TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Phylogeny of Apiaceae subtribe Daucinae and the taxonomic delineation of its genera Łu3asz Banas1a3, Aneta Wo2ewódz3a, Ja3ub Baczyńs31, Jean-P1erre Reduron, Marc1n P1wczyńs31, Renata Kurzyna-Młyn13, Rafał Guta3er, Agn1esz3a Czarnoc3a-C1ec1ura, Sylw1a Kosmala-Grzechn13 & Krzysztof Spal13 1 Department of Molecular Phylogenetics and Evolution, Institute of Botany, Faculty of Biology, University of Warsaw Biological and Chemical Research Centre, Żwirki i Wigury 4, 02-089 Warszawa, Poland 2 VIA APIA, 10 rue de l’Arsenal, 68100 Mulhouse, France 3 Chair of Ecology and Biogeography, Nicolaus Copernicus University in Toruń, Lwowska 1, 87-100 Toruń, Poland Author for correspondence: Krzysztof Spalik, spalik@biol.uw.edu.pl ORCID KS, http://orcid.org/0000-0001-9603-6793 DOI http://dx.doi.org/10.12705/653.8 Abstract Scandiceae subtribe Daucinae encompasses umbellifers that have fruits with prominent secondary ridges projecting into wings (former tribe Laserpitieae) or spines (former tribe Caucalideae pro parte). It comprises several economically or medicinally important genera including Cuminum, Daucus, Laser, Laserpitium and Thapsia among others. Recent molecular studies, based mostly on nrDNA ITS sequences, revealed that neither Daucus nor Laserpitium are monophyletic. To address issues of relationships and apply respective nomenclatural changes, we obtained additional ITS sequences as well as independent data from three plastid markers—rps16 intron, rpoC1 intron and rpoB-trnC intergenic spacer—for a comprehensive sample of the subtribe. We examined data for 260 accessions representing all genera of Daucinae and 81 of its ca. 93 species. Phylogenetic trees were estimated using maximum likelihood and Bayesian inference methods. The results indicate that former Laserpitieae constitute a paraphyletic grade at the base of the spiny-fruited members of Daucinae while traditionally delimited Daucus and Laserpitium are polyphyletic. To maintain a monophyletic Daucus, we suggest including the following genera and species into its synonymy: Agrocharis, Melanoselinum, Monizia, Pachyctenium, Pseudorlaya, Rouya, Tornabenea, Athamanta dellacellae and Cryptotaenia elegans. The species of Laserpitium occur in seven clades and only six species of the Laserpitium s.str. clade retain the generic name. Several species are transferred to Ekimia, Laser and Thapsia; additionally, a monospecific genus Siler is restored and a new genus, Silphiodaucus, is established. The inclusion of Ammodaucus into Thapsia suggested in an earlier study is not supported. The position of Laserpitium pseudomeum requires further study. Keywords cpDNA; Daucus; Laserpitieae; Laserpitium; nrDNA ITS; Scandiceae; taxonomy Supplementary Material The Electronic Supplement (Figs. S1–S3; Table S1) is available in the Supplementary Data section of the online version of this article at http://ingentaconnect.com/content/iapt/tax; DNA sequence alignment is available from TreeBase (http://purl.org/phylo/treebase/phylows/study/TB2:S18012) INTRODUCTION Umbellifers (Apiaceae) have dry fruits (schizocarps) that split into two one-seeded parts (mericarps) acting as dispersal units. These fruits exhibit an array of structural peculiarities, including various appendages that may facilitate fruit dispersal. Winged fruits are generally regarded as adapted to transportation by wind (Theobald, 1971; Jongejans & Telenius, 2001), whereas those with spines and bristles are considered as epizoochorous (Jury, 1982; Williams, 1994; Williams & Guries, 1994; Spalik & al., 2001). Fruit appendages may therefore be subject to strong selective pressure resulting in homoplasy. The diversity of winged fruits in the family is substantial, suggesting, therefore, their independent origins (Theobald, 1971; Liu & al., 2006). Multiple evolution of winged fruits has been demonstrated for several lineages of subfamily Saniculoideae (Calviño & al., 2008). Schizocarps with prominent secondary ribs that project into broad wings constitute major diagnostic characters for the apioid tribe Laserpitieae Coss. & Germ. sensu Drude (1897– 1898). Drude placed nine genera in this tribe, distributing them among three subtribes: Silerinae Tausch, Elaeoselininae Lange in Willk. & Lange and Thapsiinae Coss. & Germ. Silerinae included only Siler Crantz, Elaeoselininae incorporated Elaeoselinum W.D.J.Koch ex DC. and Margotia Boiss., and Thapsiinae encompassed Guillonea Coss., Laserpitium L., Melanoselinum Hoffm., Polylophium Boiss., Thapsia L., and Tornabenea Parl. Since Drude’s treatment, the circumscription of Laserpitieae has not generally been changed, although its subtribes are no longer recognized and some modifications at the generic level have been introduced (summarized in Table 1). Four laserpitioid genera have been sunk into Thapsia (Weitzel & al., 2014), and Melanoselinum edule (Lowe) Drude is usually recognized in Received: Aug | returned for (first) revision: Nov | (last) revision received: Feb | accepted: Feb || publication date(s): online fast track, n/a; in print and online issues, Jun || Published online “open-access” under the terms of the Creative Commons Attribution 4.0 (CC BY 4.0) License (see https://creativecommons.org/licenses/) || © International Association for Plant Taxonomy (IAPT) Version of Record 563 No. of species Nomenclatural type or species placed in Daucinae Agrocharis Hochst. A. melanantha Hochst. 4 4 (3) Townsend (1989) Ammodaucus Coss. & Durieua A. leucotrichus Coss. & Durieu 2 1 (1) Reyes-Betancort & al. (2007) A. dellacellae Asch. & Barbey ex E.A.Durand & Barratte 1 1 (1) Spalik & Downie (2001) Athamanta L., pro parte, typo excl. b All This study: all (cpDNA) Reference Genus Cryptotaenia DC., pro parte, typo excl. C. elegans Webb ex Bolle 1 1 (1) Spalik & Downie (2007) Cuminum L. C. cyminum L. 3 2 (2) Rechinger (1987a), Czerepanov (1995) b Sáenz Laín (1981), Grzebelus & al. (2011) Daucus L. D. carota L. 26 23 (20) Ekimia H.Duman & M.F.Watson E. bornmuelleri (Hub.-Mor. & Reese) H.Duman & M.F.Watson 1 1 (1) Duman & Watson (1999), Lyskov & al. (2015) Laser Borkh. ex G.Gaertn. & al. L. trilobum (L.) Borkh. ex G.Gaertn. & al. 1 1 (1) Rechinger (1987b), Reduron (2007b) Laserpitium L. L. gallicum L. 19(–35) 18 (14) Melanoselinum Hoffm. M. decipiens (Schrad. & J.C.Wendl.) Hoffm. 1 1 (1) Press & Short (1994) Monizia Lowe M. edulis Lowe 1 1 (1) Press & Short (1994) Pimenov & Leonov (1993)c Version of Record Orlaya Hoffm. O. grandiflora (L.) Hoffm. 3 3 (2) Jury (2003), Hartvig (1986) Pachyctenium Maire & Pamp. P. mirabile Maire & Pamp. 1 1 (0) Lee & al. (2001) Polylophium Boiss. P. orientale Boiss. 2 2 (1) Rechinger (1987c) Pseudorlaya (Murb.) Murb. P. pumila (L.) Murb. 2 2 (0) Rutherford & Jury (2003) Rouya Coincy R. polygama (Desf.) Coincy 1 1 (1) Reduron (2008) Thapsia L.d T. villosa L. 18 15 (8) Pujadas Salv0 & Plaza Arregui (2003), Pujadas Salv0 & Roselló (2003), Weitzel & al. (2014) *Distichoselinum García-Martín & Silvestre D. tenuifolium (Lag.) García Martín & Silvestre 1 1 (1) García Martín & Silvestre (1985), García Martín (2003a) *Elaeoselinum W.D.J.Koch ex DC. E. meoides (Desf.) W.D.J.Koch ex DC. 4(–5) 3 (2) García Martín & Silvestre (1985), García Martín (2003b), Brullo & al. (2003) G. scabra (Cav.) Coss. 1 1 (1) Montserrat (2003a) M. laserpitioides Boiss. (= M. gummifera (Desf.) Lange) 1 1 (1) García Martín & Silvestre (1985), García Martín (2003c) Tornabenea Parl. T. insularis (Parl. ex Webb.) Parl. 6 3 (1) Brochmann & al. (1997); Schmidt & Lobin (1999) a b c d Ammodaucus was recently synonymized with Thapsia based on nrDNA ITS analyses only (Weitzel & al., 2014). However, this placement has little support from morphology while the apparently long branch of this taxon within Thapsia makes this assignment problematic. Athamanta and Cryptotaenia are polyphyletic with single species placed in Daucinae; therefore, only these species are considered. Pimenov & Leonov (1993) estimated the number of species in Laserpitium at 35. However, upon verification of the local Floras and regional revisions for western Eurasia and the Mediterranean region we have found only 19 currently recognized species. The genera that are marked with asterisks have been recently sunk into the synonymy of Thapsia based on nrDNA ITS analyses only (Weitzel & al., 2014). Because that study did not include all members of the synonymized genera, we also provide their original taxonomic treatment. TAXON 65 (3) • June 2016: 563–585 *Guillonea Coss. *Margotia Boiss. Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae 564 Table 1. Genera of Apiaceae provisionally placed in tribe Daucinae and sampled for this study. TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae the monospecific genus Monizia Lowe (Press & Short, 1994). The nomenclatural type of Siler, S. montanum Crantz, has been returned to Laserpitium (as L. siler L.), whereas another member of Siler sensu Drude, S. trilobum (L.) Scop., is at present placed in the monospecific genus Laser Borkh. ex G.Gaertn. & al. and usually recognized in tribe Peucedaneae Dumort. (Pimenov & Leonov, 1993). The monospecific genus Rouya Coincy has been segregated from Thapsia. Additionally, several species have been transferred to or described in Tornabenea, Polylophium and Thapsia. Recently, phylogenetic analyses of ITS sequences suggested that an obscure endemic of Turkey, Ekimia H.Duman & M.F.Watson, is closely related to some Anatolian species of Laserpitium (Lyskov & al., 2015). Its only member, E. bornmuelleri (Hub.-Mor. & Reese) H.Duman & M.F.Watson, was originally placed in Prangos Lindl. because of its fruits with wavy wings that are superficially similar to those of the latter (Duman & Watson, 1999). Drude (1897–1898) suggested that Laserpitieae were allied with his tribe Dauceae W.D.J.Koch, whose members also possess fruits with prominent secondary ribs. In contrast to those in Laserpitieae, the ribs of Dauceae are spiny. Calestani (1905) and Koso-Poljansky (1916) went even further in uniting Laserpitieae and Dauceae. Calestani (1905) synonymized Thapsia, Elaeoselinum, Margotia, and Guillonea with Laserpitium, recognizing the latter in Ligusticeae Calest. subtribe Daucinae Dumort., although at the same time he included Siler, a segregate of Laserpitium, in subtribe Peucedaninae Tausch. For Tornabenea, he recognized a separate subtribe, although he neither indicated its content nor provided its description because he was concerned with European species only. In his worldwide revision of umbellifers, Koso-Poljansky (1916) followed Calestani’s treatment of Daucinae and included all Drude’s Laserpitieae in this subtribe: Elaeoselinum (including Margotia), Guillonea, Laserpitium s.l. (i.e., including Laser, Melanoselinum, Polylophium, Rouya, Siler, and Thapsia), Monizia, and Tornabenea. Interestingly, he transferred some species of Laserpitium, including L. prutenicum L. and L. hispidum M.Bieb., to Daucus L. placing them in a separate sect. Silphiodaucus Koso-Pol. Spiny or hairy fruits with prominent secondary ribs also occur in those umbellifers that were placed by Drude (1897– 1898) in Scandiceae subtribe Caucalidinae Tausch, and the entire tribe Scandiceae Spreng. was circumscribed based on the occurrence of calcium oxalate crystals in the parenchymatic cells surrounding the carpophore. In contrast to Drude’s treatment, the revisions of Apiaceae by Bentham (1867) and Boissier (1872) united all spiny-fruited umbellifers in tribe Caucalideae Spreng. Such an approach was also adopted by Heywood & Dakshini (1971), and their treatment served as the basis for a multivariate study of Caucalideae. It resulted in a new circumscription of the tribe (Heywood, 1982) that was generally accepted (Pimenov & Leonov, 1993). The relationships among Laserpitieae, Caucalideae, and Scandiceae have also been investigated using molecular data. First phylogenetic studies using nuclear rDNA ITS (Downie & Katz-Downie, 1996), plastid rpoC1 intron (Downie & al., 1996), and plastid matK sequences (Plunkett & al., 1996a) suggested that members of these tribes form a single clade although the relationships among them remained unresolved due to limited sampling. In these analyses, the representatives of Laserpitium grouped with some members of Caucalideae forming a Daucus clade. Subsequent molecular studies indicated that other members of Laserpitieae—Thapsia (Rasmussen & Avato, 1998; Weitzel & al., 2014), Laser and Polylophium (Katz-Downie & al., 1999), Monizia (Downie & al., 2000c), and Melanoselinum (Lee & Downie, 2000)—also belong to this clade, treated thereafter as Scandiceae subtribe Daucinae (Downie & al., 2000a, 2001, 2010). Interestingly, these former members of Laserpitieae did not form a monophyletic group within Daucinae (Downie & al., 2000c, 2001; Spalik & Downie, 2007). Moreover, some of them were placed within Daucus, making the delineation of this economically important genus problematic. The species of Daucus formed two clades with the Daucus I clade encompassing the nomenclatural type of the genus, D. carota L., which includes the cultivated carrot. This clade was sister to wingedfruited Melanoselinum and Monizia, while the closest relative of D. carota was another laserpitioid genus, Tornabenea (Spalik & Downie, 2007). The phylogenetic relationships among the species of Daucus, particularly the D. carota complex, were recently subject to intense molecular research (Iorizzo & al., 2013; Spooner & al., 2013), but these studies did not result in a new circumscription of the genus due to limited taxonomic sampling. Molecular studies that included more than one member of Laserpitium suggested that this genus is polyphyletic (KatzDownie & al., 1999; Lee & Downie, 1999, 2000; Downie & al., 2000c; Spalik & Downie, 2007; Weitzel & al., 2014). Among the major clades of umbellifers, subtribe Daucinae is one of the most important from an economical point of view because it includes the cultivated carrot, the most widely grown crop of Apiaceae (Iorizzo & al., 2013). Wild species of Daucus constitute an invaluable gene pool for developments in carrot breeding and have also been investigated with respect to the antimicrobial properties of oil components (Grzebelus & al., 2011). Subtribe Daucinae encompasses many taxa with potential medicinal value including Laser (Parlatan & al., 2009), Laserpitium (Popović & al., 2013) and Thapsia (Andersen & al., 2015). Therefore, a classification system congruent with the phylogeny of this group is of considerable practical importance. In this paper, we examine the phylogenetic relationships within Daucinae (tribe Scandiceae) with a comprehensive sampling of species traditionally classified in tribes Dauceae and Laserpitieae and using phylogenetic markers from nuclear and plastid genomes. Our aim is to provide a new taxonomic treatment of this economically important branch of umbellifers. MATERIALS AND METHODS Taxon and molecular marker sampling. — Two hundred sixty accessions representing putative members of Scandiceae subtribe Daucineae and outgroups were examined for molecular sequence variation (Appendix 1; Electr. Suppl.: Table S1). Specifically, all genera and 81 of ca. 93 ingroup species were sampled (Table 1). Outgroups included representatives of other Version of Record 565 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae subclades of tribe Scandiceae (Downie & al., 2010). If possible, voucher specimens were examined to confirm their determination (see Appendix 1 and Electr. Suppl.: Table S1 for details). For all accessions, we obtained the nuclear ribosomal DNA internal transcribed spacer (ITS) region that has already been successfully used in phylogenetic analyses of Apiaceae (Downie & al., 2001). However, our preliminary phylogenetic analyses demonstrated that this marker does not provide adequate resolution among the laserpitioid taxa. Therefore, for a representative set of accessions comprising 18 genera and 59 species of Daucinae we also included three chloroplast markers, the rpoC1 and rps16 introns and the rpoB-trnC intergenic spacer. These markers have already been employed in phylogenetic analyses of umbellifers (Downie & al., 1996; Downie & al., 2000b; Calviño & al., 2010; Panahi & al., 2015). DNA extraction, amplification, and sequencing. — Total genomic DNA was isolated from ca. 20 mg of dried leaf tissue using a DNeasy Plant Mini Kit (Qiagen, Hilden, Germany). For the nrDNA ITS sequence, the DNA samples were PCR-amplified using primers ITS4 and ITS5 (White & al., 1990) or N-nc18S10 and C26A (Wen & Zimmer, 1996). For some accessions, the ITS 1 and ITS 2 regions were amplified separately using the following pairs of primers: 18S-ITS1-F and 5.8S-ITS1-R for ITS 1, and ITS-3N and ITS4 for ITS 2 (Spalik & Downie, 2006). The rpoB-trnC intergenic spacer region was amplified with primers rpoB and trnCGCAR from Shaw & al. (2005). Because this region may include long insertions and mononucleotide repeats, three additional internal primers were designed to aid its amplification and sequencing: rpoB400 (AAG ATC AAA TGC CGA ATC CA), 400trnC (ATG GAA TTT TGT ATA GAA TAT CAA) and 400trnC2 (TTT CCT GCT TAA GAG TGG ATT). The pair rpoB and rpoB400 covers ca. 400 bp at the beginning of the spacer, while 400trcC and trnCGCAR were used to amplify the remainder of the region. The reverse primer 400trnC2 located close to the end of the spacer was used together with 400trnC to amplify the middle region for some difficult samples. The rps16 intron was amplified using external primers 5exonC and 3exonR from Calviño & al. (2006) or s16exF and s16exR from Panahi & al. (2015); additionally, two internal primers, s16inF and s16inR were also used (Panahi & al., 2015). The complete rpoC1 intron and parts of the flanking exon regions were amplified using external primers C1exF and C1exR; additionally, internal primers C1inF and C1inR were used to amplify this marker in part and to sequence difficult samples (Panahi & al., 2015). The details of the PCR reactions are provided elsewhere (Panahi & al., 2015). PCR products were checked on 1% agarose gel with ethidium bromide or Midori Green Advance DNA Stain (Nippon Genetics Europe, Dueren, Germany). Depending on their quality, they were purified using the QIAquick PCR Purification Kit (Qiagen) or gel purified using the QIAEX II Agarose Gel Extraction Kit or QIAquick Gel Extraction Kit (Qiagen). Cycle sequencing reactions were performed using the Big Dye Terminator v.3.1 Cycle Sequencing Kit (Applied Biosystems, Waltham, Massachusetts, U.S.A.) with the same primers as used for PCR amplifications. The products were purified using Sephadex columns or DyeEx 2.0 Spin Kit (Qiagen) and sequenced using an automated DNA sequencer at IBB PAS (Warsaw, Poland). The sequences were assembled and edited using SeqMan II v.4.0 (Dnastar, Madison, Wisconsin, U.S.A.). All newly obtained sequences have been deposited in GenBank (Appendix 1; Electr. Suppl.: Table S1). Sequence and phylogenetic analyses. — For each marker separately, DNA sequences were aligned using MAFFT v.7.123b (Katoh & Toh, 2008) and the resulting matrix was corrected manually if necessary. The parts of the alignment containing especially large numbers of gaps were identified using trimAl v.1.2rev59 with the “automated1” algorithm (Capella-Guti6rrez & al., 2009) and excluded from the analyses. Accessions with identical sequences (with uncorrected p distance = 0) were represented in the analyses by a single terminal (the number of unique and duplicate sequences in each analysis is given in Table 2). The congruence of the datasets was assessed using a hierarchical likelihood ratio test (hLTR) implemented in Concaterpillar v.1.8a (Leigh & al., 2008). Phylogenetic analyses were performed using the maximum likelihood (ML) method implemented in RAxML v.8.1.18 (Stamatakis, 2014) and the Bayesian inference (BI) method implemented in MrBayes v.3.2.5x64 (Ronquist & al., 2012). For Bayesian analyses of nrDNA ITS data, the models of nucleotide substitution were chosen with ModelGenerator v.0.85 (Keane & al., 2006). Partition schemes and substitution Table 2. Characteristics of the datasets used in this study. No. of unique / duplicate sequences No. of aligned positions rpoB-trnC spacer rpoC1 intron rps16 intron cpDNA Full Reduceda 212 / 48 34 /1 70 / 5 89 / 9 106 / 4 241/19 109 /1 649 1643 1139 984 3766 4415 2772 excluded (ambiguous) 49 392 111 129 632 681 289 constant 203 1010 854 672 2536 2739 1753 variable 397 241 174 183 598 995 730 containing gaps and missing data 270 243 503 99 3134 3404 2093 3.4 3.9 1.6 0.4 40.7 62.1 17.5 % of gaps and missing data in the matrix a Combined ITS Combined ITS, rpoC1 intron and rps16 intron matrix comprising only accessions, for which both ITS and cpDNA data were available. 566 Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae models for the analyses of plastid and combined plastid and nuclear data were inferred using PartitionFinder v.1.1.1 (Lanfear & al., 2012, 2014). Bayesian analyses were carried out for 40 million generations with four Monte Carlo Markov chains initiated and a sampling frequency of 1000 generations. Two independent runs were initiated for each analysis. The initial 25% of saved trees were discarded as a burn-in phase and the 50% majority-rule consensus tree and posterior probabilities (PP) of particular clades were calculated based on the remaining 60,000 trees. The effective sample size (ESS) for the estimated parameters and the convergence of the independent runs were checked using Tracer v.1.6.0 (Rambaut & al., 2014). For ML analyses, we employed the GTR + G substitution model. In RAxML, GTR(+ G + I) is the only nucleotide substitution model implemented because according to the author of the program it is better to efficiently implement and optimize this model instead of offering a spectrum of models that are programmed in a generic and thus inefficient way (Stamatakis, 2014). Similarly, GTR + G is preferred over GTR + G + I because the substitution rate heterogeneity is already accounted for with gamma distribution and, therefore, parameters G and I cannot be optimized independently from each other (Stamatakis, 2014). Best ML trees were found based on 200 independent searches starting from distinct randomized maximum parsimony trees. Bootstrap support (BS) was estimated based on 1000 rapid bootstrap replicates. In order to check whether the number of replicates was sufficient, we performed a posteriori bootstopping analysis with the extended majority-rule consensus tree as a convergence criterion. For the analyses of combined data, we used the partition schemes inferred from PartitionFinder. In order to identify terminals that introduce putative topological conflict, we used the program compat.py that compares bootstrap values for trees inferred from different partitions (Kauff & Lutzoni, 2002). As significantly unstable terminals, we regarded those that occurred in different and well-supported clades (BS ≥ 75) in trees inferred from separate analyses of nrDNA and cpDNA. Then, we excluded these terminals from the matrices, checked again the congruence of the datasets and repeated ML and BI analyses of combined data. Because the appendages of secondary ribs are principal features delimiting genera, and winged vs. spiny secondary ribs were used to distinguish tribes Dauceae and Laserpitieae in traditional classification systems of umbellifers, we performed maximum parsimony (MP) mapping of this character using a tree from the ML analyses of the combined data. This tree was transformed into a cladogram and pruned such that each taxon was represented by one accession. The reconstruction of character changes was performed using Mesquite v.3.04 (Maddison & Maddison, 2015). lower number of cpDNA sequences obtained, particularly of the rpoB-trnC intergenic spacer, resulted mostly from unsuccessful PCR amplification of these regions from low-quality DNA samples obtained from herbarium material. Aligned ITS sequences comprised a lower proportion of ambiguously aligned positions than plastid markers: only 7.6% of ITS positions were excluded from the analyses compared to 16.8% of aligned positions deleted from the combined cpDNA matrix (Table 2). Several accessions yielded identical sequences of some markers. For instance, of the 260 ITS sequences only 212 were included in the ITS matrix (Table 2). However, some accessions that had identical ITS sequences provided different cpDNA sequences and vice versa. Therefore, in the full combined matrix only 19 accessions were considered as duplicates. Five matrices were used in subsequent analyses. First, matching nrDNA ITS and cpDNA matrices with 110 accessions (denoted #1 and #2 with 98 and 106 terminals, respectively) were analysed in order to assess the congruence of markers and to identify accessions introducing topological conflicts. Large ITS matrix (#3) including 260 accessions (212 terminals) was additionally analysed and the results were compared with those from small ITS matrix (#1) in order to evaluate the effect of taxonomic sampling on tree topology. Combined data matrix (#4) with all 260 accessions (241 terminals) was used to identify major highly supported clades that may be recognised as genera. In the combined data matrix we also included those accessions for which only ITS data were available coding respective cpDNA data as missing (Table 2). In effect, this matrix comprised 62.1% missing data. In order to evaluate the impact of those missing data on tree topology, we also analysed a reduced matrix (#5) comprising 110 accessions (109 terminals) and three markers: ITS, rpoC1 intron and rps16 intron. This matrix had only 17.5% missing data. Hierarchical clustering of the markers with Concaterpillar resulted in concatenation of all three plastid loci (Table 3). In contrast, the concatenation of combined cpDNA markers and nrDNA ITS data was rejected with P < 10 –6. The comparisons of ML bootstrap analyses performed with compat.py identified 13 accessions that may cause a conflict (Figs. 1, 2). After Table 3. Hierarchical tests of congruence of phylogenetic signal performed with Concaterpillar. Dataset P-value All taxa rpoB-trnC spacer + rpoC1 intron 0.367 rpoB-trnC spacer + rpoC1 intron + rps16 intron 0.177 rpoB-trnC spacer + rpoC1 intron + rps16 intron + ITS < 10−6* Conflicting terminals excluded RESULTS Sequence analyses. — For this study, we obtained 119 sequences of ITS, 73 sequences of rps16 intron, 63 sequences of rpoC1 intron and 26 sequences of rpoB-trnC intergenic spacer from 126 accessions (Appendix 1; Electr. Suppl.: Table S1). The ITS + rpoB-trnC spacer 0.610 ITS + rpoB-trnC spacer + rpoC1 intron spacer 0.240 ITS + rpoB-trnC spacer + rpoC1 intron + rps16 intron 0.199 An asterisk indicates a statistically significant conflict between markers for α = 0.017 corrected for multiple comparisons (with initial α = 0.05). Version of Record 567 Fig. 1. Maximum likelihood tree inferred from analyses of 110 nrDNA ITS sequences of Daucinae and outgroups obtained from accessions for which plastid DNA data were also available (dataset #1, see the text for details). Accessions that yielded identical sequences are represented by a single terminal. Bootstrap support and posterior probability (for nodes that also occurred in Bayesian majority-rule consensus tree) are given along the branches. Accessions with significantly different relationships in nrDNA and cpDNA trees are marked with boldface. Major groups are bracketed. Daucus carota subsp. maritimus 448 61/0.94 Daucus carota subsp. carota 32 443 444 Daucus carota subsp. commutatus 3 Daucus carota subsp. carota G98 Daucus virgatus 659 100/1.0 Daucus carota subsp. azoricus 610 Daucus carota subsp. halophilus 9 Daucus carota subsp. gadecaei 34 100/1.0 Daucus carota subsp. gummifer 445 Daucus carota subsp. hispanicus G30 Tornabenea annua 521 97/1.0 Daucus syrticus 468 99/1.0 Rouya polygama 523 98/1.0 Athamanta dellacellae 435 Daucus mauritii G45 100/1.0 52/0.94 Daucus aureus G18 100/1.0 Daucus setifolius 237 G56 Daucus setifolius 467 37/0.73 Daucus crinitus 14 54/0.78 Daucus muricatus 271 64/0.63 Daucus muricatus G51 100/1.0 Daucus muricatus G50 61/0.69 Daucus muricatus 281 60/0.90 99/1.0 Daucus tenuisectus 266 Cryptotaenia elegans 39 Melanoselinum decipiens G72 G73 95/0.99 Monizia edulis 13 G74 88/0.81 Daucus montanus 17 95/1.0 Daucus glochidiatus 456 33/− Daucus glochidiatus 455 Daucus guttatus 238 458 31/− Daucus pusillus 466 G53 100/1.0 Daucus pusillus G52 44/− 100/1.0 Daucus arcanus 260 100/1.0 Daucus durieua G38 Daucus durieua 453 97/1.0 Daucus durieua 40 Daucus littoralis 461 99/1.0 100/1.0 98/1.0 Daucus bicolor 41 Daucus littoralis 8 Daucus involucratus 460 272 99/1.0 100/1.0 Daucus involucratus 37 Daucus