which would imply renaming more than 200 species in a different genus; second, to conserve a new type belonging to the main group of the genus. This second, more conservative (in terms of botanical nomenclature) option finally prevailed: a new type,C. paniculata L., was proposed by Greuter & al. (2001) and is now the conserved type ofCentaurea (Wiersema & al. 2015). The valid name for the genus comprising the smaller group of species isRhaponticoides Vaill. This change, in Germany, affects onlyC. ruthenica Lam., which should be known asR. ruthenica (Lam.) M. V. Agab. & Greuter. As for the segregation ofC. sect.Cyanus (Mill.) DC. as a separate genus (e.g.Greuter & al. 2001), molecular evidence, although inconclusive, points at a sister relationship ofC. sect.Cyanus andC. sect.Centaurea (e.g.Garcia-Jacas & al. 2001). The latest proposal for a classification of the entire genusCentaurea (Hilpold & al. 2014) and the revisions of tribeCardueae by Susanna and Garcia-Jacas (2007,2009) do not acceptCyanus Mill, as genetically different fromCentaurea.

Cichorieae (N. Kilian)

Lapsana L., together with the equally epappose MediterraneanRhagadiolus Juss., is nested inCrepis L., as has been shown in nuclear ribosomal (ITS) and chloroplast (matK) DNA marker phylogenies by Enke & Gemeinholzer (2008). To maintainLapsana, which is monospecific after the well-supported segregation of the E AsianLapsanastrum Pak & K. Bremer (Pak & Bremer 1995;Deng & al. 2014) and the dispecificRhagadiolus as separate monophyletic genera,Crepis would have to be split into two morphologically ill-defined entities. This is definitely no practicable solution. If only monophyletic genera should be accepted, merging of both genera withCrepis would be the more appropriate solution, although breaking with a long tradition (no combination ofLapsana inCrepis has been published). The morphological circumscription ofCrepis does not, however, preclude the inclusion ofLapsana communis L. (and ofRhagadiolus) if variation is extended to allow for the absence of a pappus. In other subtribes, parallel cases of epappose entities traditionally treated as separate genera are similarly found nested in regularly pappose genera (e.g.Deng & al. 2014).

The members ofHypochaeris L. cluster in two main clades according to the phylogenetic analyses of nuclear ribosomal (ITS) and several chloroplast DNA marker sequences by Samuel & al. (2003,2006) and Enke & al. (2012). The results, however, are inconclusive as to whether the two clades actually form a sister group and thus to the monophyly ofHypochaeris. Based on these findings, Talavera & al. (2015a) opted for splitting the genus in the forthcoming treatment ofFlora iberica, there recognizing the segregatesAchyrophorus Adans. (in its narrow sense distributed in the Mediterranean region) andTrommsdorffia Bernh. (withT. maculata (L.) Bernh. [H.maculata L.] andT. uniflora (Vill.) Soják [H. uniflora Vill.] in the German flora), a solution that necessitates recognition of at least a fourth genus for the NW African-South American clade ofHypochaeris s.l.

Leontodon L. in its traditional circumscription is at least diphyletic (Samuel & al. 2006;Enke & al. 2012).Leontodon subg.Leontodon andL. subg.Oporinia (D. Don) Peterm., which both received strong support in molecular phylogenies, are nested in two different major clades of the subtribe. This finding from phylogenetic analyses based on both nuclear ribosomal (ITS) and chloroplast (matK) DNA marker sequences necessitates the recognition ofL. subg.Oporinia (includingL. autumnalis L.,L. helveticus Mérat andL. montanus Lam. in Germany) as a separate genus,Scorzoneroides Moench, withS. autumnalis (L.) Moench as type (Greuter & al. 2006) andS. helvetica (Mérat) Holub andS. montana (Lam.) Holub as additional species in the German flora. The authorship ofScorzoneroides should be attributed to Moench (Meth.: 549. 1794), because an earlier place of publication of that name and other genus names (in a German translation dated 1754–1756 of a pre-Linnaean work by Vaillant) is expected to be added to the list of suppressed works by the next International Botanical Congress (Applequist 2014: 1370).Leontodon s.str., moreover, is paraphyletic with respect to the small, chiefly Mediterranean genusHedypnois Mill., not present in the flora of Germany (Enke & al. 2012). The nrlTS phylogeny by Samuel & al. (2006) and Enke & al. (2012) also provide initial indication (without statistical support, and not supported by thematK phylogeny) thatL. sect.Thrincia (Roth) Benth. (with onlyL. saxatilis Lam.[Thrincia saxatilis (Lam.) Holub & Moravec] in the German flora) forms a clade not sister to the remainder ofLeontodon s.str. Based on these findings, Talavera & al. (2015b) revived the genusThrincia Roth for this clade.