conchitae 236 97/1.0 71/0.94 Daucus broteri 440 93/1.0 100/1.0 Daucus broteri 277 Daucus broteri 276 90/1.0 72/0.95 Agrocharis incognita 46 100/1.0 Agrocharis pedunculata 247 Agrocharis incognita 245 Laserpitium hispidum 494 100/1.0 Laserpitium prutenicum subsp. prutenicum 649 27/− Orlaya daucoides G75 100/1.0 Orlaya grandiflora 258 Laserpitium siler 517 100/1.0 Laserpitium siler G70 Laserpitium siler 516 33/− Laserpitium siler 515 Laserpitium pseudomeum 513 27/− Laserpitium archangelica 481 96/1.0 55/0.65 Laserpitium affine 480 Polylophium panjutinii G82 13/− Laserpitium carduchorum 483 29/− Ammodaucus leucotrichus 12 47/− 26/− Laserpitium stevenii 518 100/1.0 Laser trilobum G61 13/− Laser trilobum 611 100/1.0 Cuminum cyminum 339 100/1.0 Cuminum setifolium 20 Cuminum cyminum G7 Thapsia garganica 525 100/1.0 Thapsia garganica 526 33/− 95/1.0 Thapsia garganica 527 88/0.98 Thapsia transtagana 524 Thapsia tenuifolia 469 87/1.0 100/1.0 Thapsia thapsioides 477 Thapsia thapsioides 478 52/0.69 Thapsia sp. 471 Thapsia gummifera 520 88/1.0 Thapsia meoides 473 82/1.0 81/0.50 Laserpitium eliasii subsp. ordunae 485 100/1.0 Laserpitium eliasii subsp. eliasii 484 94/1.0 Laserpitium eliasii subsp. thalictrifolium 486 95/1.0 Laserpitium nestleri subsp. nestleri 501 28/− Laserpitium nestleri subsp. flabellatum 505 Laserpitium nestleri subsp. lainzii 506 100/1.0 97/1.0 Thapsia villosa var. dissecta 528 98/1.0 Thapsia scabra 479 Laserpitium petrophilum 509 100/1.0 Ekimia bornmuelleri 655 100/1.0 Laserpitium gallicum subsp. orospedanum 489 94/0.98 Laserpitium gallicum subsp. paradoxum 490 100/1.0 Laserpitium gallicum subsp. gallicum 487 Laserpitium gallicum subsp. angustissimum 488 60/− Laserpitium krapfii subsp. krapfii 496 49/0.50 Laserpitium krapfii subsp. gaudinii 497 66/0.56 Laserpitium latifolium 498 Leutea avicennae 67 96/1.0 Ferula olivacea 363 Ferula communis 195 94/1.0 Glaucosciadium cordifolium 221 100/1.0 72/0.99 Glaucosciadium insigne 74 Anthriscus sylvestris 83 0.01 Daucus I Macaronesian Daucus II Agrocharis Silphiodaucus Orlaya Siler Laser Cuminum Thapsia Ekimia Laserpitium s.str. Outgroup 568 Version of Record Daucus s.l. TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae TAXON 65 (3) • June 2016: 563–585 Fig. 2. Maximum likelihood tree inferred from analyses of combined sequences of the rps16 and rpoC1 introns and the rpoB-trnC intergenic spacer for the same 110 accessions as in Fig. 1 (dataset #2, see the text for details). Accessions that yielded identical sequences are represented by a single terminal. Bootstrap support and posterior probability (for nodes that also occurred in Bayesian majority-rule consensus tree) are given along the branches. Accessions with significantly different relationships in nrDNA and cpDNA trees are marked with boldface. Major groups are bracketed. Daucus syrticus 468 Daucus carota subsp. gadecaei 34 Daucus virgatus 659 Daucus carota subsp. carota G98 Tornabenea annua 521 95/0.97 Daucus carota subsp. carota 32 Daucus carota subsp. halophilus 9 Daucus carota subsp. commutatus 3 Daucus carota subsp. azoricus 610 Daucus carota subsp. carota 443 444 89/0.85 Daucus carota subsp. gummifer 445 Daucus carota subsp. maritimus 448 54/0.69 Daucus carota subsp. hispanicus G30 2/− Rouya polygama 523 Daucus tenuisectus 266 Daucus setifolius G56 94/0.58 Daucus setifolius 467 62/0.53 Daucus crinitus 14 79/− 6/− Daucus setifolius 237 32/0.92 Athamanta dellacellae 435 Daucus aureus G18 93/0.84 Daucus mauritii G45 Daucus muricatus G51 G50 99/1.0 Daucus muricatus 281 Daucus muricatus 271 72/1.0 95/1.0 Monizia edulis G74 Monizia edulis 13 70/0.95 Melanoselinum decipiens G72 Melanoselinum decipiens G73 Cryptotaenia elegans 39 Daucus broteri 440 49/0.97 18/0.52 Daucus broteri 277 Daucus conchitae 236 42/0.96 Daucus broteri 276 53/0.97 Daucus involucratus 272 88/1.0 Daucus involucratus 460 Daucus involucratus 37 85/1.0 Daucus littoralis 461 /− 45/0.57 Daucus bicolor 41 44/0.96 Daucus littoralis 8 Daucus pusillus G53 Daucus arcanus 260 86/1.0 Daucus pusillus G52 Daucus pusillus 466 79/1.0 Daucus guttatus 238 64/0.88 Daucus guttatus 458 92/1.0 100/1.0 Daucus durieua G38 98/1.0 Daucus durieua 453 Daucus durieua 40 97/1.0 Daucus glochidiatus 455 41/0.82 Daucus glochidiatus 456 Daucus montanus 17 26/0.88 Agrocharis incognita 46 100/1.0 Agrocharis pedunculata 247 Agrocharis incognita 245 Laserpitium prutenicum subsp. prutenicum 649 88/0.99 Laserpitium hispidum 494 Laserpitium pseudomeum 513 21/0.95 94/0.98 Polylophium panjutinii G82 96/1.0 Laser trilobum G61 50/0.70 Laserpitium stevenii 518 94/1.0 Laserpitium archangelica 481 93/1.0 Laser trilobum 611 Laserpitium carduchorum 483 17/0.90 85/1.0 64/0.67 Laserpitium affine 480 Orlaya grandiflora 258 99/1.0 Orlaya daucoides G75 49/0.73 Laserpitium siler 515 Laserpitium siler 516 37/0.67 Laserpitium siler 517 62/0.99 Laserpitium siler G70 Thapsia thapsioides 477 95/1.0 61/0.99 Thapsia meoides 473 59/0.98 Thapsia thapsioides 478 88/1.0 Thapsia sp. 471 Thapsia tenuifolia 469 88/1.0 Thapsia gummifera 520 75/0.92 99/1.0 Thapsia garganica 525 93/1.0 Thapsia garganica 526 100/1.0 Thapsia garganica 527 Thapsia transtagana 524 73/0.97 Thapsia villosa var. dissecta 528 64/0.97 Thapsia scabra 479 86/1.0 93/1.0 Laserpitium eliasii subsp. ordunae 485 Laserpitium nestleri subsp. lainzii 506 34/− Laserpitium nestleri subsp. flabellatum 505 37/0.50 Laserpitium eliasii subsp. thalictrifolium 486 91/1.0 Laserpitium eliasii subsp. eliasii 484 23/− Laserpitium nestleri subsp. nestleri 501 Laserpitium gallicum subsp. gallicum 487 100/1.0 61/0.82 Laserpitium gallicum subsp. paradoxum 490 40/0.68 Laserpitium gallicum subsp. angustissimum 488 80/0.96 Ekimia bornmuelleri 655 24/0.68 Laserpitium gallicum subsp. orospedanum 489 96/1.0 74/1.0 Laserpitium krapfii subsp. gaudinii 497 Laserpitium latifolium 498 94/0.99 Laserpitium petrophilum 509 Ammodaucus leucotrichus 12 Laserpitium krapfii subsp. krapfii 496 Cuminum cyminum 339 100/1.0 99/1.0 Cuminum setifolium 20 Cuminum cyminum G7 Glaucosciadium cordifolium 221 100/1.0 100/1.0 Glaucosciadium insigne 74 Anthriscus sylvestris 83 Ferula communis 195 94/0.99 79/1.0 Ferula olivacea 363 90/0.99 Leutea avicennae 67 0.001 Daucus I Macaronesian Daucus s.l. Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Daucus II Agrocharis Silphiodaucus Laser Orlaya Siler Thapsia Laserpitium s.str. Cuminum Outgroup Version of Record 569 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae exclusion of these accessions from the matrix, we could not reject the null hypothesis of congruence of all four molecular markers (P = 0.199). ModelGenerator selected the GTR + G and GTR + G + I models of nucleotide substitution for small (#1) and large (#3) ITS matrices, respectively. PartitionFinder recognized all three cpDNA markers as a single partition and returned the GTR + G + I model. When all markers were combined, ITS was recognized as a separate partition with the SYM + G + I model. These models and partitions were applied in subsequent BI analyses. In ML analyses, these partitions were analysed with the GTR + G model. The data for this study have been deposited with TreeBASE, study No. 18012. Phylogenetic analyses of separate nrDNA and cpDNA data. — Phylogenetic analyses of small ITS matrix (#1) and cpDNA matrix (#2) with the same accessions resulted in phylogenetic trees with similar major clades, while the discrepancies occurred mostly within those clades or in their relationships to each other (Figs. 1, 2). The species of former Laserpitieae did not form a monophyletic group but most of them constituted a paraphyletic group with respect to the clade of spiny-fruited members of Daucinae, and some winged-fruited species were nested within the latter. Several major groups were apparent within the ITS tree (Fig. 1); with the exception of the Ekimia group, these clades also occurred in the cpDNA tree (Fig. 2). The relationships among these clades were equivocal and the internal support for some of them was moderate. Several species of Laserpitium including L. gallicum L., the nomenclatural type of the genus, formed the Laserpitium s.str. clade that occurred in both the ITS and cpDNA trees with low BS (60% and 24%) and PP support (< 0.5 and 0.68). The relationships among the members of this clade differed between the ITS and cpDNA trees. Two accessions of L. gallicum were identified as introducing topological conflict, i.e., in each tree their position was different and highly supported with BS > 75% and PP = 1.0. Additionally, the accession of L. krapfii Crantz subsp. krapfii was included in this clade only in the nrDNA tree but not in the cpDNA tree. However, its position did not receive high support in either of these trees and, therefore, this terminal was not identified as introducing significant topological conflict. Two Anatolian endemics, Ekimia bornmuelleri and Laserpitium petrophilum Boiss. & Heldr., formed a highly supported Ekimia clade in the ITS trees (BS = 100%, PP = 1.0) while in the cpDNA trees they were placed in separate branches. However, this alternative placement was not highly supported either by ML bootstrap value or by high posterior probability from Bayesian inference. The species of Thapsia, including former members of Distichoselinum García Martín & S.Silvestre, Elaeoselinum, Guillonea, Margotia, and two species of Laserpitium, L. eliasii Sennen & Pau and L. nestleri Soy.-Will., formed the Thapsia clade. It is noteworthy that Ammodaucus leucotrichus Coss. & Durieu, which had been included in this group in previous analyses of ITS data (Weitzel & al., 2014), did not group with Thapsia in the trees inferred from the small ITS matrix (#1) and from the cpDNA data (#2). This species was placed in Thapsia 570 only in trees obtained from the analyses of the large ITS matrix (#3, Electr. Suppl.: Fig. S1). Within the Thapsia clade, several species took notably different positions in the ITS and cpDNA trees but the support for the entire clade was high (BS > 80%, PP = 1.0) with the exception of the tree from the analyses of all ITS sequence data (Electr. Suppl.: Fig. S1). In this tree, with Ammodaucus leucotrichus included in the Thapsia clade, its bootstrap support was 30% (as compared to 82% in small ITS tree) and its posterior probability was 0.78 (as compared to 1.0 in small ITS tree). The accessions of Cuminum cyminum L. and C. setifolium (Boiss.) Koso-Pol. formed a highly supported clade (BS = 99 or 100%, PP = 1.0) with ambiguous affinities. In the cpDNA trees, this genus was sister to the remaining members of the ingroup and this position received high support from ML bootstrap and BI analyses: BS = 94% and PP = 0.99 for the ingroup and BS = 80% and PP = 0.96 for the sister group relationship of Cuminum to the rest of the ingroup. In the ITS trees, Cuminum was sister to the Laser clade but this relationship had little support (BS = 13%, PP < 0.5). The Laser clade encompassed a monospecific genus Laser and some species of Polylophium and Laserpitium. It was highly supported in the cpDNA trees (BS = 93%, PP = 1.0) but poorly in the ITS trees (BS = 47%, PP < 0.5). In the latter, it also included Ammodaucus leucotrichus. In the analyses of the large ITS dataset (#3), with Ammodaucus Coss. & Durieu placed outside this clade, its internal support was higher (BS = 55%, PP = 0.95). All accessions of Laserpitium siler formed a clade supported in ITS (BS = 100%, PP = 1.0) and cpDNA trees (BS = 62%, PP = 0.99). Because this species was once placed in the separate genus Siler, and it is the generitype of this name, we denote this branch as the Siler clade. A single accession of Laserpitium pseudomeum Orph., Heldr. & Sart. ex Boiss. constituted a separate lineage and its position differed between the cpDNA and ITS trees. The monophyly of Orlaya Hoffm. was strongly supported in both the ITS (BS = 100%, PP = 1.0) and cpDNA trees (BS = 99%, PP = 1.0). This genus with spiny fruits was, however, placed separate from other spiny-fruited taxa: Agrocharis Hochst. and Daucus. The core Daucinae were highly supported in all trees (BS ≥ 90%, PP = 1.0). This clade included the Daucus s.l. clade and the Silphiodaucus clade. The former comprised four groups identified in previous molecular studies: Agrocharis, Daucus I subclade, Daucus II subclade and the Macaronesian endemic group (Lee & Downie, 1999, 2000; Spalik & Downie, 2007). The Silphiodaucus clade included two species of Laserpitium, L. prutenicum and L. hispidum, that were placed by KosoPoljansky (1916) in Daucus sect. Silphiodaucus. The monophyly of this clade and its sister position to the Daucus s.l. clade was strongly supported in all analyses (BS > 85%, PP > 0.95). The species of Daucus were placed in two distinct subclades that received various support: low for the Daucus I group (BS = 54%, PP = 0.78 in ITS analyses and BS = 6%, PP < 0.5 in cpDNA analyses) and high for the Daucus II group (BS = 100%, PP = 1.0 in ITS analyses and BS = 79%, PP = 1.0 in cpDNA analyses). A group of three Macaronesian endemics Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae was paraphyletic with respect to the Daucus I subclade with the two winged-fruited genera, Monizia and Melanoselinum, forming a highly supported branch. Cryptotaenia elegans Webb ex Bolle, endemic to the Canary Islands, was placed either in a polytomy or sister to the Daucus I subclade. The latter also included two other genera of former Laserpitieae: Tornabenea and Rouya. The species of Tornabenea was placed within the D. carota complex and this relationship received strong support in both the cpDNA and ITS data analyses. The placement of Athamanta dellacellae Asch. & Barbey ex E.A.Durand & Baratte in the Daucus I subclade, previously inferred based on ITS data only, has been confirmed with cpDNA data. The monophyly of Agrocharis and its sister position to the Daucus II subclade were strongly supported in all analyses. Phylogenetic analyses of combined data. — ML and BI analyses of combined nrDNA ITS and cpDNA sequences (dataset #4) were performed with and without the terminals introducing topological conflict that had been identified with compat.py. Additionally, to assess the impact of missing data on tree topology, we repeated the analyses for a subset of accessions for which both ITS and cpDNA sequences (rpoC1 intron and/or rps16 intron) were available (dataset #5). The topologies of trees from all analyses were generally similar with the same major clades identified. Because our major aim is to delimit major clades and provide a new taxonomic treatment for Daucinae, we here present the results of the analyses of all data (matrix #4 with all terminals, Figs. 3, 4), while the trees obtained from the reduced matrices (matrix #4 without the terminals introducing topological conflicts and matrix #5) are provided in the Electronic Supplement (Figs. S2 and S3, respectively). In the combined analyses, Cuminum was placed in a sister position to the remaining Daucinae (Fig. 3). However, this relationship and most relationships among the major clades received very weak BS and PP support. Ammodaucus formed a separate branch with no direct relationship to Thapsia or Laser as in the previous analyses of nrDNA ITS data. The Ekimia clade received high support (BS = 100%, PP = 1.0) and it also included Laserpitium glaucum Post, another Anatolian endemic, for which only the ITS sequence was available. The Laserpitium s.str. clade was well supported in ML analyses (BS = 85%) and in Bayesian inference (PP = 1.0). All accessions of L. gallicum formed a clade sister to L. halleri Crantz. Two subspecies of L. krapfii were not placed as sister taxa but subsp. gaudinii was grouped with L. nitidum Zanted. and L. peucedanoides L., and this relationship was strongly supported (BS = 97%, PP = 1.0). Several species of Thapsia that were represented only by ITS data grouped within the Thapsia clade, and the relationships among them were generally well resolved. As in the previous analyses, L. nestleri and L. eliasii were placed within this group. Noteworthy is the distinct position of Thapsia sp. 471, an accession from Corsica of uncertain specific affinity. It was provisionally classified as Elaeoselinum asclepium subsp. meoides (Desf.) Fiori (Reduron, 2007b) or Thapsia meoides Guss. but in our analyses it did not group with conspecific accessions. The combined analyses confirmed the isolated positions of two species of Laserpitium, L. siler and L. pseudomeum, which are not related to their congeners. These analyses also supported the sister position of the Silphiodaucus clade to the Daucus s.l. clade. The relationships among the major subclades within the Daucus s.l. clade were well resolved and highly supported (Fig. 4). They were generally similar to those inferred from the nrDNA ITS analyses. As before, Cryptotaenia elegans was placed in sister position to the Daucus I subclade rather than to the two endemics from Madeira. Within the Daucus I subclade, conspecific accessions usually grouped together, confirming the current delineation of these species. The exception was the D. carota complex that also included three species of Tornabenea and Daucus virgatus (Poir.) Maire. Similar to separate analyses, Rouya polygama (Desf.) Coincy was closely related to D. carota. A distinct group nested within the Daucus I subclade was formed by two species of Pseudorlaya (Murb.) Murb. Athamanta dellacellae was sister to Pachyctenium mirabile Maire & Pamp, the only species of Pachyctenium Maire & Pamp. Within the Daucus II subclade, two taxa that are usually treated as conspecific, D. broteri Ten. and D. bicolor Sibth. & Sm., were distantly related. The recently described D. conchitae Greuter was placed sister to D. involucratus Sm. with very high support (BS = 99%, PP = 1.0). Morphological analyses. — The morphology of secondary ribs was coded as a discrete unordered character with four states (secondary ribs: obsolete, winged, spiny, keeled). Maximum parsimony mapping of this character onto a pruned cladogram from the ML analyses of combined data inferred 14 character changes (Fig. 5). The reconstruction of the ancestral condition was ambiguous because all character states occurred at basal branches. Throughout the evolutionary history of the group, winged fruits dominated and fruits with spiny secondary ribs are generally derived from the former. However, in the spiny-fruited clade of Daucinae (encompassing the Daucus s.l. clade, the Silphiodaucus clade and Orlaya) the ancestral condition was ambiguous: winged or spiny. Therefore, spiny fruits in Orlaya, the Daucus I subclade and the Daucus II plus Agrocharis subclade may have arisen independently from winged fruits; alternatively, winged fruits in the Silphiodaucus clade and in the Madeiran endemics subclade are derived from spiny fruits. In contrast, winged fruits in some members of the Daucus I subclade represent evolutionary reversions. DISCUSSION The incongruence between plastid and nuclear data. — Incongruence among molecular markers may result from diverse evolutionary processes such as hybridization, introgression and incomplete lineage sorting. Alternatively, it may be an analytical artefact resulting from lack of phylogenetic signal, i.e., missing data and possible sampling error of characters, taxa, or both (Salichos & al., 2014, and references therein). Discrepancy between cpDNA and nrDNA ITS data has been reported in several studies of Apiaceae (e.g., Lee & Downie, 2006; Spalik & al., 2009; Zhou & al., 2009; Bone & al., 2011; Yi & al., 2015). In this study, 13 accessions Version of Record 571 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Daucus s.l. clade 86/1.0 Laserpitium hispidum G65 98/1.0 Laserpitium hispidum G66 Laserpitium hispidum 495 99/1.0 Laserpitium hispidum G64 Laserpitium hispidum 494 100/1.0 100/1.0 Laserpitium prutenicum subsp. prutenicum G69 Laserpitium prutenicum subsp. prutenicum 649 95/0.95 Laserpitium prutenicum subsp. duforianum 511 72/0.99 Orlaya daucoides G79 72/0.99 Orlaya daucoides 264 98/1.0 Orlaya daucoides G111 Orlaya daucoides 229 99/1.0 Orlaya daucoides G75 Orlaya daucorlaya G76 100/1.0 87/0.93 Orlaya daucoides 263 Orlaya grandiflora 258 100/1.0 Orlaya grandiflora 259 G78 Orlaya grandiflora G77 Polylophium panjutinii G84 55/0.97 99/1.0 Polylophium panjutinii G83 100/1.0 Polylophium panjutinii G82 87/1.0 Polylophium involucratum 395 G81 100/1.0 Laserpitium stevenii 518 34/0.67 Laserpitium stevenii G71 Laserpitium archangelica 481 55/1.0 100/0.98 Laser trilobum 611 98/1.0 Laser trilobum G112 99/1.0 49/0.80 Laser trilobum G61 100/1.0 Laserpitium carduchorum 482 Laserpitium carduchorum 483 59/1.0 35/0.52 100/1.0 Laserpitium affine G62 Laserpitium affine 480 Laserpitium pseudomeum 513 4 /− Laserpitium siler 515 Laserpitium siler 516 32/− Laserpitium siler 517 100/1.0 Laserpitium siler G113 Laserpitium siler G70 Thapsia garganica G128 94/− Thapsia garganica 525 100/1.0 Thapsia garganica G92 Thapsia garganica 526 100/1.0 Thapsia garganica 527 Thapsia gymnesica G115 100/1.0 Thapsia platycarpa G93 Thapsia transtagana G117 100/1.0 Thapsia transtagana G94 Thapsia transtagana G116 Thapsia transtagana 524 Fig. 3. Maximum likelihood tree 82/0.92 Thapsia smittii G120 82/0.99 Thapsia smittii G119 inferred from analyses of combined 96/1.0 Thapsia meoides G59 9/− nrDNA ITS and cpDNA data for 260 Thapsia meoides 473 52/0.52 Thapsia meoides 472 accessions (241 terminals) representing Thapsia thapsioides 477 85/0.87 90/1.0 84/1.0 89/1.0 Thapsia thapsioides 478 Daucinae and outgroups (dataset #4, Thapsia sp. 471 87/1.0 Thapsia gummifera 520 see the text for details). The Daucus s.l. Thapsia tenuifolia 469 100/1.0 Thapsia asclepium 470 clade is marked as a single terminal and Thapsia asclepium G121 89/1.0 Thapsia minor G129 illustrated in detail in Fig. 4. Bootstrap 77/0.98 Thapsia minor G128 69/0.94 Thapsia minor G131 support and posterior probability (for Thapsia minor G130 98/1.0 Thapsia villosa G126 nodes that also occurred in Bayesian 66/0.91 Thapsia villosa 530 G127 99/1.0 Thapsia villosa G125 majority-rule consensus tree) are given Thapsia villosa G124 100/1.0 along the branches. Major groups are Thapsia villosa var. dissecta 529 Thapsia laciniata G132 G133 100/1.0 bracketed. Thapsia villosa var. dissecta 528 99/1.0 Thapsia villosa G95 Thapsia maxima G122 G123 /− Thapsia scabra 479 92/1.0 96/0.99 Laserpitium nestleri subsp. flabellatum 505 99/1.0 48/0.71 Laserpitium nestleri subsp. lainzii 506 Laserpitium nestleri subsp. nestleri 501 100/1.0 Laserpitium eliasii subsp. ordunae 485 Laserpitium eliasii subsp. eliasii 484 99/1.0 Laserpitium eliasii subsp. thalictrifolium 486 95/0.94 Laserpitium nitidum 507 97/1.0 Laserpitium peucedanoides 510 73/0.95 Laserpitium krapfii subsp. gaudinii 497 92/1.0 Laserpitium latifolium 498 Laserpitium krapfii subsp. krapfii 496 Laserpitium gallicum subsp. gallicum 487 100/1.0 85/1.0 41/− 99/1.0 Laserpitium gallicum subsp. paradoxum 490 96/1.0 Laserpitium gallicum subsp. angustissimum 488 96/1.0 Laserpitium gallicum subsp. orospedanum 489 Laserpitium halleri 493 Laserpitium petrophilum G87 72/0.86 Laserpitium petrophilum G88 Laserpitium petrophilum G86 Laserpitium petrophilum 508 G85 100/1.0 Laserpitium petrophilum G67 96/1.0 Laserpitium petrophilum 509 100/1.0 100/1.0 Laserpitium glaucum 491 Ekimia bornmuelleri 655 100/1.0 Ammodaucus leucotrichus 12 Ammodaucus leucotrichus G114 Cuminum cyminum G9 63/0.99 Cuminum cyminum G11 G13 G8 Cuminum cyminum G12 74/0.97 Cuminum cyminum G10 Cuminum cyminum 339 96/0.99 Cuminum setifolium 20 100/1.0 Cuminum setifolium G6 Cuminum cyminum G7 96/1.0 Athamanta cretensis 617 94/1.0 Conopodium arvense 630 97/1.0 Anthriscus sylvestris 83 Glaucosciadium insigne 74 100/1.0 100/1.0 Glaucosciadium cordifolium 221 84/1.0 Ferula olivacea 363 99/1.0 Ferula communis 195 Leutea avicennae 67 0.001 100/1.0 Silphiodaucus Orlaya Laser Siler Thapsia Laserpitium s.str. Ekimia Cuminum Outgroup 572 Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Fig. 4. The relationships among the representatives of the Daucus s.l. clade inferred from maximum likelihood analyses of combined nrDNA ITS and cpDNA data. See Fig. 3 for details. The asterisk indicates a sequence that was identical to three other sequences obtained from Daucus carota subsp. maximus 269 and 449, and Daucus virgatus 50. Daucus carota subsp. commutatus 3 Daucus carota subsp. carota G23 Daucus carota subsp. capillifolius G21 Daucus carota subsp. maximus G31 Daucus carota subsp. hispanicus var. linearis 38 57/0.92 Daucus carota subsp. maximus 450 Daucus carota subsp. carota 443 444 Daucus carota subsp. maritimus 448 1/− Daucus carota subsp. carota G98 Daucus carota subsp. maximus 451 3/− Daucus carota subsp. carota G25 Daucus carota subsp. carota 442 Daucus carota subsp. maximus 274 65/0.74 Daucus virgatus 659 Daucus carota subsp. carota 32 Daucus carota subsp. hispanicus G30 Daucus carota subsp. hispanicus 35 Daucus carota subsp. hispanicus 447 1 /− Daucus carota subsp. drepanensis G26 100/1.0 Tornabenea insularis G106 Daucus carota subsp. gadecaei 34 84/0.98 Tornabenea annua G96 65/0.98 Tornabenea annua 521 Tornabenea tenuissima G97 Daucus carota subsp. gummifer 445 Daucus carota subsp. gummifer G28 99/1.0 Daucus carota subsp. azoricus G24 Daucus carota subsp. gadecaei G27 Daucus carota subsp. halophilus G29 Daucus carota subsp. carota G22 62/0.91 Daucus carota subsp. azoricus 610 Daucus carota subsp. halophilus 9 52/0.88 94/0.96 Daucus syrticus 265 100/1.0 Daucus syrticus 468 Daucus syrticus G57 56/0.86 100/1.0 Daucus gracilis G41 Daucus gracilis 270 58/0.92 81/0.91 Daucus sahariensis G54 100/1.0 Rouya polygama G107 Rouya polygama 523 86/1.0 Daucus biseriatus G89 Pseudorlaya minuscula G90 100/1.0 Pseudorlaya pumila G108 82/1.0 Pseudorlaya pumila 227 83/0.99 Pseudorlaya pumila 228 G91 100/1.0 Pachyctenium mirabile 244 Pachyctenium mirabile G80 85/0.99 Athamanta dellacellae 435 87/1.0 86/1.0 Daucus aureus 273 439 Daucus aureus G18 87/0.98 Daucus aureus G19 100/1.0 Daucus aureus G16 100/1.0 Daucus aureus G17 79/1.0 100/1.0 Daucus mauritii G46 Daucus mauritii G45 93/0.92 Daucus setifolius 237 Daucus setifolius G56 100/1.0 Daucus setifolius G55 95/1.0 Daucus setifolius 467 100/1.0 Daucus crinitus 14 86/0.96 Daucus crinitus G33 G34 65/0.98 Daucus muricatus G49 Daucus muricatus G50 63/0.76 Daucus muricatus G51 Daucus muricatus 281 100/1.0 67/0.74 Daucus muricatus 271 Daucus muricatus G48 70/0.96 Daucus tenuisectus 266 100/1.0 Daucus tenuisectus 36 Daucus tenuisectus G58 100/1.0 Cryptotaenia elegans G105 G99 100/1.0 Cryptotaenia elegans 39 Melanoselinum decipiens 522 613 30/− Melanoselinum decipiens G72 Melanoselinum decipiens 612 95/1.0 Melanoselinum decipiens G109 96/1.0 Melanoselinum decipiens G73 edulis G74 97/1.0 Monizia Monizia edulis 13 Monizia edulis G110 100/1.0 Daucus bicolor G42 99/1.0 Daucus bicolor 41 100/1.0 Daucus littoralis 461 100/1.0 Daucus littoralis 8 46/0.62 Daucus littoralis 275 99/0.97 Daucus guttatus 458 100/1.0 Daucus guttatus 238 Daucus guttatus G43 49/0.69 93/0.99 Daucus arcanus G15 63/0.85 Daucus arcanus G14 Daucus arcanus 260 Daucus pusillus 267 100/1.0 Daucus pusillus G53 Daucus pusillus G52 64/0.96 Daucus pusillus 466 95/1.0 Daucus involucratus 272 Daucus involucratus G44 79/1.0 Daucus involucratus 460 99/1.0 Daucus involucratus 37 72/0.98 Daucus conchitae 236 99/1.0 Daucus conchitae G32 90/1.0 Daucus broteri 440 100/1.0 Daucus broteri 277 100/1.0 Daucus broteri 276 Daucus montanus 262 95/1.0 Daucus montanus 230 Daucus montanus G47 90/0.58 Daucus montanus 17 95/1.0 Daucus glochidiatus 456 64/0.98 96/1.0 Daucus glochidiatus 455 Daucus glochidiatus G40 Daucus glochidiatus G39 64/0.90 Daucus durieua G38 100/1.0 100/1.0 Daucus durieua 453 100/1.0 Daucus durieua G37 durieua G36 98/1.0 Daucus Daucus durieua 454 Daucus durieua 40 Agrocharis incognita 46 51/0.70 Agrocharis pedunculata 247 55/0.83 95/1.0 Agrocharis pedunculata G3 Agrocharis pedunculata G4 Agrocharis incognita 245 100/1.0 Agrocharis melanantha 248 Agrocharis melanantha G2 89/1.0 Agrocharis gracilis G1 * Daucus I Macaronesian 100/1.0 Daucus II Agrocharis 0.001 Version of Record 573 Morphology of secondary ribs obsolete winged spiny keeled Daucus carota subsp. maritimus Daucus carota subsp. carota Daucus carota subsp. commutatus Daucus carota subsp. capillifolius Daucus carota subsp. maximus Daucus virgatus Daucus carota subsp. hispanicus Daucus carota subsp. drepanensis Tornabenea insularis Tornabenea tenuissima Tornabenea annua Daucus carota subsp. azoricus Daucus carota subsp. halophilus Daucus carota subsp. gadecaei Daucus carota subsp. gummifer Daucus syrticus Daucus sahariensis Daucus gracilis Rouya polygama Daucus biseriatus Pseudorlaya minuscula Pseudorlaya pumila Athamanta della−cellae Pachyctenium mirabile Daucus mauritii Daucus aureus Daucus