Picris L. is monophyletic after exclusion of the small Mediterranean-SW Asian genusHelminthotheca Zinn (Samuel & al. 2006;Enke & al. 2012). Its segregation has previously been concluded for morphological reasons by Lack (1975). The only species of the latter genus in Germany isII. echioides (L.) Holub (Picris echioides L.), which also provides the type of the genus name.

Scorzonera L. is polyphyletic in all current circumscriptions according to the initial molecular phylogenetic investigations in the subtribe by Mavrodiev & al. (2004) and Owen & al. (2006), using nuclear ribosomal (ITS and ETS) DNA markers and Amplified Fragment Length Polymorphisms (AFLP) variation, respectively. According to these analyses, the clade ofScorzonera s.str. (including the type of the name,S. humilis L., as well asS. purpurea L.) is sister to a clade comprisingPodospermum DC. (of which the only member in the German flora,P. laciniatum (L.) DC. [S. laciniata L.], provides the type of that name). The other members ofScorzonera in its wider circumscription, as far as included in the analyses, are distributed over at least three further clades. Two of them, which form a clade sister to the clade comprisingKoelpinia Pall, and thePodospermum andScorzonera s.str. clades (Owen & al. 2006), each include one species in the German flora: S.austriaca L. andS. hispanica L. The third clade is the“Lasiospora clade” (includingS. hirsuta L., the type ofLasiospora Cass.), which is sister to all other lineages of the subtribe but has no representatives in the German flora. Apart from the segregation ofPodospermum DC. fromScorzonera s.str., which is supported as an option (but not a necessity), the current state of our knowledge ofScorzonera s.l. is still far too preliminary to draw taxonomic conclusions.

Sonchus L. has turned out to be paraphyletic with respect to various smaller Mediterranean-Macaronesian and Australian-New Zealand segregates as well as to the SE Pacific Ocean island endemicsDendroseris D. Don andThamnoseris Phil, in a series of molecular phylogenetic analyses based on both nuclear ribosomal and chloroplast DNA markers (Kim & al. 2007 and references therein). The preferred and envisaged taxonomic solution is the broadening of the generic concept forSonchus and (re)inclusion of all these genera (Mejias & Kim 2012). A splitting approach would inevitably dismember even the four German representatives of the genus, the congenerity of which has never been questioned.

The systematics of theLactuca alliance, which is represented in the German flora by the generaCicerbita Wallr.,Mycelis Cass, andLactuca L., has been in lively debate for more than 200 years. The first molecular phylogenetic analyses published (Koopman & al. 1998;Wang & al. 2013) explained the difficulties in arriving at a natural classification with frequent convergent evolution of morphological characters. Consequences for the generic classification of the species in Germany are to be expected, but phylogenetic reconstruction is still in progress and any reclassification would be premature at present.

Prenanthes L. has been redefined completely on the basis of molecular phylogenetics, now being understood as a probably monospecific genus, accommodating the chiefly EuropeanP. purpurea L. (Kilian & Gemeinholzer 2007;Kilian & al. 2009;Wang & al. 2013).