crinitus Daucus setifolius Daucus tenuisectus Daucus muricatus Cryptotaenia elegans Monizia edulis Melanoselinum decipiens Daucus littoralis Daucus bicolor Daucus guttatus Daucus pusillus Daucus arcanus Daucus conchitae Daucus involucratus Daucus broteri Daucus glochidiatus Daucus montanus Daucus durieua Agrocharis melanantha Agrocharis gracilis Agrocharis incognita Agrocharis pedunculata Laserpitium prutenicum subsp. duforianum Laserpitium prutenicum subsp. prutenicum Laserpitium hispidum Orlaya daucorlaya Orlaya daucoides Orlaya grandiflora Polylophium involucratum Polylophium panjutinii Laserpitium stevenii Laserpitium archangelica Laser trilobum Laserpitium carduchorum Laserpitium affine Laserpitium pseudomeum Laserpitium siler Laserpitium nestleri subsp. flabellatum Laserpitium nestleri subsp. lainzii Laserpitium nestleri subsp. nestleri Laserpitium eliasii subsp. ordunae Laserpitium eliasii subsp. eliasii Laserpitium eliasii subsp. thalictrifolium Thapsia laciniata Thapsia villosa var. dissecta Thapsia minor Thapsia villosa Thapsia maxima Thapsia scabra Thapsia meoides Thapsia smittii Thapsia thapsioides Thapsia gummifera Thapsia tenuifolia Thapsia platycarpa Thapsia transtagana Thapsia gymnesica Thapsia garganica Thapsia asclepium Laserpitium nitidum Laserpitium peucedanoides Laserpitium krapfii subsp. gaudinii Laserpitium latifolium Laserpitium krapfii subsp. krapfii Laserpitium gallicum subsp. gallicum Laserpitium gallicum subsp. paradoxum Laserpitium gallicum subsp. angustissimum Laserpitium gallicum subsp. orospedanum Laserpitium halleri Laserpitium glaucum Laserpitium petrophilum Ekimia bornmuelleri Ammodaucus leucotrichus Cuminum setifolium Cuminum cyminum Athamanta cretensis Conopodium arvense Anthriscus sylvestris Glaucosciadium insigne Glaucosciadium cordifolium Ferula olivacea Ferula communis Leutea avicennae Daucus I Macaronesian Daucus s.l. TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Daucus II Agrocharis Silphiodaucus Orlaya Laser Thapsia Laserpitium s.str. Ekimia Cuminum Outgroup Fig. 5. Maximum parsimony reconstruction of ancestral secondary rib characters for subtribe Daucinae using an ML tree inferred from combined data. This tree was transformed into a cladogram and pruned such that each taxon was represented by one accession. The morphology of secondary ribs was coded as a multistate (unordered) character with four states: obsolete, winged, spiny, keeled. 574 Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae introducing topological conflict were identified and removal of these terminals from the analyses resulted in a significant decrease in incongruence among the markers. Although these accessions had different positions in ITS and cpDNA trees, they were placed within the same major clades suggesting that this discrepancy may have resulted from reticulation among closely related taxa or from incomplete lineage sorting. Five of them are members of the Thapsia clade that comprises diploid and polyploid species; the latter have been recorded in the T. villosa L. complex and may represent allopolyploids (Avato & al., 1996). Among the species of Daucus, polyploids include tetraploid Australian D. glochidiatus (Labill.) Fisch. & al. and hexaploid South American D. montanus Humb. & Bonpl. (Grzebelus & al., 2011). These species are closely related in our analyses (Fig. 4). The homoplasy of fruits with winged secondary ribs. — The fruits of umbellifers are schizocarps that, despite their overall similarity, vary with respect to both their external features, e.g., their shape and compression, the type of appendages and ribs, as well as anatomical characters, including the number and size of vittae, the shape of the endosperm, the form of fruit sclerification, etc. These characters served as basis for nearly all traditional classification systems of the family including the most influential system by Drude (1897–1898). This system had remained almost intact throughout the 20th century until the first molecular phylogenetic studies demonstrated that nearly all tribes recognized by Drude are polyphyletic (Downie & Katz-Downie, 1996; Downie & al., 1996; Plunkett & al., 1996a, b). Subsequent studies of fruit evolution based on molecular phylogenies confirmed that fruit characters in Apiaceae are homoplastic, although less so than other morphological features (Lee & al., 2001; Spalik & Downie, 2001; Spalik & al., 2001; Calviño & al., 2008). Our study unequivocally demonstrates that umbellifers with secondary ribs projecting into wings do not constitute a single lineage: the majority of species form a grade at the base of Daucinae while some are nested within the clade of spiny-fruited taxa. Therefore, winged fruits are of limited value for the identification of monophyletic groups. Similarly, spiny fruits are homoplastic due to several reversals to winged fruits and they should be used with caution for delimiting genera. In particular, the traditional delineation of Daucus and Laserpitium, the two largest genera of Daucinae, does not correspond to the phylogenetic relationships in the subtribe. The phylogeny and taxonomy of Daucus. — The cultivated carrot, Daucus carota subsp. sativus, is one of the most important crops worldwide, and wild species of Daucus constitute an invaluable gene pool for improving its cultivars (Grzebelus & al., 2011). A successful example for this was the introduction of resistance to carrot fly (Chamaepsila rosae), the most devastating pest of carrot, parsnip, parsley and celery, from wild D. carota subsp. capillifolius (Gilli) Arbizu (Ellis & al., 1993). The latter taxon was formerly recognized as a separate species but included in D. carota based on the analyses of phylogenomic and morphological data (Arbizu & al., 2014a). This case shows the importance of a phylogeny-based taxonomy of Daucus for plant breeders. The separation of Daucus into two distinct subclades, subsequently named Daucus I and II, was inferred from the analyses of nrDNA ITS (Lee & Downie, 1999) and reassessed with cpDNA data (Lee & Downie, 2000). These studies demonstrated that the Daucus I subclade also includes the representatives of Pachyctenium and Pseudorlaya, while the Daucus II subclade is sister to Agrocharis. Successive analyses with extended taxonomic sampling added Athamanta dellacellae and two species of Tornabenea to the Daucus I subclade and indicated that this group is sister to a clade of Macaronesian endemics (Spalik & Downie, 2007). The subclades of Daucus received very strong support from phylogenomic studies (Arbizu & al., 2014b); however, these analyses did not include representatives of the Macaronesian endemics group and Agrocharis, i.e., sister groups to Daucus I and II subclades, respectively. Our study reinforces these previous findings and demonstrates the need for a new taxonomic delineation of Daucus. However, the reconciliation of the taxonomy of Daucus with its phylogeny is not an easy task because the clades inferred from molecular data lack obvious morphological synapomorphies that facilitate the recognition of their members. Canonical variate analysis and hierarchical cluster analysis based on 40 morphological characters scored for representatives of both Daucus clades failed to separate these groups; moreover, none of the characters was diagnostic (Arbizu & al., 2014a). Similarly, the study of fruit anatomy and morphology of Daucinae did not provide any distinctive features for these two lineages (A. Wojewódzka & K. Spalik, unpub. data). In the absence of good diagnostic characters discriminating those two clades, a broad concept of Daucus should be considered. In order to maintain the monophyly of the genus, spiny-fruited genera Agrocharis and Pseudorlaya and the winged-fruited genera Melanoselinum, Monizia, Pachyctenium, Rouya and Tornabenea must be sunk into synonymy of Daucus. We propose to recognize the major clades within Daucus s.l. as sections of the genus. The Daucus I clade includes the nomenclatural type of the genus and therefore constitutes sect. Daucus. The members of the Daucus II clade were traditionally placed in three sections: sect. Daucus, sect. Platyspermum (Hoffm.) DC. and sect. Anisactis DC. (Sáenz Laín, 1981; Heywood, 1982). The genus Platyspermum was described by Hoffmann (1814) based on a single species, D. muricatus L., which is now placed in the redefined sect. Daucus; sect. Platyspermum is, therefore, a taxonomic synonym of sect. Daucus. Section Anisactis was described by Candolle (1830) with four names: D. brachiatus Sieber ex DC. (= D. glochidiatus), D. toriloides DC. (= D. montanus), D. montevidensis Link ex Spreng. (= D. pusillus Michx.), and D. australis Poepp. ex DC. (= D. montanus). In our analyses, all these taxa are placed in the Daucus II clade; therefore, we formally apply the name Daucus sect. Anisactis to this clade, and designate D. brachiatus as the nomenclatural type of this section. The names of the two remaining sections of Daucus are based on the generic names Agrocharis and Melanoselinum, which are here reduced to synonymy under Daucus. Rouya. — This genus includes one species, Rouya polygama (Desf.) Coincy that was described in Thapsia with the phrase name indicating finely divided leaves as its diagnostic Version of Record 575 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae character (Desfontaines, 1798). It occurs in littoral habitats in the southwest of the Mediterranean region (Algeria, Tunisia) and on the main islands (Corse, Sardinia). In habit it resembles some littoral species of Daucus, but differs in having winged rather than spiny fruits (Reduron, 2008). Our results unanimously place this species in Daucus and support its formal transfer to this genus. The epithet polygamus is not available in Daucus because it is occupied by D. polygamus Gouan (1773), an unresolved name typified by an illustration in Boccone (1674) that may represent D. carota subsp. drepanensis. However, both its locus classicus and the only extant herbarium specimen suggest that the name is synonymous with D. bicolor (Reduron, 2007a). With the new epithet, rouyi, we acknowledge the intention of Coincy, who dedicated the genus to Georges C.C. Rouy, an eminent French botanist and the principal author of the most exhaustive 14-volume Flore de France. Pseudorlaya. — This genus comprises two annual species, P. pumila (L.) Grande and P. minuscula (Pau ex Font Quer) Laínz, that are morphologically similar to each other (Rutherford & Jury, 2003). The genus supposedly differs from the traditionally delimited Daucus in having two rows of spines instead of one on secondary ridges; however, this character is variable in P. pumila (Lee & al., 2001). Molecular analyses confirmed the close relationship between the two species and unanimously indicated that the genus is nested within Daucus; therefore, Lee & Park (2014) suggested that it should be included in the latter. The names for these species in Daucus are therefore D. pumilus (L.) Hoffmanns. & Link and D. minusculus Pau ex Font Quer. Cryptotaenia, Athamanta and Pachyctenium. — Two species, Athamanta dellacellae and Cryptotaenia elegans, that are placed in the Daucus I clade lack spines or wings on secondary ridges. Therefore, their placement in Daucinae came as a surprise. They are incorrectly classified in their respective genera. Athamanta dellacellae has hairy, oblong fruits with a short beak that are superficially similar to those of its congeners; the genus Athamanta L., however, is now placed in Scandiceae subtribe Scandicinae (Spalik & al., 2001). The sister species to A. dellacellae is Pachyctenium mirabile and this affinity is supported by their geographic distribution: both are endemic to the northern African region of Cyrenaica (Qu6zel, 1978). The placement of Pachyctenium in Daucus had already been postulated based on phylogenetic analyses of morphological data (Lee & al., 2001). An endemic of the Canary Islands, Cryptotaenia elegans has been retained in this genus due to a lack of alternative placement rather than its similarity to distantly related congeners (Spalik & Downie, 2007). Its fruits are glabrous and have no secondary appendages, which is exceptional in Daucinae. Because spiny fruits seem to be plesiomorphic for the Daucus clade, Spalik & Downie (2007) speculated that in the absence of dispersing agents, like terrestrial mammals, this insular descendant of epizoochorous species has switched to gravity dispersal. The winged-fruited endemics of Macaronesia. — Monizia and Melanoselinum are two monospecific endemics of Madeira characterised by woody stems crowned by a rosette of leaves. Monizia was once included in Melanoselinum along with the 576 species of Tornabenea (Press & Dias, 1998; Fernandes & Carvalho, 2014). The last genus includes up to six endemics of Cape Verde that are characterised by fruits with narrowly winged secondary ridges with reduced spines. Two species, T. bischoffii J.A.Schmidt and T. tenuissima (A.Chev.) A.Hansen & Sunding, have slender woody stems at flowering time that make them superficially similar to the Madeiran endemics; however, a relationship of Tornabenea to Daucus has also been suggested (Brochmann & al., 1997). The results of the molecular phylogenetic studies place the three examined species of Tornabenea within the D. carota complex. Moreover, experimental crosses between Daucus carota subsp. sativus (Hoffm.) Arcang. and Tornabenea tenuissima (A.Chev.) A.Hansen & Sunding successfully demonstrated that there is no crossing barrier between those species (J.