The placement of the C and SE EuropeanTolpis staticifolia (All.) Sch. Bip., the only representative ofTolpis L. in the flora of Germany, is not settled yet.Tolpis staticifolia and the S and tropical AfricanT. capensis (L.) Sch. Bip. (plus its close allyT. mbalensis G. V. Pope) have been excluded from that chiefly Mediterranean-Macaronesian genus based on palynological differences (Blackmore & Jarvis 1986) and on the results of a chloroplastndhF sequence phylogeny by Park & al. (2001), which placed the two species as sister toTaraxacum F. H. Wigg. (T. capensis) andCrepis (T. staticifolia), respectively.

Recent molecular phylogenetic analyses of theHieracium alliance using nuclear ribosomal, low-copy nuclear and chloroplast DNA markers (Fehrer & al. 2007,2009;Krak & al. 2013) revealed conflicting topologies between the different gene trees in particular due to both reticulate evolution and incomplete lineage sorting during the rapid evolution of the alliance. Discussing the available evidence, the authors concluded that the nuclear ribosomal DNA gene trees provide the best approximation for the reconstruction of the species tree. Accordingly,Hieracium L. in the wide sense is polyphyletic.Hieracium subg.Pilosella (Hill.) Fr. is sister to the W Mediterranean genusHispidella Lam., both are sister toH. subg.Hieracium and the AmericanH. subg.Chionoracium Sch. Bip. (=Stenotheca Monnier), the four taxa in turn are sister to the chiefly Mediterranean-Macaronesian genusAndryala L., and, finally,H. intybaceum All., which is restricted to the siliceous Alps, forms the sister group to all of them. The taxonomic consequences already widely drawn are the recognition as separate genera ofHieracium andPilosella Hill (for taxonomy and new combinations needed seeBräutigam & Greuter 2007; for the authorship ofPilosella the above notes onScorzoneroides also apply). The further consequence in order to arrive at monophyletic entities is the resurrection of the genusSchlagintweitia Griseb. to accommodateH. intybaceum (asS. intybacea (All.) Griseb.) and its few allies (Gottschlich & Greuter 2007;Greuter & Raab-Straube 2008).

Senecioneae (J. W. Kadereit)

Phylogenetic analyses ofSenecioneae (Pelser & al. 2002,2007,2010) have shown thatSenecio L. in its traditional circumscription is not monophyletic but rather both poly and paraphyletic. As regards species in the German flora, it is evident that those species that lack outer involucral bracts, i.e.S. congestus (R. Br.) DC.,S. gaudinii Gremli,S. helenites (L.) Schinz & Thell.,S. integrifolius (L.) Clairv. andS. rivularis (Waldst. & Kit.) DC., need to be segregated asTephroseris (Rchb.) Rchb., in which they are known asT. palustris (L.) Rchb. (forS. congestus), T. tenuifolia (Gaudin) Holub (forS. gaudinii), T. helenites (L.) B. Nord, (forS. helenites), T. integrifolia (L.) Holub (forS. integrifolius) andT. crispa (Jacq.) Rchb. (forS. rivularis).Tephroseris is only very distantly related toSenecio s.str. and even belongs to a different subtribe ofSenecioneae.

Species related toSenecio jacobaea L. should be segregated asJacobaea Mill., which again is only distantly related toSenecio s.str. These, besidesS. jacobaea (J. vulgaris Gaertn.), includeS. abrotanifolius L. (J. abrotanifolia (L.) Moench),S. alpinus (L.) Scop. (J. alpina (L.) Moench),S. aquaticus Hill (J. aquatica (Hill) G. Gaertn. & al.),S. erraticus Bertol. (J. erratica (Bertol.) Fourr.),S. erucifolius L. (J. erucifolia (L.) G. Gaertn. & al.),S. incanus subsp.carniolicus (Willd.) Braun-Blanq. (J. incana subsp.carniolica (Willd.) B. Nord.; for a recent account of theS. carniolicus aggregate seeFlatscher & al. 2015),S. paludosus L. (J. paludosa (L.) G. Gaertn. & al.) andS. subalpinus Koch. (J. subalpina (W. D. J. Koch) Pelser & Veldkamp). Combinations inJacobaea are available for all these species (Pelser & al. 2006).