-P. Reduron, unpub. data). The phylogenetic position of the remaining congeners, particularly the woody species, awaits further study. Recent morphological studies demonstrated that Monizia edulis comprises four distinct morphotypes deserving taxonomic recognition at the rank of subspecies (Fernandes & Carvalho, 2014). In fact, morphological differences among those populations are similar to those among the morphological forms of Tornabenea that are recognized as separate species (Brochmann & al., 1997). Apparently, detailed morphological and molecular studies are required to assess the status of those insular taxa of Daucus. Agrocharis. — With its spiny fruits, Agrocharis is similar to Daucus (Lee & al., 2001). The genus comprises up to four species that are distributed in tropical regions of Africa (Townsend, 1989). Until it was revised by Heywood (1973, 1982), most of its species had been recognized in Caucalis L. Based on the molecular phylogeny of Agrocharis and allied genera, Lee (2002) discussed their morphological delimitation and suggested its inclusion in Daucus at the rank of subgenus or section, contingent upon further taxonomic and molecular sampling. Our results justify a formal transfer of Agrocharis to Daucus. We refrain from a formal recognition of A. gracilis Hook.f. in Daucus because this species was only represented by one ITS sequence that was identical to that of A. melanantha Hochst. The Silphiodaucus clade. — Two species of Laserpitium— L. prutenicum and L. hispidum—form the clade sister to the Daucus s.l. clade. Although these species were traditionally placed in Laserpitium due to their winged fruits, their similarity to Daucus had also been noticed. Koso-Poljansky (1916) included these two species in Daucus as D. prutenicus (L.) E.H.L.Krause (= L. prutenicum) and D. pilosus (Willd.) KosoPol. (= L. hispidum) in sect. Silphiodaucus Koso-Pol. He also placed L. latifolium L. in this section, but this transfer is not supported by molecular data. With their finely divided leaves and prominent indumentum, L. prutenicum and L. hispidum resemble the species of Daucus, while their winged fruits are somewhat similar to those of Laserpitium. We propose to recognize this clade as a new genus based on the section described by Koso-Poljansky. Since this author did not indicate its type, we select D. prutenicus as the nomenclatural type of the section and, consecutively, the genus. Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae The Laser clade. — This clade comprises representatives of three genera, Laser, Laserpitium and Polylophium. Laser is presently recognized as having one species, L. trilobum (Pimenov & Leonov, 1993), distributed in Europe and western Asia (Schischkin, 1951a; GBIF, 2013). The fruits of this species are used in traditional medicine and as a condiment with confirmed antimicrobial activity (Parlatan & al., 2009). The other members of this clade have rather narrow distributions. Laserpitium stevenii Fisch. & al., L. affine Ledeb. and Polylophium panjutinii Manden. & Schischk. are all endemics of the Caucasus (Schischkin, 1951b, c), L. carduchorum Hedge & Lamond and P. involucratum (Pall.) Boiss. are IranoTuranian elements (Hedge & Lamond, 1972; Rechinger, 1987c), and L. archangelica Wulfen occurs in the southeastern part of Central Europe. This clade is supported in molecular analyses and we propose to recognize it as a single genus. Of the two available names, Laser Borkh. ex G.Gaertn. & al. 1799 has priority over Polylophium Boiss. 1844. Thapsia. — The phylogeny of Thapsia has recently been examined based on ITS sequence variation, resulting in the restoration of a broad Linnaean concept of the genus (Weitzel & al., 2014). Its former segregates, Elaeoselinum, Distichoselinum and Margotia, as well as monospecific Guillonea have been sunk into synonymy. The species of Thapsia s.l. are characterized by relatively large fruits with broad wings. The clade encompasses species of predominantly western Mediterranean distribution, with the centre of endemism in the Iberian Peninsula. Our results support such a definition of the genus with the addition of two other Iberian endemics, Laserpitium nestleri and L. eliasii. These species have fruits with relatively small lateral wings that make their fruits externally more similar to the species of Laserpitium rather than to the broad-winged fruits of Thapsia. However, the fruits of L. nestleri and L. eliasii have distinct extravallecular vittae that are obsolete in other congeners but occur in some species of Thapsia s.l. (Arenas Posada & García Martín, 1993). Both species include several subspecies each (Montserrat, 2003b, c). Some of them were included in our analyses but they were represented only by single accessions. Within each of these species, there seems to be rather low sequence variation suggesting very close relationships among their inclusive taxa. More detailed morphological and molecular studies are required to ascertain whether the recognition of these subspecies is justified. Within Thapsia, there are several species aggregates, including diploid, tetraploid and hexaploid forms that are morphologically hardly distinguishable. However, they are distinguishable based on chemotaxonomy and nrDNA ITS sequences, and some of these chemotypes deserve taxonomic recognition. Recently, one of these was described as a new species, T. smittii Simonsen & al., which is a segregate of T. maxima Mill. (Weitzel & al., 2014). A detailed morphological and molecular study may unravel new species. The accession from Corsica (No. 471), provisionally determined as T. meoides, may represent such a case based on its placement in our molecular trees, where it stands separate from presumably conspecific accessions. The Ekimia clade. — Ekimia bornmuelleri was originally described in Prangos and later transferred to a separate genus of presumed affinity to Prangos (Duman & Watson, 1999). However, a detailed comparison of fruit anatomy of Ekimia and its presumed relatives (two species of Prangos, P. ferulacea Lindl. and P. lophoptera Boiss., and an Anatolian representative of Laserpitium, L. petrophilum) demonstrated that Ekimia shares several characters with Laserpitium petrophilum (Lyskov & al., 2015). Its position in Daucinae is therefore firmly established both by molecular and morphological data. The three members of the Ekimia clade are eastern Mediterranean elements but have allopatric distributions: Ekimia occurs in southwestern Turkey (Özhatay & al., 2008), L. petrophilum inhabits mountains of central southern Anatolia (Hedge & Lamond, 1972), while L. glaucum was described from the Nur (Amanus) Mountains in southeastern Turkey (Post, 1891). Laserpitium petrophilum was also recognized in Polylophium (Pimenov & Leonov, 2004), but our molecular analyses do not confirm this relationship. We propose to include L. petrophilum and L. glaucum in Ekimia. Laserpitium s.str. — Six species, including Laserpitium gallicum with four subspecies, and L. krapfii with two subspecies, are placed in the Laserpitium s.str. clade. This clade is stable and supported in all molecular analyses although only in combined analyses this support is high. Moreover, it is supported by fruit morphology and anatomy: its members differ from former congeners in having non-elongated sclerified cells in the endocarp (A. Wojewódzka & K. Spalik, unpub. data). Because this clade includes the nomenclatural type of the genus, L. gallicum, it retains the name Laserpitium. Unstable and isolated lineages. — Several species either formed separate lineages or their phylogenetic positions were unstable. These are Ammodaucus leucotrichus, Laserpitium siler, L. pseudomeum and the genus Cuminum. The accessions of L. siler included in this study formed a highly supported monophyletic group with a firmly established sister position to the spiny-fruited clade (i.e., Daucus s.l., Silphiodaucus and Orlaya). Therefore, restitution of the genus Siler Mill. with a single species, Siler montanum Crantz, is justified. This montane species is highly diverse with several infraspecific taxa that are sometimes recognized as species including L. garganicum (Ten.) Bertol., L. ochridanum Micevski and L. zernyi Hayek (Stevanović & al., 1993). Detailed studies are necessary to elucidate their taxonomic status. Ammodaucus leucotrichus is an annual species distributed in northern Africa. Recently, A. nanocarpus (Beltrán) P.P6rez & Velasco has been recognized as a separate species. This taxon was formerly recognized as a subspecies of A. leucotrichus and differs from the latter in having smaller fruits; it has a Canarian-Moroccan distribution (Reyes-Betancort & al., 2007). Based on the analyses of ITS sequences only, Weitzel & al. (2014) included A. leucotrichus in Thapsia. However, our analyses suggest that the position of this species is unstable and that its inclusion in the analyses decreases the support for Thapsia. This may be due to the elevated rate of molecular evolution in this species as demonstrated by its long branch in the trees. Moreover, the phylogenetic analyses of cpDNA Version of Record 577 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae do not support a relationship of A. leucotrichus to Thapsia but place it in an isolated lineage. We therefore propose to retain Ammodaucus as a separate genus. Cuminum includes three species, two of which, C. cyminum and C. setifolium, were included in this study. The former has glabrous fruits and is widely cultivated but also sometimes occurs adventitiously, while the second has setose fruits and occurs in the wild in the Irano-Turanian region and central Asia (Rechinger, 1987a). Given their low sequence variation, these two species probably represent cultivated and wild varieties of the same species. Due to a high rate of molecular evolution, the phylogenetic position of this clade was unstable. However, it was always placed as a separate lineage among the laserpitioid clades, from which it differs in being annual and in having fruits without prominent secondary ribs. Therefore, its taxonomic status as a separate genus is undisputed. Laserpitium pseudomeum is endemic to the montane regions of Peloponnesus, Greece (Hartvig, 1986). The species does not seem to have any distinct characters that separate it from other congeners (A. Wojewódzka & K. Spalik, unpub. data). Its isolated phylogenetic position in all analyses suggests that it deserves to be placed in a separate genus. However, more data from morphology are necessary before this genus is established. NOMENCLATURAL CHANGES The following list contains species with their new generic status. A synopsis of tribe Daucinae with all recognised genera and species is provided in Appendix 2. Daucus L. sect. Daucus – Type: Daucus carota L. Daucus annuus (B6g.) Wojew., Reduron, Banasiak & Spalik, comb. nov. ≡ Tornabenea annua B6g. in Ann. Mus. Civico Storia Nat. Giacomo Doria, ser 3, 8: 39. 1918. Daucus dellacellae (Asch. & Barbey ex E.A.Durand & Barratte) Spalik, Banasiak & Reduron, comb. nov. ≡ Athamanta dellacellae Asch. & Barbey ex E.A.Durand & Barratte, Fl. Libyc. Prodr.: 108. 1910. Daucus elegans (Webb ex Bolle) Spalik, Banasiak & Reduron, comb. nov. ≡ Cryptotaenia elegans Webb ex Bolle in Braun, Append. Pl. Nov. Hort. Berol. 1861: 9: 1862. Daucus insularis (Parl. ex Webb) Spalik, Wojew., Banasiak & Reduron, comb. nov. ≡ Tetrapleura insularis Parl. ex Webb in Hooker, Niger Fl.: 131. 1849 ≡ Tornabenea insularis (Parl. ex Webb) Parl. in Hooker’s J. Bot. Kew Gard. Misc. 2: 370. 1850. Daucus mirabilis (Maire & Pamp.) Reduron, Banasiak & Spalik, comb. nov. ≡ Pachyctenium mirabile Maire & Pamp. in Arch. Bot. (Forlì) 12: 176. 1936. 578 Daucus rouyi Spalik & Reduron, nom. nov. ≡ Thapsia polygama Desf., Fl. Atlant. 1: 261, t. 75. 1798 ≡ Rouya polygama (Desf.) Coincy in Naturaliste, ser. 2, 15: 213. 1901, non Daucus polygamus Gouan, Ill. Observ. Bot.: 9. 1773. Daucus tenuissimus (A.Chev.) Spalik, Wojew., Banasiak & Reduron, comb. nov. ≡ Melanoselinum tenuissimum A.Chev. in Bull. Mus. Natl. Hist. Nat., s6r. 2. 7: 143. 1935 ≡ Tornabenea tenuissima (A.Chev.) A.Hansen & Sunding, Fl. Macaronesia, ed. 2, 1: 92. 1979. Daucus sect. Melanoselinum (Hoffm.) Spalik, Wojew., Banasiak & Reduron, comb. & stat. nov. ≡ Melanoselinum Hoffm., Gen. Pl. Umbell.: 156. 1814, pro gen. – Type: Selinum decipiens Schrad. & J.C.Wendl. (≡ Melanoselinum decipiens (Schrad. & J.C.Wendl.) Hoffm.). Daucus decipiens (Schrad. & J.C.Wendl.) Spalik, Wojew., Banasiak & Reduron, comb. nov. ≡ Selinum decipiens Schrad. & J.C. Wendl., Sert. Hannov. 3: 23, t. 13. 1797 ≡ Melanoselinum decipiens (Schrad. & J.C.Wendl.) Hoffm., Gen. Pl. Umbell.: 156. 1814. Daucus edulis (Lowe) Wojew., Reduron, Banasiak & Spalik, comb. nov. ≡ Monizia edulis Lowe in Hooker’s J. Bot. Kew Gard. Misc. 8: 295. 1856. Daucus sect. Anisactis DC. – Lectotype (designated here): Daucus brachiatus Sieber ex DC. = Daucus glochidiatus (Labill.) Fisch., C.A.Mey. & Av6-Lall. (≡ Scandix glochidiata Labill.) Daucus sect. Agrocharis (Hochst.) Spalik, Banasiak & Reduron, comb. & stat. nov. ≡ Agrocharis Hochst. in Flora 27: 19. 1844, pro gen. – Type: Agrocharis melanantha Hochst. Daucus incognitus (C.Norman) Spalik, Reduron & Banasiak, comb. nov. ≡ Caucalis incognita C.Norman in J. Bot. 72: 205. 1934 ≡ Agrocharis incognita (C.Norman) Heywood & Jury in Launert, Fl. Zambes. 4: 573. 1978. Daucus melananthos (Hochst.) Reduron, Spalik & Banasiak, comb. nov. ≡ Agrocharis melanantha Hochst. in Flora 27: 19. 1844. Daucus pedunculatus (Baker f.) Banasiak, Spalik & Reduron, comb. nov. ≡ Caucalis pedunculata Baker f. in Trans. Linn. Soc. London, Bot. 4: 15. 1894 ≡ Agrocharis pedunculata (Baker f.) Heywood & Jury in Launert, Fl. Zambes. 4: 573. 1978. Silphiodaucus (Koso-Pol.) Spalik, Wojew., Banasiak, Piwczyński & Reduron, stat. nov. ≡ Daucus sect. Silphiodaucus Koso-Pol. in Bull. Soc. Imp. Naturalistes Moscou 29: 211. 1916 – Lectotype (designated here): Daucus prutenicus (L.) E.H.L.Krause ≡ Laserpitium prutenicum L. Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Silphiodaucus prutenicus (L.) Spalik, Wojew., Banasiak, Piwczyński & Reduron, comb. nov. ≡ Laserpitium prutenicum L., Sp. Pl.: 248. 1753. Silphiodaucus hispidus (M.Bieb.) Spalik, Wojew., Banasiak, Piwczyński & Reduron, comb. nov. ≡ Laserpitium hispidum M.Bieb., Fl. Taur.-Caucas. 1: 221. 1808. Laser Borkh. ex G.Gaertn., B.Mey. & Scherb., Oekon. Fl. Wetterau 1: 244, 384. 1799 – Type: Laser trilobum (L.) Borkh. ex G.Gaertn. B.Mey. & Scherb. Laser affine (Ledeb.) Wojew. & Spalik, comb. nov. ≡ Laserpitium affine Ledeb., Fl. Ross. 2: 335. 1844. Laser archangelica (Wulfen) Spalik & Wojew., comb. nov. ≡ Laserpitium archangelica Wulfen in Jacquin, Collectanea 1: 214. 1787. Laser carduchorum (Hedge & Lamond) Wojew. & Spalik, comb. nov. ≡ Laserpitium carduchorum Hedge & Lamond in Notes Roy. Bot. Gard. Edinburgh 31(1): 76. 1971. Laser involucratum (Pall. ex Schult.) Spalik & Wojew., comb. nov. ≡ Cachrys involucrata Pall. ex Schult. in Roemer & Schultes, Syst. Veg. 6: 447–448. 1820 ≡ Polylophium involucratum (Pall. ex Schult.) Boiss., Fl. Orient. 2: 1066. 1872. Laser panjutinii (Manden. & Schischk.) Banasiak, Wojew. & Spalik, comb. nov. ≡ Polylophium panjutinii Manden. & Schischk. in Bot. Zhurn. (Moscow & Leningrad) 33: 318. 