Endocellion Turcz. ex Herder, containing two species in Asia, is clearly nested inPetasites Mill. (Steffen 2013) and should be treated as part of that genus. This does not affect the generic identity of thePetasites species in Germany.

Gnaphalieae (M. Galbany-Casals)

Phylogenetic analyses and morphological data show thatFilago L. is not monophyletic, and that the species involved should now be placed in two separate genera not closely related to each other (Galbany-Casals & al. 2010;Andrés-Sánchez & al. 2011):Logfia Cass, includesL. minima (Sm.) Dumort. (F. minima (Sm.) Pers.) andL. gallica (L.) Coss. & Germ. (F. gallica L.), andFilago includes the rest of the species present in Germany.Filago neglecta (Soyer-Willemet) DC. has been claimed to be of hybrid origin betweenL. gallica andGnaphalium uliginosum L. (Holub 1976;Jäger 2011), but this is currently considered highly doubtful (Andrés-Sánchez, pers. comm.). However, it is not clear yet if this rarely collected species belongs toFilago orLogfia.

Bombycilaena erecta (L.) Smoljan. has not been treated in Jäger (2005,2011), but there exists at least one old record of this species from Germany (Andrés-Sánchez & al. 2014). The genusBombycilaena (DC.) Smoljan. has been shown to be a lineage separate fromMicropus L. andFilago in a molecular phylogeny and is currently considered to include only two species from the Old World (Galbany-Casals & al. 2010;Andrés-Sánchez & al. 2014).

Omalotheca Cass, (sensuHolub 1976) has often been considered a synonym ofGnaphalium L. (e.g.Anderberg 1991;Jäger 2005,2011). However, a molecular phylogeny (Galbany-Casals & al. 2010) has shown thatGnaphalium s.l. is not monophyletic and that these two genera should be considered separate, given thatG. supinum L. — the type ofOmalotheca — is not closely related toG. uliginosum — the type ofGnaphalium. Additionally, Blöch & al. (2010) showed thatG. hoppeanum W. D. J. Koch,G. norvegium Gunnerus andG. sylvaticum L., three species also present in Germany, form a clade withG. supinum. In conclusion, with regard to the German flora,Gnaphalium should be restricted toG. uliginosum, and the other four species named above should be considered to belong toOmalotheca, asO. hoppeana (W. D. J. Koch) Sch. Bip. & F. W. Schultz,O. norvegica (Gunnerus.) Sch. Bip. & F. W. Schultz,O. supina (L.) DC. andO. sylvatica (L.) Sch. Bip. & F. W. Schultz. Smissen & al. (2011) noted thatGnaphalium s.str. includes diploid species (2n = 14), whereasOmalotheca species are all polyploids, and that the latter genus is part of a large clade of ancient allopolyploid origin, together with, among others, genera such asAntennaria Gaertn,Bombycilaena, Filago, Gamochaeta Wedd.,Leontopodium R. Br. ex Cass, andLogfia (Galbany-Casals & al. 2010).

Helichrysum Mill, is not monophyletic. Some Australasian species had already been transferred to other genera for morphological reasons (seeBayer 2001 andWard & al. 2009 for a review) and later were shown not to be part of the mainHelichrysum clade (Galbany-Casals & al. 2004;Ward & al. 2009;Smissen & al. 2011). This affectsH. bracteatum (Vent.) Willd., an ornamental species (Jäger 2005), which should be known asXerochrysum bracteatum (Vent.) Tzvelev (Bayer 2001).Anaphalis DC. andPseudognaphalium Kirp., two genera of hypothesized allopolyploid origin, are embebbed in the mainHelichrysum clade (Galbany-Casals & al. 2014). The need for a generic re-circumscription of these three genera, plus others, was extensively discussed by Galbany-Casals & al. (2014), who recommended maintainingAnaphalis, Helichrysum andPseudognaphalium as independent genera until more data are available. This affects two taxa present in Germany,A. margaritacea (L.) Benth. & Hook. f., an ornamental but naturalized (Jäger 2011) species native to Asia and North America, andP. luteoalbum (L.) Hilliard & B. L. Burtt. The latter species was treated asGnaphalium luteoalbum L. in Jäger (2005). At present it remains unclear if this species should be included inHelichrysum orPseudognaphalium, or if it should be treated asLaphangium Tzvelev as was done in Jäger (2011).

Astereae (C. Oberprieler)

The most comprehensive molecular phylogenetic analysis of tribeAstereae based on nrDNA ITS sequences was made by Brouillet & al. (2009). To a large extent, its results are supportive of the generic delimitation proposed by Greuter (2003) for the Euro+Med plantbase treatment of the tribe and of Nesom & Robinson's (2007) treatment ofAstereae in Kubitzki'sThe families and genera of vascular plants (Kadereit & Jeffrey 2007).

In subtribeSolidagininae, results by Brouillet & al. (2009) confirm thatSolidago L. is polyphyletic and that the naturalizedS. graminifolia (L.) Salisb. should be transferred toEuthamia (Nutt.) Cass. asE. graminifolia (L.) Nutt. because it belongs to another lineage than the type ofSolidago (i.e.S. virgaurea L.). While in subtribeBellidinae the monophyly of bothBellis L. andBellium L. was repeatedly found in molecular phylogenetic studies based on nrDNA ITS sequences (Fiz & al. 2002;Brouillet & al. 2009;Fiz-Palacios & Valcarcel 2011), phylogenetic analyses in subtribeAsterinae have led to extensive generic rearrangements due to the consistently demonstrated polyphyly ofAster L. in its classical circumscription. According to nrDNA ITS-based analyses by Brouillet & al. (2009), a more narrowly and more naturally circumscribed genusAster in Germany would only compriseA. alpinus L. andA. amellus L., whileA. linosyris (L.) Bernh. should be transferred to the Eurasian genusGalatella Cass. asG. linosyris (L.) Rchb. f., the halophilicA. tripolium L. to the genusTripolium Nees asT. pannonicum (Jacq.) Dobrocz., andA. bellidiastrum (L.) Scop. not only to the separate and monospecific genusBellidiastrum Scop. (asB. michelii Scop.) but also to another subtribe (Bellidinae). The last has also been confirmed by the phylogenetic analyses by Fiz & al. (2002) and Fiz-Palacios & Valcarcel (2011). Finally, molecular phylogenies based on nrDNA ITS alone (Brouillet & al. 2009) or on nrDNA ITS + ETS complemented by the intergenic spacer regiontrnL-trnF of the chloroplast genome (Li & al. 2012b) support the transfer of the naturalized “New World asters” (i.e.A. laevis L.,A. lanceolatus Willd.,A. novae-angliae L.,A. novi-belgii L.,A. parviflorus Nees,A. salignus Willd.,A. versicolor Willd.) to the genusSymphyotrichum Nees (subtribeSymphyotrichinae). On the other hand, Li & al. (2012b) found no evidence for a close relationship betweenCallistephus chinensis (L.) Nees and any other genus of subtribeAsterinae and supported its independent generic status. Finally, in subtribeConyzinae, it has been repeatedly demonstrated (Noyes 2000;Andrus & al. 2009;Brouillet & al. 2009) that neitherConyza Less. norErigeron L. as previously defined are monophyletic; a situation that is best accommodated by merging the two genera intoErigeron, as was already suggested by Greuter (2003). This requires the transfer ofC. bonariensis (L.) Cronquist,C. canadensis (L.) Cronquist andC. sumatrensis (Retz.) E. Walker to this more broadly circumscribed genus (asE. bonariensis L.,E. canadensis L., andE. sumatrensis Retz., respectively).