1948. Laser stevenii (Fisch., C.A.Mey. & Trautv.) Wojew. & Spalik, comb. nov. ≡ Laserpitium stevenii Fisch., C.A.Mey. & Trautv., Index Sem. Hort. Petrop. 4: 40. 1838. Thapsia L., Sp. Pl.: 261. 1753 – Type: Thapsia villosa L. Thapsia nestleri (Soy.-Will.) Wojew., Banasiak, Reduron & Spalik, comb. nov. ≡ Laserpitium nestleri Soy.-Will., Observ. Pl. France: 87. 1828 Thapsia eliasii (Sennen & Pau) Wojew., Banasiak, Reduron & Spalik, comb. nov. ≡ Laserpitium eliasii Sennen & Pau, Bol. Soc. Aragonesa Ci. Nat. 6: 25. 1907. Ekimia H.Duman & M.F.Watson in Edinburgh J. Bot. 56: 200. 1999 – Type: Prangos bornmuelleri Hub.-Mor. & Reese (≡ Ekimia bornmuelleri (Hub.-Mor. & Reese) H.Duman & M.F.Watson). Ekimia petrophila (Boiss. & Heldr.) Baczyński, Banasiak & Spalik, comb. nov. ≡ Laserpitium petrophilum Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., Ser. 1, 10: 46. 1849. 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Evol. 53: 56–68. http://dx.doi.org/10.1016/j.ympev.2009.05.029 Appendix 1. Accessions of Scandiceae subtribe Daucinae and outgroups used in this study with corresponding accession identifiers, voucher information and GenBank reference numbers. Newly generated sequences are identified with an asterisk behind the accession numbers. See Electr. Suppl.: Table S1 for detailed information on vouchers and references. Taxon name, accession ID, voucher, GenBank accession numbers: ITS, rpoB-trnC spacer, rps16 intron, rpoC1 intron Agrocharis gracilis Hook.f., G001, Cameroun, Letouzey 12123 (K), AY065344/AY065345, –, –, –. Agrocharis incognita (C. Norman) Heywood & Jury, 0046, Kenya, DNA supplied by E. Knox (coll. 2578) via S.R. Downie, KT347648*, –, KT347774*, KT347842*; 0245, Tanzania, Massawe, Gobbo & Mwiga 250 (E), KT347649*, –, KT347775*, –. Agrocharis melanantha Hochst, G002, Kenya, DNA supplied by E. Knox (coll. 2579), AF077794, –, –, –; 0248, Yemen Arab Republic, Miller 286 (E), KT347650*, –, –, –. Agrocharis pedunculata (Baker f.) Heywood & Jury, G004, Tanzania, Gereau & Kayombo 3870 (E), AY065342/ AY065343, –, –, –; G003, Malawi, Hillard Burtt 4131 (E), AF077792/AF077107, –, –, –; 0247, Tanzania, Gereau & Kayombo 3870 (E), KT347651*, –, KT347776*, –. Ammodaucus leucotrichus Coss. & Durieu, 0012, Morocco, cult. Cons. bot. Mulhouse no. F98003, –, KT347642*, KT347762*, KT347831*; G114, Spain, Canary Islands, Hansen 557 (C), KF160676, –, –, –. Anthriscus sylvestris (L.) Hoffm., 0083, France, Reduron s.n. (Reduron, pers. coll.), KT347715*, KT347743*, KT347806*, KT347879*. Athamanta cretensis L., 0617, Switzerland, Eckardt 1001 (B), KT347716*, –, –, –. Athamanta dellacellae Asch. & Barbey ex E.A. Durand & Barratte, 0435, Libya, Davis 50209 (E), AF073565/AF073566, –, –, KT347866*. Conopodium arvense (Coss.) Calest., 0630, Spain, Reverchon 1218 (B), KT347717, –, –, –. Cryptotaenia elegans Webb ex Bolle, 0039, Spain, Canary Islands, Danton s.n. (Reduron, pers. coll.), KT347674*, –, KT347791*, KT347860*; G099, Spain, Kunkel 17692 (G), DQ516354, –, –, –; G105, Spain, Canary Islands, Jarvis 617A (MO), DQ516355, –, –, –. Cuminum cyminum L., G007, cult., Lee 120 (ILL), U78362, –, U72436, –; 0339, cult. Turkey, (E 00328012), KT347713*, –, KT347804*, KT347878*; G008, HM176650, –, –, –; G009, HM176651, –, –, –; G010, HM176652, –, –, –; G011, HM176653, –, –, –; G012, HM176654, –, –, –; G013, HM176655, –, –, –. Cuminum setifolium (Boiss.) Koso-Pol., G006, Afghanistan, Hedge & al. 7083 (E), AF077796/AF077111, –, –, –; 0020, Iran, Rechinger 55742 (G), KT347714*, –, KT347805*, –. Daucus arcanus García-Martín & Silvestre, G014, Spain, García & Silvestre s.n. (E), AY065338/AY065339, –, –, –; G015, Spain (ABH 53887), JQ290118, –, –, –; 0260, Mexico, Davis & Lightowlers 66231 (E), KT347671*, –, –, KT347858*. Daucus aureus Desf., G017, Reading Univ./B.M.Exped. 1076 (RNG), HE602378, –, –, –; G016, cult., Lee 57 (ILL), AF077784/AF077099, –, –, –; G018, Spain, Triano & Castro s.n. (ABH 55117), JQ290119, –, –, JQ290131; G019, Algeria, Juan s.n. (ABH 57786), JQ290120, –, –, –; 0273, Tunisia, Davis & Lamond 57901 (E), KT347685*, –, –, –; 0439, Algeria, Reverchon 203 (B), KT347686*, –, –, –. Daucus bicolor Sibth. & Sm., G042, Israel, Lee 270 (ILL), AF077791/AF077106, –, –, –; 0041, Israel, Cohen s.n. (WA), KT347652*, –, KT347777*, KT347843*. Daucus biseriatus Murb., G089, Algeria, cult. Univ. of California no. C-958, AY065328/AY065329, –, –, –. Daucus broteri Ten., 0276, Turkey, Coode & Jones 2575 582 Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Appendix 1. Continued. (E), KT347653*, –, KT347778*, KT347844*; 0277, Turkey, Alava & Bocquet 4988 (E), KT347654*, –, KT347779*, KT347845*; 0440, Turkey, Bornmüller & Bornmüller 14160 (B), KT347655*, –, –, KT347846*. Daucus carota subsp. azoricus Franco, G024, Morocco (RNG), AY065312/AY065313, –, –, –; 0610, Portugal, Azores, Danton s.n. (Cons. bot. Mulhouse no. 08-092A, Reduron, pers. coll.), KT347693*, –, –, KT347869*. Daucus carota subsp. capillifolius (Gilli) Arbizu, G021, Libya (E), AY065318/AY065319, –, –, –. Daucus carota L. subsp. carota, G098, Germany, Downie 164 (ILL), U27589/U30315, –, U36290, –; G022, Jury 17848 (RNG), HE602376, –, –, –; G025, Kazakhstan, Lee 167 (ILL), AF077779, –, –, –; G023, AY552527, –, –, –; 0032, France, Reduron s.n. (Cons. bot. Mulhouse no. 08-114A, Reduron, pers. coll.), KT347694*, –, KT347798*, KT347870*; 0443, Greece, Böhling 10118 (B), KT347695*, –, –, KT347871*; 0444, Romania, Paucǎ 2255 (B), KT347696*, –, –, KT347872*; 0442, Iran, Wojewódzka & Zych s.n. (WABG), FJ415158, –, –, –. Daucus carota subsp. commutatus (Paol.) Thell., 0003, France, Bocquet 16283 (G), KT347697*, –, KT347799*, KT347873*. Daucus carota subsp. drepanensis (Arcang.) Heywood, G026, Portugal, AY065314/AY065315, –, –, –. Daucus carota subsp. gadecaei (Rouy & Camus) Heywood, G027, France, cult. Univ. of Reading, AY065316/ AY065317, –, –, –; 0034, France, Reduron s.n. (Reduron, pers. coll.), KT347698*, –, KT347800*, –. Daucus carota subsp. gummifer Hook.f., G028, cult. UIUC from seeds from France, Lee 47 (ILL), AF077782/AF077097, –, –, –; 0445, France, Kohlmeyer 1448 (B), KT347699*, –, –, KT347874*. Daucus carota subsp. halophilus (Brot.) Okeke, G029, cult. UIUC from seeds from France, Lee 81 (ILL), AF077781/AF077096, –, –, –; 0009, Portugal, cult. Cons. bot. Mulhouse no. 9305*, KT347700*, –, KT347801*, KT347875*. Daucus carota subsp. hispanicus (Gouan) Thell., G030, Spain, Llorenç Saéz s.n. (ABH 53270), JQ290121, –, –, JQ290132; 0035, France, Reduron, Dumet & Le Clerc s.n. (Reduron, pers. coll.), KT347701*, –, –, –; 0447, Spain, Benedi & Molero 28 (B), KT347703*, –, –, –. Daucus carota subsp. hispanicus var. linearis Reduron, 0038, France, Reduron s.n. (Reduron, pers. coll.), KT347702*, –, –, –. Daucus carota subsp. maritimus (Lam.) Batt., 0448, France, Mory s.n. (B), KT347704*, –, –, KT347876*. Daucus carota subsp. maximus (Desf.) Ball., G031, cult., Lee 64 (ILL), AF077778/AF077093, –, –, –; 0269, Algeria, Davis 52992 (E), KT347705*, –, –, –; 0274, Turkey, Davis 47038 (E), KT347706*, –, –, –; 0449, Turkey, Kehl s.n. (B), KT347707*, –, –, –; 0450, Algeria, Reverchon 45 (B), KT347708*, –, –, –; 0451, Greece, Shay 80-394 (B), KT347709*, –, –, –. Daucus conchitae Greuter, G032, Turkey, Jury & Warren 366 (RNG), AY065332/AY065333, –, –, –; 0236, Turkey, Gardner, Knees, Barker-Mill & Layman 8120 (E), KT347656*, –, KT347780*, KT347847*. Daucus crinitus Desf., G033, Ait Lafkih & al. 70 (RNG), HE602443, –, –, –; G034, Spain, Martínez-Flores s.n. (ABH 52065), JQ290122, –, –, –; 0014, cult. UIUC from seeds from Portugal, Lee 49 (ILL), KT347675*, –, KT347792*, KT347861*. Daucus durieua Lange, G037, Díez 3482/94 (RNG), HE602377, –, –, –; G036, Israel, cult. UIUC, Lee 271 (ILL), AF077790/AF077105, –, –, –; G038, Spain, Navarro Reyes s.n. (ABH 53919), JQ290123, –, –, JQ290133; 0040, Israel, Cohen s.n. (WA), KT347657*, –, KT347781*, KT347848*; 0453, Spain, Valdés & al. 256/88 (B), KT347658*, –, –, KT347849*; 0454, Israel, Danin s.n. (B), KT347659*, –, –, –. Daucus glochidiatus (Labill.) Fisch., C.A.Mey. & Av6-Lall., G039, New Zealand (AK 297601), EU331132, –, –, –; G040, Australia, New South Wales, Lepschi 449 (CANB), AY065340/AY065341, –, –, –; 0456, Australia, New South Wales, Eichler 22856 (B), KT347660*, –, –, KT347851*; 0455, Australia, North West Plains, Blaylock 938 (KRAM), FJ415160*, KT347740*, KT347782*, KT347850*. Daucus gracilis Steinh., G041, Algeria, Davis 52098 (RNG), AY065322/AY065323, –, –, –; 0270, Algeria, Davis 52098 (E), KT347687*, –, –, –. Daucus guttatus Sm., G043, Greece, Jury & Warren 209 (RNG), AY065336/AY065337, –, –, –; 0238, Greece, Gardner, Knees & Amor 8226 (E), KT347661*, –, KT347783*, –; 0458, Greece, Risse 2101 (B), KT347662*, –, KT347784*, KT347852*. Daucus involucratus Sm., G044, Greece, Bowen 8896 (E), AY065334/AY065335, –, –, –; 0037, Greece, Charpin 25294 (G), KT347663*, –, KT347785*, KT347853*; 0272, Turkey, Davis 41352 (E), KT347664*, –, KT347786*, –; 0460, Greece, Greuter & Matthäs 19630 (B), KT347665*, –, KT347787*, KT347854*. Daucus littoralis Sm., 0008, Israel, cult. Cons. bot. Mulhouse no. 99080*, Reduron s.n. (Reduron, pers. coll.), KT347666*, –, KT347788*, KT347855*; 0275, Egypt, Mashaly s.n. (E), KT347667*, –, –, –; 0461, Iran, Wojewódzka & Zych s.n. (WABG), FJ415159*, KT347741*, –, KT347856*. Daucus mauritii Sennen, G045, Morocco, Crespo & al. s.n. (ABH 55659), JQ290124, –, –, JQ290134; G046, Morocco, Crespo & al. s.n. (ABH 55656), JQ290125, –, –, –. Daucus montanus Humb. & Bonpl., G047, Argentina, cult. Botanical Garden of the University of California, Berkeley 94.0563, AF077789/AF077104, –, –, –; 0017, Chile, cult. Cons. bot. Mulhouse no. 98050, Reduron s.n. (Reduron, pers. coll.), KT347668*, –, KT347789*, KT347857*; 0230, Mexico, Gardner & Knees 5108 (E), KT347669*, –, –, –; 0262, Peru, Hutchinson 1659 (E), KT347670*, –, –, –. Daucus muricatus L., G049, Jury 16748 (RNG), HE602379, –, –, –; G048, cult. UIUC, Lee 36 (ILL), AF077785/AF077100, –, –, –; G050, Morocco, Crespo & al. s.n. (ABH 55634), JQ290126, –, –, JQ290135; G051, Spain, Meneses Sores s.n. (ABH 53894), JQ290127, –, –, JQ290136; 0271, Algeria, Davis 58080 (E), KT347676*, –, KT347793*, KT347862*; 0281, Portugal, Gibbs 69.148 (E), KT347677*, –, –, KT347863*. Daucus pusillus Michx., G052, cult. Botanical Garden of the University of California, Berkeley 92.0891, AF077788/AF077103, –, –, AF123729; G053, Argentina, Camadro s.n. (ABH 57683), JQ290128, –, –, JQ290137; 0267, USA, California, Thorne, Wallace & Haefs 48769 (E), KT347672*, –, –, –; 0466, Argentina, Leuenberger & Arroyo 3867 (B), KT347673*, –, KT347790*, KT347859*. Daucus sahariensis Murb., G054, Algeria, JGR & AA 129-108 (RNG), AY065320/AY065321, –, –, –. Daucus setifolius Desf., G055, Jury 17514 (RNG), HE602375, –, –, –; G056, Spain, Aparicio s.n. (ABH 53906), JQ290129, –, –, JQ290138; 0237, Spain, Gardner, Knees & Read 4837 (E), KT347678*, –, KT347794*, –; 0467, Algeria, Reverchon 234 (B), KT347679*, –, –, KT347864*. Daucus syrticus Murb., G057, Libya (RNG), AY065324/AY065325, –, –, –; 0265, Libya, Davis 49612 (E), KT347688*, –, –, –; 0468, Libya, Bornmüller 711 (B), KT347689*, –, KT347797*, KT347867*. Daucus tenuisectus Coss. ex Batt., G058, Morocco, Jury & Springate s.n. (RNG), AY065326/AY065327, –, –, –; 0266, Morocco, Balls 2504 (E), KT347681*, –, KT347795*, –; 0036, Morocco, Charpin 27029 (G; Reduron, pers. coll.), KT347680*, –, –, –. Daucus virgatus (Poir.) Maire, 0050, Algeria, Maire (Duffour 4456) (G), KT347710*, –, –, –; 0659, Algeria, Véla s.n. (Reduron, pers. coll.), KT347711*, –, KT347802*, –. Ekimia bornmuelleri (Hub.-Mor. & Reese) H. Duman & M.F. Watson, 0655, Turkey, Duman & Karaveliogullari 5071 (E), KT347640*, –, –, KT347810*. Ferula communis L., 0195, Spain, Sánchez-Gómez s.n. (Univ. of Zaragoza, Spain), DQ379392, KJ660616, KJ660477, KJ698369. Ferula olivacea (Diels) H. Wolff, 0363, China, Chamberlain, Ming, Yuan & al. 229 (E), KJ660802, KJ660688, KJ660547, KJ660382. Glaucosciadium cordifolium (Boiss.) B.L. Burtt & P.H. Davis, 0221, Cyprus, cult. Cons. bot. Mulhouse no. 98112, Reduron s.n. (WA), DQ379459, KJ660745, KJ660458, KJ660439. Glaucosciadium insigne (Pimenov & Maassoumi) Spalik & S.R. Downie, 0074, Iran, Mozaffarian 77099 (TARI), KJ660839, KJ660746, KJ660459, KJ660440. Laser trilobum (L.) Borkh. ex G. Gaertn., B. Mey. & Scherb., G061, Azerbaijan, cult. Moscow State University Botanical Garden, Russia, Pimenov & al. s.n. (MW), AF008644/AF009123, –, –, AF123735; 0611, France, cult. Cons. bot. Mulhouse no. 98020B, Reduron s.n. (Reduron, pers. coll.), KT347638*, –, KT347744*, KT347807*; G112, Jelitto Staudensamen GmbH, Schwarmstedt, Germany, 2011, Lot-No. 70016019AA1AA0s. LA017, KF160678, –, –, –. Laserpitium affine Ledeb., G062, Georgia, Pimenov 1454 (MW), JQ305145, –, –, –; 0480, Georgia, Shreter & Pimenov 394 (LE), FJ415151*, KT347718*, KT347745*, KT347808*. Laserpitium archangelica Wulfen, 0481, Slovenia, Mayer 63416 (KRAM), FJ415153, KJ832093, KJ832098, KJ832103. Laserpitium carduchorum Hedge & Lamond, 0482, Turkey, Davis & Polunin 24567 (E 00042009), FJ415116*, –, –, –; 0483, Turkey, Davis & Polunin 22551 (E), FJ415117, KJ832094, KJ832099, KJ832104. Laserpitium eliasii Sennen & Pau subsp. eliasii, 0484, Spain, Alejandre s.n. (VIT 27070), FJ415118*, –, –, KT347823*. Laserpitium eliasii subsp. ordunae P. Monts., 0485, Spain, Montserrat s.n. (JACA 733483), FJ415119, KJ832095, KJ832100, KJ832105. Laserpitium eliasii subsp. thalictrifolium (Samp.) P. Monts., 0486, Spain, Rico s.n. (JACA ex SALA 41460), FJ415120*, KT347727*, KT347755*, KT347824*. Laserpitium gallicum L. subsp. gallicum, 0487, France, Reduron s.n. (Reduron, pers. coll.), FJ415128*, KT347722*, KT347749*, KT347816*. Laserpitium gallicum subsp. angustissimum (Willd.) Lange, 0488, Morocco, Jury & al. 17635 (E 00065531), FJ415129*, KT347723*, KT347750*, KT347817*. Laserpitium gallicum subsp. orospedanum M.B. Crespo & al., 0489, Spain, Crespo & al. s.n. (isotype, ABH 43500), FJ415126*, KT347724*, KT347751*, KT347818*. Laserpitium gallicum subsp. paradoxum (O. Bolòs & Font Quer) P. Monts., 0490, Spain, Viñas s.n. (HGI 15292), FJ415127*, KT347725*, KT347752*, KT347819*. Laserpitium glaucum Post, 0491, Turkey, Darrah 552 (E 00042002), FJ415115*, –, –, –. Laserpitium halleri Crantz, 0493, France, Reverchon s.n. (KRAM), FJ415130*, –, –, –. Laserpitium hispidum M. Bieb., G064, Russia, Krasnodar, Pimenov 316 (MW), JQ305142, –, –, –; G065, Turkey, Pimenov & Kljuykov s.n. (MW), JQ305143, –, –, –; G066, Russia, Krasnodar, Ostroumova 19 (MW), AF077898, –, –, –; 0494, Turkey, Davis, Coode & Yaltirik 38795 (E 00042016), FJ415154*, –, KT347772*, –; 0495, Ukraine, Crimea, Werblan-Jakubiec & al. s.n. (WABG), FJ415155*, –, –, –. Laserpitium krapfii Crantz subsp. krapfii, 0496, Romania, Nyárády 976 (WA), FJ415124*, –, –, KT347821*. Laserpitium krapfii subsp. gaudinii (Moretti) Thell., 0497, Italy, Charpin s.n. (G), FJ415125*, KT347726*, KT347753*, KT347820*. Laserpitium latifolium L., 0498, Poland, Sudnik s.n. (WA), FJ415131*, –, KT347754*, KT347822*. Laserpitium nestleri Soy.