Anthemideae (C. Oberprieler)

In the S hemisphere subtribeCotulinae, phylogenetic analyses by Himmelreich & al. (2012) based on sequence variation of nrDNA ITS and intergenic spacer regions (psbA-trnH, trnC-petN) of the chloroplast genome have shown thatCotula L. is non-monophyletic, even when the MediterraneanC. cinerea Delile is excluded as the independent and monospecific genusBrocchia Vis. (asB. cinerea (Delile) Vis.) following results by Oberprieler (2004a). Being the type ofCotula, sinking ofLeptinella Cass. andSoliva Ruiz & Pav. into a broader, then monophyletic genus would not affect the name ofC. coronopifolia L., naturalized in the N hemisphere. Of subtribeArtemisiinae, as circumscribed by Oberprieler & al. (2007a,2009), onlyArtemisia L. andLeucanthemella Tzvelev are represented in our area. In the case of the former genus, there is a consistent tendency supported by many molecular phylogenetic studies of the last years (e.g.Vallès & al. 2003;Sanz & al. 2008;Pellicer & al. 2010,2011;Garcia & al. 2011) for lumping the numerous small to large segregate genera (i.e.Crossostephium Less.,Filifolium Kitam.,Mausolea Poljakov,Neopallasia Poljakov,Picrothamnus Nutt.,Seriphidium Fourr.,Sphaeromeria Nutt. andTuraniphytum Poljakov) into a broadly defined and monophyleticArtemisia. On the other hand, studies focusing on phylogenetic relationships among the remainder of theArtemisiinae sensu Oberprieler & al. (2007a,2009) in general and on the generic delimitation ofAjania Poljakov versusChrysanthemum L. in particular, presented no consistent and well-supported evidence for the affiliation ofLeucanthemella Tzvelev to any other genus of the subtribe (Masuda & al. 2009;Zhao & al. 2010). As a consequence,Leucanthemella with its sole European speciesL. serotina (L.) Tzvelev should be treated as an independent evolutionary unit at genus rank. After inclusion, motivated by molecular phylogenetic studies, of the Mediterranean monospecificOtanthus Hoffmanns. & Link and the equally monospecific Turkish endemicLeucocyclus Boiss. in subtribeMatricariinae (Guo & al. 2004;Oberprieler 2004b;Ehrendorfer & Guo 2005),Achillea L. constitutes a monophyletic genus. Support from a comprehensive molecular phylogenetic analysis for the monophyly of the Eurasian and Mediterranean Matricaria L. with its presently accepted six species (Oberprieler & al. 2007b,2009) is still missing. However, the transfer of the AegeanM. macrotis Rech. f. toAnthemis L. (asA. macrotis (Rech. f.) Oberpr. & Vogt) based on nrDNA sequence information (Oberprieler & Vogt 2006) and the repeatedly shown support for the generic independence ofMatricaria (subtribeMatricariinae) fromTripleurospermum Sch. Bip. (subtribeAnthemidinae; e.g.Oberprieler 2004b,2005;Oberprieler & al. 2007a) and fromMicrocephala Pobed. (subtribeHandeliinae; e.g.Oberprieler & al. 2007a;Sonboli & al. 2012) contributed strong evidence for the naturalness ofMatricaria in its present circumscription. In subtribeAnthemidinae sensu Oberprieler & al. (2007a,2009) with its species-rich core generaAnthemis L. andTanacetum L., considerable efforts have been made to achieve a natural delimitation of genera based on molecular phylogenies. After Oberprieler (2001) had shown, with a limited taxon sample, thatAnthemis in its traditional circumscription is paraphyletic, and that the species ofA. sect.Cota (J. Gay) Rchb. f., distinct in their achene morphology, should be transferred to the independent genusCota J. Gay ex Guss. (Greuter & al. 2003), Lo Presti & al. (2010) corroborated this finding based on a comprehensive species sampling (c. 75 % of the described species) and sequence information from both nuclear and plastid markers. With the exclusion of further four species from the Caucasus region (i.e.A. calcarea Sosn.,A. fruticulosa M. Bieb.,A. marschalliana Willd. andA. trotzkiana Bunge) from Anthemis and their accommodation in the newly described genusArchanthemis Lo Presti & Oberpr., and the abovementioned inclusion ofMatricaria macrotis (Oberprieler & Vogt 2006), three natural, morphologically distinct genera were established (Lo Presti & al. 2010;Sonboli & al. 2012). To reflect these phylogenetic findings,Anthemis austriaca Jacq. andA. tinctoria L., hitherto treated asAnthemis in Germany, should be transferred toCota, asC. austriaca (Jacq.) Sch. Bip. andC. tinctoria (L.) J. Gay.

The natural circumscription ofTanacetum L. remains problematic even after considerable taxon and marker sampling. Based on nrDNA ITS and cpDNAtrnH-psbA sequence information, Sonboli & al. (2012) could demonstrate that there is no support for a generic separation of the yellow-rayed or rayless species of Tanacetum from the white- and red-rayed species ofPyrethrum Zinn. On the other hand, even after exclusion of several enigmatic species fromTanacetum based on phylogenetic analyses (i.e.T. annuum L. andT. microphyllum DC. transferred to the newly establishedVogtia Oberpr. & Sonboli [Sonboli & al. 2012];T. paradoxum Bornm. transferred toArtemisia [Sonboli & al. 2011];T. semenovii Herder transferred toRichtera Kar. & Kir. [Sonboli & Oberprieler 2012]) and the suggested inclusion inTanacetum of many satellite genera (e.g.Balsamita Mill.,Gonospermum Less.,Gymnocline Cass.,Hemipappus K. Koch,Lugoa DC.,Spathipappus Tzvelev andXylanthemum Tzvelev), support for a monophyleticTanacetum remains weak and awaits phylogenetic reconstructions based on a broader sampling of molecular markers (Sonboli & al. 2012). For the time being, this argues for the presently used broad generic concept ofTanacetum in Germany.

After having been raised from sectional rank inTanacetum to an independent genus by Heywood (1975),Leucanthemopsis (Giroux) Heywood was considered to be related toLeucanthemum Mill. by Bremer & Humphries (1993) until molecular phylogenetic studies revealed its even closer relationships with three monospecific genera endemic to the Iberian Peninsula,Castrilanthemum Vogt & Oberpr.,Hymenostemma Willk. andProlongoa Boiss. This resulted in its accommodation in the new subtribeLeucanthemopsidinae (Oberprieler & al. 2007a,2009). More recently, a multilocus phylogenetic analysis of all species of the subtribe in a coalescent-based species-tree reconstruction clearly demonstrated the well-supported monophyly ofLeucanthemopsis (Tomasello & al. 2015).

As already discussed by Vogt (1991) in his revision ofLeucanthemum Mill. in the Iberian Peninsula, the genus in its traditional circumscription contained species that are only remotely related to its type,L. vulgare Lam. Accommodation of these divergent species in the independent generaMauranthemum Vogt & Oberpr. andRhodanthemum B. H. Wilcox & al. by Vogt & Oberprieler (1995) and Bremer & Humphries (1993), respectively, has led to a well-circumscribed and strongly supported monophyleticLeucanthemum, as was recently corroborated by a multi-locus phylogenetic analysis by Konowalik & al. (2015).