-Will. subsp. nestleri, 0501, Italy, Fabregat & Udias 2007 (BCC), FJ415123, KJ832097, KJ832102, KJ832107. Laserpitium nestleri subsp. flabellatum var. tensinum P. Monts., 0505, Version of Record 583 TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Appendix 1. Continued. Spain, Montserrat s.n. (JACA R265475), FJ415121*, KT347728*, KT347756*, KT347825*. Laserpitium nestleri subsp. lainzii P. Monts., 0506, Spain, Garcia s.n. (JACA 466085), FJ415122*, KT347729*, KT347757*, KT347826*. Laserpitium nitidum Zanted., 0507, Italy, Cobau 2715 (KRA), FJ415132*, –, –, –. Laserpitium petrophilum Boiss. & Heldr., 0508, Turkey, Southam s.n. (RNG), FJ415114*, –, –, –; 0509, Turkey, Davis 18412 (E), KT347641*, –, KT347747*, KT347811*; G067, Turkey, Spalik & Żochowska s.n. (WA), AF073567, –, –, –; G085, Turkey, Eren s.n. (B), JQ305150, –, –, –; G086, Turkey, Davis 18412 (E), JQ305151, –, –, –; G087, Turkey, Hartvig 23613 (EGE), JQ305152, –, –, –; G088, Turkey, Kotschy 186/242 (MW), JQ305153/JQ305154, –, –, –. Laserpitium peucedanoides L., 0510, Montenegro, Zarzycki s.n. (KRAM), FJ415133*, –, –, –. Laserpitium prutenicum L. subsp. prutenicum, 0649, Poland, Nowak s.n. (WA), AF336374, –, KT347773*, KT347841*; G069, Russia, Maiorov s.n. (MW), JQ305144, –, –, –. Laserpitium prutenicum subsp. duforianum (Rouy & Camus) Braun-Blanq., 0511, Spain, Montserrat s.n. (JACA 701083), FJ415156*, –, –, –. Laserpitium pseudomeum Orph., Heldr. & Sart. ex. Boiss., 0513, Greece, Gustavsson 9672 (G), FJ415134*, –, KT347748*, KT347812*. Laserpitium siler L., 0515, Italy, Davis & Sutton D 65739 (E), FJ415111*, –, –, KT347813*; 0516, Spain, Gardner & Gardner 781 (E 00043183), FJ415112*, KT347720*, –, KT347814*; 0517, Montenegro, Gardner & Gardner 2455 (E 00043177), FJ415113*, KT347721*, –, KT347815*; G070, Germany, cult. Johannes Gutenberg University (no. 1112), Downie 71 (ILL), U30528/U30529, –, U36296, AF123734; G113, Jelitto Staudensamen GmbH, Schwarmstedt, Germany, 2011, Lot-No. 80011018100g. LA021, Simonsen 2013-09 (C), KF160679, –, –, –. Laserpitium stevenii Fisch. & Trautv., 0518, Georgia, Muibaniani & al. s.n. (LE), FJ415152*, KT347719*, KT347746*, KT347809*; G071, Georgia, Pimenov 886 (MW), JQ305146, –, –, –. Leutea avicennae Mozaff., 0067, Iran, Mozaffarian 65041 (TARI), KJ660830, KJ660736, KJ660449, KJ660430. Melanoselinum decipiens (Schrad. & J.C. Wendl.) Hoffm., G109, cult. Botanical Garden, Copenhagen, Denmark, Hansen 13407 (C), KF160680, –, –, –; 0522, Portugal, Madeira, cult. Cons. bot. Mulhouse no. 2027, Reduron s.n. (Reduron, pers. coll.), FJ415161, –, –, –; 0612, Seeds from Botanischer Garten der Universit2t Kiel, Univ. of Warsaw Bot. Garden nos. KS 124/97, 163/245 (WA), KT347682*, –, –, –; 0613, Portugal, Madeira, cult. University of Oslo Botanical Garden, seeds no. 447, garden no. 257, KT347683*, –, –, –; G072, Portugal, Madeira, cult. University of Reading, England, APE 605, plant A EF016755, –, –, AF123737; G073, Portugal, Madeira, cult. University of Reading, England, APE 605, plant B, EF016756, –, –, AF123738. Monizia edulis Lowe, 0013, Portugal, Madeira, cult. Cons. bot. Mulhouse no. 98141, KT347684*, –, KT347796*, KT347865*; G074, Madeira, cult. Madeira Botanic Garden, F. & O. Baets 08655 (E), AF073569, –, –, AF123739; G110, Portugal, Madeira, Hansen 2478 (C), KF160681, –, –, –. Orlaya daucoides (L.) Greuter, 0229, Greece, Gardner, Knees & Amor 8225 (E), KT347643*, –, –, –; 0263, Italy, Davis & Sutton 65766 (E), KT347644*, –, –, –; 0264, Spain, Galiano, Gibbs, Silvestre & Valdes 1284.69 (E), KT347645*, –, –, –; G075, cult. UIUC form seeds from Hungarian Academy of Sciences, Vácrátót, Lee 7 (ILL), AF077797, –, –, AF123733; G079, Germany, cult., Downie 20 (ILL), U30526/U30527, –, –, –; G111, Greece, Strid 42072 (C), KF160682, –, –, –. Orlaya daucorlaya Murb., G076, Macedonia, Edmonston 27 (E), AF077798/AF077113, –, –, –. Orlaya grandiflora (L.) Hoffm., 0258, Slovakia, Smejkal & Vicherek 1544 (E), KT347646*, –, KT347771*, KT347840*; 0259, Italy, Davis & Sutton 65886 (E), KT347647*, –, –, –; G077, France, cult. Jardin botanique de Caen (no. 1474), Downie 309 (ILL), U30524/U30525, –, –, –; G078, Spain, Lopes & Javier s.n. (ABH 55673), JQ290130, –, –, –. Pachyctenium mirabile Maire & Pamp., 0244, Libya, Davis 50249 (E), KT347690*, –, –, –; G080, Libya, Davis 50249 (E), AF077787/AF077102, –, –, –. Polylophium involucratum (Pall. ex Schult.) Boiss., 0395, Iran, Klein 3699 (W 06547), KT347639*, –, –, –; G081, Iran, Mozaffarian s.n. (TARI), JQ305147, –, –, –. Polylophium panjutinii Manden. & Schischk., G082, Georgia, Daushkevich. s.n. (MW), AF008645/AF009124, –, –, AF123736; G083, Georgia, Arzinba s.n. (AA), JQ305148, –, –, –; G084, Georgia, Ostroumova s.n. (MW), JQ305149, –, –, –. Pseudorlaya minuscula (Pau ex Font Quer) Laínz, G090, Spain (RNG), AY065330/AY065331, –, –, –. Pseudorlaya pumila (L.) Grande, G091, cult. University of Oldenburg Botanic Garden (no. 20), Downie 138 (ILL), U30522, –, –, –; 0227, Algeria, Davis 51712 (E), KT347691*, –, –, –; 0228, Tunisia, Davis 56740b (E), KT347692*, –, –, –; G108, Greece, Strid 38276 (C), KF160683, –, –, –. Rouya polygama (Desf.) Coincy, 0523, France, cult. Cons. bot. Mulhouse no. 99143, 9 September 2000, FJ415157, –, –, KT347868*; G107, Italy, Greuter 9736 (C), KF160684, –, –, –. Thapsia asclepium L., G121, Greece, Strid 39066 (C), KF160685, –, –, –; 0470, Italy, Optima Iter VIII (RNG), FJ415135, –, –, –. Thapsia garganica L., G092, Italy, AJ007930, –, –, –; G128, Greece, Strid 26821 (C), KF160688, –, –, –; 0525, Algeria, Davis 53034 (E), FJ415143*, KT347730*, KT347758*, KT347827*; 0526, Tunisia, Davis & Lamond D 56827 (E), FJ415144*, KT347731*, KT347759*, KT347828*; 0527, Greece, Edmondson & McClintock E 2579 (E), FJ415145*, KT347732*, KT347760*, KT347829*. Thapsia gummifera (Desf.) Spreng., 0520, Portugal, cult. Cons. bot. Mulhouse no. 9309, FJ415139, KT347733*, KT347761*, KT347830*. Thapsia gymnesica Rosselló & A. Pujadas, G115, Spain, Smitt 94-01 (C), KF160693, –, –, –. Thapsia laciniata Rouy, G132, France, Smitt 90-01 (C), KF160694, –, –, –; G133, Spain, Smitt 87-12 (C), KF160696, –, –, –. Thapsia maxima Mill., G122, Spain, Smitt 87-31 (C), KF160698, –, –, –; G123, Portugal, Smitt 88-19 (C), KF160699, –, –, –. Thapsia meoides Guss., 0472, Morocco, Jury s.n. (RNG), FJ415137*, –, –, –; 0473, Morocco, Davis 54321 (E), FJ415138*, –, KT347763*, KT347832*; G059, Jury & Upson 20572 (RNG), HE602456, –, –, –. Thapsia minor Hoffmanns. & Link, G128, Portugal, Smitt 81-v-10 (C), KF160701, –, –, –; G129, Portugal, Smitt 88-17 (C), KF160702, –, –, –; G130, Portugal, Smitt 88-25 (C), KF160703, –, –, –; G131, Portugal, Smitt 88-30 (C), KF160704, –, –, –. Thapsia platycarpa Pomel, G093, Jury 15837 (RNG), HE602373, –, –, –. Thapsia scabra (Cav.) Simonsen, Rønsted, Weitzel & Spalik, 0479, Spain, Aran & Tohá s.n. (VAL 118242), FJ415150, –, KT347764*, KT347833*. Thapsia smittii Simonsen, Rønsted, Weitzel & Spalik, G119, Morocco, Montserrat FC-9169 (C), KF160706, –, –, –; G120, Portugal, Smitt 81-v-11 (C), KF160708, –, –, –. Thapsia tenuifolia Lag., 0469, Portugal, cult. Cons. bot. Mulhouse no. 9372, FJ415140*, KT347735*, KT347766*, KT347835*. Thapsia thapsioides (Desf.) Simonsen, Rønsted, Weitzel & Spalik, 0477, Tunisia, Davis & Lamond D 57768 (E 00040997), FJ415141*, KT347736*, KT347767*, KT347836*; 0478, Algeria, Davis 53419 (E), FJ415142, KT347737*, KT347768*, KT347837*. Thapsia transtagana Brot., 0524, Morocco, Davis & Davis D 48431 (E), FJ415146*, KT347738*, KT347769*, KT347838*; G094, Jury 16325 (RNG), HE602372, –, –, –; G116, Portugal, Smitt 81-t-16 (C), KF160714, –, –, –; G117, Spain, Smitt 87-15 (C), KF160719, –, –, –. Thapsia villosa L., 0530, Portugal, Bowen s.n. (RNG), FJ415149*, –, –, –; G095, Lambinon 94/Ma/350 (RNG), HE602371, –, –, –; G124, Portugal, Smitt 88-33 (C), KF160723, –, –, –; G125, Portugal, Smitt 88-32 (C), KF160726, –, –, –; G126, Spain, Smitt 87-11 (C), KF160730, –, –, –; G127, Portugal, Weitzel 2011-10 (C), KF160734, –, –, –. Thapsia villosa var. dissecta Boiss., 0528, Morocco, Davis & King D 68336 (E), FJ415147*, KT347739*, KT347770*, KT347839*; 0529, Spain, Davis 61691 (E 00042013), FJ415148*, –, –, –. Thapsia sp., 0471, France, Corsica, cult. Cons. bot. Mulhouse no. 98138, FJ415136*, KT347734*, KT347765*, KT347834*. Tornabenea annua B6g., 0521, Cape Verde Islands, cult. Cons. bot. Mulhouse no. 99145, KT347712*, KT347742*, KT347803*, KT347877*; G096, Cape Verde Islands, Hildenbrand, Meyer & Reduron 01007 (ILL, Reduron, pers. coll.), DQ516356, –, –, –. Tornabenea insularis (Parl. ex Webb.) Parl., G106, cult. Botanical Garden, Copenhagen, Denmark, Hansen 53 (C), KF160739, –, –, –. Tornabenea tenuissima (A. Chev.) A. Hansen & Sunding, G097, Cape Verde Islands, Hildenbrand, Meyer & Reduron 2036 (ILL, Reduron, pers. coll.), DQ516357, –, –, –. 584 Version of Record TAXON 65 (3) • June 2016: 563–585 Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae Appendix 2. Synopsis of tribe Scandiceae subtribe Daucinae. The arrangement of genera and species reflects their phylogenetic position in the combineddata tree (see Figs. 3 and 4). For the species that have changed their generic position, their previous (commonly used) name is also given. Species, which were not checked for molecular markers, are indicated with asterisks. Daucus L. sect. Daucus: D. carota L., D. annuus (B6g.) Wojew. & al. [≡ Tornabenea annua B6g.], D. insularis (Parl. ex Webb) Spalik & al. [≡ Tornabenea insularis (Parl. ex Webb) Parl.], D. tenuissimus (A.Chev.) Spalik & al. [≡ Tornabenea tenuissima (A.Chev.) A.Hansen & Sunding], D. virgatus (Poir.) Maire, D. syrticus Murb., D. gracilis Steinh., D. sahariensis Murb., D. rouyi Spalik & Reduron [≡ Rouya polygama (Desf.) Coincy], D. biseriatus Murb., D. pumilus (L.) Hofmanns. & Link [≡ Pseudorlaya pumila (L.) Grande], D. minusculus Pau ex Font Quer [≡ Pseudorlaya minuscula (Pau ex Font Quer) Laínz], D. mirabilis (Maire & Pamp.) Reduron & al. [≡ Pachyctenium mirabile Maire & Pamp.], D. dellacellae (E.A.Durand & Barratte) Spalik & al. [≡ Athamanta dellacellae E.A.Durand & Barratte], D. aureus Desf., D. mauritii Sennen, D. setifolius Desf., D. crinitus Desf., D. muricatus L., D. tenuisectus Coss. ex Batt., D. elegans (Webb ex Bolle) Spalik & al. [≡ Cryptotaenia elegans Webb ex Bolle]; sect. Melanoselinum (Hofm.) Spalik & al.: D. decipiens (Schrad. & J.C.Wendl.) Spalik & al. [≡ Melanoselinum decipiens (Schrad. & J.C.Wendl.) Hofm.], D. edulis (Lowe) Wojew. & al. (≡ Monizia edulis Lowe); sect. Anisactis DC.: D. bicolor Sibth. & Sm., D. littoralis Sm., D. guttatus Sm., D. arcanus García-Martín & Silvestre, D. pusillus Michx., D. involucratus Sm., D. conchitae Greuter, D. broteri Ten., D. montanus Humb. & Bonpl., D. glochidiatus (Labill.) Fisch., C.A.Mey. & Av6-Lall., D. durieua Lange; sect. Agrocharis (Hochst.) Spalik & al. : D. incognitus (C.Norman) Spalik & al. [≡ Agrocharis incognita (C.Norman) Heywood & Jury], D. melananthos (Hochst.) Reduron & al. [≡ Agrocharis melanantha Hochst.], D. pedunculatus (Baker f.) Banasiak & al. [≡ Agrocharis pedunculata (Baker f.) Heywood & Jury]; Incertae sed1s: *Tornabenea bischoffii J.A.Schmidt, *T. humilis Lobin & K.H.Schmidt, *T. ribeirensis K.H.Schmidt & Lobin, Agrocharis gracilis Hook.f., *Daucus jordanicus Post, *D. microscias Bornm. & Gauba, *D. reboudii Coss. Silphiodaucus (Koso-Pol.) Spalik & al. S. prutenicus (L.) Spalik & al. [≡ Laserpitium prutenicum L.], S. hispidus (M.Bieb.) Spalik & al. [≡ Laserpitium hispidum M.Bieb.]. Orlaya Hofm. O. daucoides (L.) Greuter, O. daucorlaya Murb., O. grandiflora (L.) Hofm., Laser Borkh. ex G.Gaertn. & al. L. panjutinii (Manden. & Schischk.) Banasiak & al. [≡ Polylophium panjutinii Manden. & Schischk.], L. involucratum (Pall. ex Schult.) Spalik & Wojew. [≡ Polylophium involucratum (Pall. ex Schult.) Boiss.], L. stevenii (Fisch., C.A.Mey. & Trautv.) Wojew. & Spalik [≡ Laserpitium stevenii Fisch., C.A.Mey. & Trautv.], L. archangelica (Wulfen) Spalik & Wojew. [≡ Laserpitium archangelica Wulfen], L. trilobum (L.) Borkh. ex G.Gaertn. & al., L. carduchorum (Hedge & Lamond) Wojew. & Spalik [≡ Laserpitium carduchorum Hedge & Lamond], L. affine (Ledeb.) Wojew. & Spalik [≡ Laserpitium affine Ledeb.]. Siler Mill. Siler montanum Crantz [≡ Laserpitium siler L., inc. L. garganicum (Ten.) Bertol., L. zernyi Hayek, L. ochridanum Micevski]. Thapsia L. T. garganica L., T. gymnesica Rosselló & A.Pujadas, T. platycarpa Pomel, T. transtagana Brot., T. smittii Simonsen & al., T. meoides Guss., T. thapsioides (Desf.) Simonsen & al., T. gummifera (Desf.) Spreng., T. tenuifolia Lag., T. asclepium L., T. minor Hofmanns. & Link, T. villosa L., T. laciniata Rouy, T. maxima Mill. [= T. nitida Lacaita?], T. scabra (Cav.) Simonsen & al., T. nestleri (Soy.-Will.) Wojew. & al. [≡ Laserpitium nestleri Soy.-Will.], T. eliasii (Sennen & Pau) Wojew. & al. [≡ Laserpitium eliasii Sennen & Pau], *T. cinerea A.Pujadas., *T. foetida L. Laserpitium L. L. nitidum Zanted., L. peucedanoides L., L. krapfii Crantz, L. latifolium L., L. gallicum L., L. halleri Crantz. Ekimia H.Duman & M.F.Watson E. bornmuelleri (Hub.-Mor. & Reese) H.Duman & M.F.Watson, E. petrophila (Boiss. & Heldr.) J.Baczyński & al. [≡ Laserpitium petrophilum Boiss. & Heldr.], E. glauca (Post) Banasiak & al. [≡ Laserpitium glaucum Post]. Ammodaucus Coss. & Durieu A. leucotrichus Coss. & Durieu, *A. nanocarpus (Beltrán) P.P6rez & Velasco. Cuminum L. C. cyminum L., C. setifolium (Boiss.) Koso-Pol., Incertae sed1s: *Cuminum borsczowii (Regel & Schmalh.) Koso-Pol., *Elaeoselinum tunetanum Brullo, Minissale & Terrasi, Laserpitium pseudomeum Orph., Heldr. & Sart. ex. Boiss., *L. longiradium Boiss. Version of Record 585