In subtribeSantolininae sensu Oberprieler & al. (2007a,2009), genus delimitations were studied in molecular phylogenetic analyses by Oberprieler (2002). Based on nrDNA ITS and cpDNAtrnL-trnF sequence variation, this study demonstrated the paraphyly ofChamaemelum Mill. relative to the monospecificCladanthus Cass. Transfer of four W MediterraneanChamaemelum species toCladanthus led to two well-supported monophyletic sister genera, with the widely cultivated and sporadically naturalizedC. nobile (L.) All. and the W MediterraneanC. fuscatum (Brot.) Vasc. as the only members ofChamaemelum.

Glebionis Cass. with the naturalizedG. segetum (L.) Fourr. comprises only two species and is the type genus of the small subtribeGlebionidinae (Oberprieler & al. 2007a,2009). Phylogenetic relationships within this subtribe were studied by Francisco-Ortega & al. (1997), who found little support for the monophyly of the subtribe and for the genus (subChrysanthemum) in a maximum-parsimony analysis based on nrDNA ITS sequence variation. While more recent studies using model-based sequence analysis methods (maximum likelihood) gained strong support for the monophyly of the subtribe (Oberprieler 2005;Oberprieler & al. 2007a), relationships among the genera of Glebionidinae, i.e. the species-richArgyranthemum Webb (24 spp.),Glebionis (two spp.), and the two monospecific generaHeteranthemis Schott andIsmelia Cass., remain unclear, especially after a recent study based on nrDNA ITS sequence variation by Imamura & al. (2015), who foundG. coronaria (L.) Spach nested in a group of Argyranthemum species. If future studies should corroborate the non-monophyly of the four genera of Glebionidinae, and their merging would be necessary to arrive at a monophyletic genus, the oldest genus name for this entity would beHeteranthemis Schott. For the time being, however, retaining the four genera in their present circumscriptions appears preferable due to their morphological and geographical distinctness.

Inuleae (J. W. Kadereit)

Phylogenetic analyses of tribeInuleae have shown that neitherInula L. norPulicaria Gaertn. are monophyletic (Anderberg & al. 2005;Englund & al. 2009), but this has not yet been translated into formal taxonomic changes, although possible taxonomic consequences were discussed by Englund & al. (2009). The species ofInula present in Germany fall into at least four different clades, of whichI. graveolens (L.) Desf. is more closely related toPulicaria and its relatives than toInula and its relatives and has been treated asDittrichia Greuter. Maintainance of this genus will depend on future treatment of the various lineages ofPulicaria. If, after exclusion of some lineages as suggested by Englund & al. (2009), a broad concept ofPulicaria is adopted,Dittrichia will have to be included in that genus. If, on the other hand, a narrow concept ofPulicaria is adopted,Dittrichia would remain an independent genus and the two species of Pulicaria present in Germany (P. dysenterica (L.) Bernh. andP. vulgaris Gaertn.) would remain inPulicaria. Adoption of a broad concept ofInula would require inclusion ofCarpesium L. andTelekia Baumg. Adoption of a narrow concept would require distribution of the German species in probably several genera, dependent on treatment, and onlyI. helenium L., as the type, would remain inInula.

Helenieae (J. W. Kadereit)

BothBidens L. andCoreopsis L. have been shown not to be monophyletic (Mort & al. 2008), but this has not yet been translated into taxonomic changes.

Heliantheae (J. W. Kadereit)

BothAmbrosia L. andIva L. have been found not to be monophyletic (Miao & al. 1995).Ambrosia becomes monophyletic after inclusion ofHymenoclea Torr. & A. Gray, as proposed by Panero (2007), whereas parts ofIva are better accommodated in other genera. This affects the GermanI. xanthiifolia Nutt., which, according to Panero (2007), should be considered a species ofEuphrosyne DC. and calledE. xanthiifolia (Nutt.) A. Gray.

Madieae (J. W. Kadereit)

AlthoughEriophyllum Lag. does not appear to be monophyletic (Baldwin & al. 2002),E. lanatum (Push) Forbes, a naturalized ornamental in Germany, is part of the perennial clade, which also contains the type of the genus name. In consequence, no change of name will be necessary should the genus be split.

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